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A TEXT-BOOK
OF
MYCOLOGY AND PLANT

PATHOLOGY
HARSHBERGER
A TEXT-BOOK
OF
MYCOLOGY AND PLANT

PATHOLOGY
BY
JOHN W. HARSHBERGER, Ph.D.

PROFESSOR OF BOTANY, UNIVERSITY' OF PENNSYLVANIA; MEMBER OF
THE BOTANICAL SOCIETY OF AMERICA; VICE-PRESIDENT OF THE
ECOLOGICAL SOCIETY OF AMERICA, ETC.
WITH 271 ILLUSTRATIONS
PHILADELPHIA
P. BLAKISTON'S SON & CO.
1012 WALNUT STREET
Copyright, 191 7, by P. Blakiston's Son & Co.
THE MAPLE PRESS YORK PA

PREFACE
This book is the outcome of twenty-seven years' experience, as a
teacher of botany, during which fifteen years have been given to a
graduate course on the morphology, classification and physiology of the
fungi, and five years to a course which combined with this considera-
tion a parallel study of the most important cultural and inoculation
methods used by the practical bacteriologist and mycologist at the
present day. The English and Germans have led in the production
of text-books on mycology and pathology; Berkeley, Smith, Cooke
and Massee in England, Frank, Sorauer, von Tubeuf and Kiister in
Germany. Americans have been behind in this important field,
notwithstanding, that American plants harbor some of the most
destructive fungi, which, through our careless methods of agriculture
and horticulture up to the present, are annually destructive to the
extent of millions of dollars. This lack is being rapidly remedied and
the appearance of text-books by Duggar, Stevens, Hall and' Stevens,
Mel T. Cook and general monographs by Erwin T. Smith, and others,
augurs well for the future of this line of literary and scientific labor.
The bacteriologists have led and mycologists should follow.
The following pages represent in a much extended form the lectures
and laboratory exercises given by the author before his botanic classes
at the University of Pennsylvania, and before public audiences else-
where, especially, Farmers' Institutes with which he has had three
years' experience as a lecturer in Pennsylvania. The arrangement of
the text has been suggested by the needs of the classroom and from an
acquaintance with similar work in other colleges and universities in
America. It is hoped that the book and the suggestions, as to teaching
which it contains, will appeal to those responsible for similar courses.
The keys are given with the anticipation that they will prove useful
to the student and teacher who desire exercises in the classification of
the fungi: The illustrations have been chosen with care, and credit is
given in all cases for those borrowed from other books and monographs.
The author hopes that the book is reasonably free from misleading
<^
A(ii^ US'? 00
statements, and that prove useful to the teaching and student
it will
body. The which are given in detailed form are designed to

exercises,
acquaint the student with the methods that are used in the cultural
investigation of the bacteria and fungi. It is also designed to introduce
the student to the highly important subject of Technical Mycology.
The modern demands for investigators trained in technical my-
cology are many. The health bureaus of our large cities need men and
women, who can make a study of the milk, water and food supplies.
The men, who are engaged in the fermentation industries, frequently
demand expert information on the bacterial and fungal organisms, that
are either useful, or harmful, in the fermentation process. The bread
baker should have someone to whom questions relative to his, one of the
oldest, arts could be referred. The canner also needs such expert
advice. The farmer depends upon the fertility of his soil for the growth
of crops, and the character of that fertility determines whether his
crop shall be a large or a small one. It is conceded on all sides at
present that fertility due not alone to the chemical character of the
is
soil, but also to other conditions which are quite as influential, such as,
the physical state, the bacterial and fungous flora and the presence or
absence of toxic substances. A study of the mycologic flora of the soil
can only be pursued satisfactorily by those who have been trained in
cultural methods.
Then too the study of plant diseases and animal diseases rests funda-
mentally upon technical mycologic laboratory methods. The alarm-
ing increase of plant diseases has attracted a larger and ever growing
number of young men into the study of bacteriology and fungology.
There seem to be unlimited opportunities for such carefully trained men
and women to get profitable employment in health bureaus, manufac-
turing plants, agricultural experiment stations, and as plant doctors
stationed in our larger towns and cities, ready, as a medical doctor is
ready, to give for a monetary consideration expert advice and treatment.
Lastly, there are chances for men and women trained in technical
mycology to become professors, or teachers, of the subject in our col-
leges and agricultural high schools. Such trained specialists can help
to increase the crop-producing capacity of our farms by eliminating
the prevalent diseases, which reduce seriously the farmers' profits.
Such specialists are conservationists in the truest sense of the term.
PREFACE Vll
The author, with great pleasure, thanks the following persons for
suggestive help in the preparation of the text-book: Professor J. C.
Arthur read the proof of the chapter on the rust fungi; Prof. D. H.
Bergey of the University of Pennsylvania the pages dealing with
laboratory methods. Prof. Mel T. Cook and his associates J. C.
Helyar and C. A. Schwarze of the New Jersey Agricultural Experiment
Station, New Brunswick, read the galley proofs throughout and made
valuable suggestions. Dr. J. S. Hepburn read the pages dealing with
bio-chemistry, Messrs. H. R. Fulton and Donald Reddick also made
valuable suggestions as to the arrangement of the contents of the book,
while Prof. L. R. Jones and Dr. C. L. Shear furnished illustrations for
reproduction in the text. Prof. A. H. Reginald Buller of the University
of Manitoba gave permission to use five illustrations in his book,
Researches on Fungi. The au thor desires to express his thanks for the
'
'
uniform courtesy of members of the firm of P. Blakiston's Son & Co.,

especially to Mr. C. V. Brownlow, whose unfailing interest has done
so much to forward the publication of the work.
J. W. H.
————
CONTENTS
PART I. MYCOLOGY
Page
i
CHAPTER I. General Statement and Classification
CHAPTER II.— Slime Moulds (Myxomycetes) 7
21
CHAPTER III.—The Bacteria in General
Name; Size; Locomotion; Cell Division and Reproduction; Photogens;
Chromogens; Thermogens; Aerobism and Anaerobism.
CHAPTER IV.— Classification of Bacteria • 28

Bacteria;
According to Nutrition; Prototrophic Bacteria; Metatrophic
Bibliography.
Paratrophic Bacteria; Systematic Account of the Bacteria;
CHAPTER V. Characteristics of the True Fungi 42
CHAPTER VI. Histology and Chemistry of Fungi 52
Histology; Cell Contents; Colors; Physiology; Enzymes;

Classification of
Enzymes in Fungi; Chemotaxis.

61
CHAPTER VII.— General Physiology of Fungi
Influence of Light; Luminosity; Liberation of Spores.
CHAPTER VIII.— Ecology of Fungi 69

Sclerotia; Galls; Habitats; Xerophytism;
Saprophytes and Parasites;
lachen Fungi.
Fungi and Geographic Distribution 82
CHAPTER IX.— Fossil
of Lichens; Distribution of
Fossil Fungi; Geographic Distribution; Habitats
Chestnut Blight; Laboulbeniaceae; Family Clathraceae.
^9
CHAPTER X. Phylogeny of Fungi
CHAPTER XL— Mould Fungi ' ' ^^
Order Zygomycetales; Sexual Reproduction; Spores and
Sporangia; Fer-
Mucoracese;
mentation; Key to Families of the Order Zygomycetales;
IÎortierellacea;; Choanephoracea;; Chaetocladiacea;; Piptocephalidaceae;
Entomophthoraceae; Bibliography.
CHAPTER XII.— Oospore-producing Algal Fungi 107

to Families; Mono-
Sexual Reproduction; Haploid and Diploid State; Key
blepharidaces; Saprolegniaceae; Peronosporaceae; Generic Key
to Family
Peronosporaceee.
ix'
— ——
X CONTENTS
Page
CHAPTER XIII.— OoMYCETALES (Continued) ii6
Chytridiaceae; Ancyclistaceae; Bibliography.
CHAPTER XIV.— Higher Fungi 120
Ascomycetales; Sexuality, Claussen and Harper; Life Cycle; Bibliography.
CHAPTER XV. Sac Fungi in Particular (Yeasts, etc.) 131
Endomycetaceae, Exoascaceae; Saccharomycetaceas; Yeasts, cells and fer-
mentation, etc.; Systematic Position.
CHAPTER XVI.— Sac Fungi (Continued) 143
Gymnoascaceae; Aspergillaceae; Elaphomycetaceae; Terfeziaceae; Tuberaceae
(TruflBes) ; Myriangiaceae.
CHAPTER XVIL— Mildews and Related Fungi 154
Erysiphaceae (Mildews); Perisporiaceae; Microthyriaces; Hypocreaceae;
Dothideaceae; Sordariaceae; Chaetomiaceae; Sphaeriaceae; Valsaceae; Melo-
grammataceae; Xylariaceae; Hysteriaceae; Phacidiaceae; Pyronemaceae;
Ascobolaceae; Pezizaceae; Helotiaceae; Mollisiaceae; Geoglossaceae; Helvel-
laceae; Cyttariaceae; Rhizinaceae; Phylogeny of Ascomycetales; General
Bibliography.
CHAPTER XVIII.— Basidia-bearing Fungi (Smuts) 177
Key to Suborders; Ustilaginaceae (Smuts); Bibliography of Smuts.
CHAPTER XIX.— Rust Fungi 187
General Structure; Forms; Life Cycles; Cytology; Phylogeny; Endophyl-
lacese; Coleosporiaceae; Pucciniacese; Bibliography of Rusts; Auriculariaceae;
Tremellaceae (Trembling Fungi).
CHAPTER XX.— Fleshy and Woody Fungi 218
Cytology; Dacryomycetaceae; Exobasidiaceae; Hypochnaceae; Thele-
phoraceae; Clavariaceae; Hydnaceae; Polyporaceae; Manuals.
CHAPTER XXI.— Mushrooms and Toadstools 231
Agaricaceae;Development of Fruit Bodies; Cultivation of Mushrooms;
Chemistry and Toxicology of Mushrooms; Gasteromycetes; Hymeno-
gastraceae; Tylostomaceae; Lycoperdaceae; Nidulariaceae; Key to; Sclero-
dermacese; Sphserobolaceae; Phallomycetes; Development of Carrion Fungi;

Clathraceae; Phallaceae; Bibliography of Eubasidii.
CHAPTER XXII. Fungi Imperfecti (Deuteromycetes) 258
General Characters; Systematic Position; Sphaeropsidales ; Melanconiales;
Hyphomycetales.
PART II. GENERAL PLANT PATHOLOGY
CHAPTER XXIII. General Consideration of Plant Diseases . . .271

Etiology; Predisposing Causes; Determining Causes; Physical Character of
Soil; Climatic and Meteorologic Factors, Effect of Smoke, etc.; Trauma-
tism; Animate Agents of Disease; Insects.
————
CONTENTS XI
Page
CHAPTER XXIV. Plants as Disease Producers, Epiphytotism, Prophy-
laxis 298
Vegetal Agents of Disease; Parasitic Flowering Plants; Fungous Organisms
as the Cause of Disease; Mechanic Injuries; Injuries Due to Meteorologic
Causes; Infection; Incubation; Duration of Disease; Dissemination of
Fungi; Epiphytotisms (Epidemics); Prophylaxis.
CHAPTER XXV.— Practical Tree Surgery 319
Preventive Measures; Character of Work; Cavity Treatment; Mixing and
Placing the Cement; Metal-covered Cavities; Guying.
CHAPTER XXVI.— Internal Causes of Disease .^26
Enzymes; Panaschiering; Calico; Nutritive Disturbances; Mutations; Mal-

formations and Monstrosities; Graft Hybrids; Chimaeras.
CHAPTER XXVII. Classification of Abnormalities 331
CHAPTER XXVIII. Symptoms of Disease (Symptomatology) 341
Symptoms of Disease; Discolorations; Shot-holes; Wilting; Necrosis,
Dwarfing; Hypertrophy; Replacement; Mummification; Alteration of
Position; Destruction of Organs; Excrescences and Malformations;
Exudations; Rotting; Bibliography of Diseases in General.
CHAPTER XXIX.— Pathologic Plant Anatomy 354
Restitution; Hypoplasia; Metaplasia.
CHAPTER XXX. Pathologic Plant Anatomy (Continued) 364
Hypertrophy; Excrescences; Intumescences; Callous Hypertrophy; Ty-
loses; Gall Hypertrophies; Hyperplasia; Homooplasia; Heteroplasia;
Callus; Conditions of Callous Formation; Wound Wood; Wound Cork.
CHAPTER XXL— Galls 384
Kinds Cataplasms; Histology of Cataplasms; Histology of Galls.
of Galls;
Cecidial Tissue Forms; Bibliography of Galls.
CHAPTER XXXII.^ Mechanic Development of Pathologic Tissues . . 403
General Consideration; Bibliography of Developmental Mechanice, Sug-
gestions to Teachers and Students.
PART III. SPECIAL PLANT PATHOLOGY
CHAPTER XXXIIL— Specific Diseases OF Plants 411

Common and Important
General Statement; Principal Publications; List of
Diseases of Economic Plants in "the United States and Canada Arranged
according to Host Plants.
CHAPTER XXXIV. Detailed Account of Specific Diseases of Plants 475 .
Alfalfa to Grape.
CHAPTER XXXV.— Detailed Account of Specific Plant Diseases
(Continued) 517
Hemlock to Wheat.
— ——
Xll CONTENTS
Page
CHAPTER XXXVI. Non-parasitic, or Physiologic Plant Diseases.. . 564
Root Asphyxiation; Desiccation; Water-logging;
Classification; Stag-head;
Oidema; Frost Necrosis; Apple Fruit Spots; Water-core of Apple; Die-
back, or Exanthema; Mottle-leaf; Curly-top of Sugar Beets; Peach Yel-
lows; Tip-burn of Potato; Leaf -casting; Curly-dwarf of Potato; Bean
Mosaic; Mosaic of Tobacco; Bibliography.
PART IV. LABORATORY EXERCISES IN THE

CULTURAL STUDY OF FUNGI
CHAPTER XXXVIL—Laboratory and Teaching Methods 581
Introductory Remarks; Lesson Micrometry; Lesson 2, Plugging Test-
i,
tubes, etc.; Lesson 3, Microscopic Study of Culture Material; Stains;

Lesson 4, Liquid Nutrient Solutions; Lesson 5, Potatoes as Medium; Lesson
6, Solid Vegetable Substances; Lesson 7, Plant Juices; Lesson 8, Milk, Beer-
wort; Lesson 9, Bouillon; Lesson 10, Eggs; Lesson 11, Nutrient Gelatin;
Lesson 12, Agar-Agar; Lesson 13, Various Nutrient Agars; Lesson 14,
General Directions for Making Plant Agars; Lesson 15, Potato Juice Agar;
Lesson 16, Starch Agar; Lesson 17, Culture Media for Nitric Organisms;
Lesson 18,Standardization of Culture Media; Lesson 19, Germination
Studies; Lesson 20, Counting of Yeasts and Bacteria; Lesson 21, Cultiva-
tion of Yeasts on Gypsum Blocks, Method of Pouring Plates, Streak
Method; Lesson 22, Isolation of Fungi; Lesson 23, Water Analysis; Lesson
24, Methods of Identification; Lesson 25, Plate Counter; Lesson 26, Sys-
tematic Bacteriology; Lesson 27; Scheme for the Study of Bacteria; Lesson
. 2.8, Detailed Study of Bacteria; Lesson 29, Directions for the Study of
Pathogenic Fungi.
CHAPTER XXXVIII. Laboratory and Teaching Methods (Continued) 643
Lesson 30, Inoculation Experiments; Lesson 31, Do.; Lesson 32, Do.;
Lesson ^^, Do.; Lesson 34, Do.; Lesson 35, Experiments with Artificial
Wounding of Plants; Lesson 36, Gas Injuries; Lesson 37, Enzyme Diseases;
Lesson 38, Study of Mistletoe; Lesson 39, Wire Worms in Plants; Lesson 40,
Relation of Light to Pathogenic Conditions; Lesson 41, Withering, or Wilt-
ing of Plants; Lesson 42, Methods of Sectioning, Celloidin, Paraffin;
Lesson 43, Freezing and Cutting of Material; Lesson 44, Use of Drawing
and Projection Apparatus, Drawing Methods; Lesson 45, Suggestions to
Teachers and Students; Lesson 46, Content of Field Trips and Excursions.
APPENDIX I.— Fungicides 669
Bordeaux Mixture, etc.
APPENDIX II.— Spray Calendar 680
APPENDIX III. Antisepsis and Disinfection ............ 692
Preservation of Woods.
————
CONTENTS Xlll
Page
APPENDIX IV. Culture of Mushrooms 693
APPENDIX V. Synopsis of Families and Principal Genera of Myxo-
GASTRALES 693
APPENDIX VI. Key for the Determination of Species of Mucor . . . 695
APPENDIX VII. Keys for the Determination of Species of Asper-
gillus and Penicillium 702
APPENDIX VIII. Keys to the Genera of the Erysiphace^ 721
APPENDIX IX.— Collection and Preservation of Fleshy Fungi . . .726
APPENDIX X. List of Keys to Fleshy Fungi and Selected Keys of
Fleshy Fungi 729
APPENDIX XI.— Key to Agaricace^ 732
Index 753
PART I
MYCOLOGY
CHAPTER I
GENERAL STATEMENT AND CLASSIFICATION

The lower plant organisms which concern the mycologist, or the
student of the fungi, may be considered in a general sense, or in a
narrow way. A general definition would include all those thallo-
phytes, or lower cellular plants (lacking archegonia), which are
destitute of chlorophyll and in its absence become dependent, with
the exception of the prototrophic bacteria, upon extraneous supplies
of organic food, either Hving or dead. This broad definition compels
the mycologist to study the slime moulds, the bacteria and the true
fungi, both as to their morphology and their physiology. He finds on
such study, that broadly speaking, there are similarities of structure
and function in both groups of dependent plants, in fact, he finds that
the function of these plants is connected with cell organization and
structure and vice versa. With this clearly in view, the mycologist
finds that he has to deal with three distinct classes of chlorophylless
plants, namely:
Class Myxomycetes (slime moulds).

Class Schizomycetes (bacteria).
Class Eumycetes (true fungi).
The classification of these colorless (chlorophylless) lower plants
has been elaborated in recent years with considerable detail by various
authors, 30 that the broad fundamental facts both of taxonomy and
phylogeny are known fairly well, but much remains to be done along
the classificatory fines, especially, since the life histories of many
of the bacteria and fungi are incompletely known. It may be
many years before a generally acceptable nomenclature and classifi-
cation win be an accompHshed fact. The choice of a classification by

any worker in mycology depends largely on his training and bias and
on his detailed study of the various groups. No two men would
entirely agree as to which was the best sequence to adopt in a system-
atic treatment of the different forms. The classification adopted in this
fHOPERTfÛBRARY
N. C. State College
2 MYCOLOGY
treatise is based on that of Engler and Gilg, as published in the seventh

illustrated edition of "Syllabus der PflanzenfamiUen," Berlin, 1912, and
on that of Wettstein in his "Handbuch der Systematischen Botanik,"
Leipzig and Vienna, 191 1. Where consistent, the classificatory sys-
tems of these two books are harmonized and any departures which the
student will find from the taxonomic arrangements of Engler and
Wettstein have been made to simplify them by the omission of cer-
tain group names, or to bring the two systems into line with the facts
as at present known. The author has not hesitated to make changes,
where from his experience as a teacher, he has found it best to make
such alterations, especially where, for example, Wettstein uses Ordnung
and Engler Reihe for the same classificatory group, and where in
American usage order and family are used. Then, too, the author has
found it convenient to replace the name of a family, or order, as given
by Engler for one used by Wettstein, or some other author, where such
replacement is recommended by American usage, or where etymolog-
ically the name is more suggestive of the character of the group, and,
therefore, best for the use of students who do not expect to follow out
the intricacies of any system of classification.As the statements and
views of Engler and Wettstein are generally dependable and as their
classification is founded on long experience, as systematic botanists,
it will be found that with respect to the larger subdivisions of the
fungi their classifications are remarkably harmonious. The attempt
has been made simphfy for student use
in the pages that follow to
the facts of classificatory importance and while the groups are ar-
ranged in lineal sequence, it should be explained that true relationship
is expressed better by a family tree with main trunk, larger and smaller
branches. It will be noted that the arrangement of the famiUes, as
given in the two systematic works above mentioned, are sometimes
reversed. The simple groups are given first place, followed by the
more complex.
CLASS I. MYXOMYCETES.
ORDER I. ACRASIALES.
Family i. Guttulinaceae.
Family 2. Dictyosteliaceae.
ORDER II. PHYTOMYXALES.

Family i. Plasmodiophoraceae.
GENERAL STATEMENT AND CLASSIFICATION
ORDER III. MYXOGASTRALES.

Suborder. Exospore^.
Family i. Ceratiomyxacese.
Suborder. ENDOspoREiE.
Family 2. Physaraceae.
Family 3. Didymiaceag.
Family 4. Stemonitaceae.
Family 5. Brefeldiaceae.
Family 6. Cribrariaceae.
Family 7. Liceacese.
Family 8. Tubiferaceae.
Family 9. Reticulariaceae.
Family 10. Trichiaceae.
CLASS II. SCHIZOMYCETES.

ORDER I. EUBACTERIALES.
Family i. Coccaceae.
Family 2. Bacteriaceae.
Family 3. Spirillaceae.
Family 4. Phycobacteriaceae (Chlamydobacteriaceae).

Family 5. Thiobacteriaceae (Beggiatoaceae).
Family 6. Actinomycetaceje (position doubtful).
ORDER II. MYXOBACTERIALES.

Family i. Myxobacteriaceae.
CLASS III. EUMYCETES.

Subclass. Phycomycetes.
ORDER I. ZYGOMYCETALES.
Family i. Mucoraceae.
Family 2. Mortierellaceae.
Family 3. Choanephoraceae.
Family 4. Chaetocladiaceae.
Family 5. Piptocephalidaceae.
Family 6. Entomophthoraceae.
MYCOLOGY
ORDER II. OOMYCETALES.
Family i. Monoblepharidacejr.
Family 2. Saprolegniaceae.
Family 3. Peronosporaceae,
Family 4. Chytridiaceae.
Family 5. Ancyclistaceas.
Subclass. Mycomycetes.
ORDER III. ASCOMYCETALES.

Suborder A. Protoasciine^.
Family i. Endomycetaceae.
Family 2. Exoascaceae.
Suborder B. Saccharomycetiine;1':
Family i. Saccharomycetaceas.
Suborder C. Plectasciine^.
GEN-ERAL STATEMENT AND CLASSIFICATION
Suborder F. Discomycetiine^,
Family i. Hysteriaceae.
Family 2. Phacidiaceae.
Family 3. Pyronemaceae.
Family 4. Ascobolaceae.
Family 5. Pezizaceae.
Family 6. Helotiaceae.
Family 7. Mollisiaceae.
Family 8. Celidiaceae.
Family 9. Patellariaceas.
Family 10. Cenangiaceas.
Suborder G. HELVELLiiNEiE.
Family i. Geoglossaceae.
Family 2. Helvellaceae.
Family 3. Cyttariaceae.
Family 4. Rhizinaceae.
Suborder H. Laboulbeniine^.
Family i. Peyritschiellaceae.
Family 2. Laboulbeniaceae.
Family 3. Zodiomycetaceae.
ORDER IV. BASIDIOMYCETALES.

Suborder. Hemibasidiine^.
Family i. Ustilaginaceas.
Family 2. Tilletiaceas.
Suborder. Uredine^. (Usually Order Uredinales).

Family i. Endophyllaceae.
Family 2. Melamsporaceae.
Family 3. Pucciniaceae.
Family 4. Coleosporaccce.
Suborder. Auricularin^.
Family i. Auriculariaceae.
Family 2. Pilacraceae.
Suborder. Tremellin^.
Family i. Tremellaceae.
6 MYCOLOGY
Suborder. Eubasidiine^.
A. Hymenomycetes.
Family i. Dacryomycetaceae.
Family 2. Exobasidiaceae.
Family 3. Hypochnaceae.
Family 4. Thelephoraceae.
Family 5. Clavariaceas.
Family 6. Hydnaceae.
Family 7. Polyporaceae.
Family 8. Agaricacese.
B. Gasteromycetes.
Family i. Hymenogastraceae
Family 2. Tylostomaceae.
Family 3. Lycoperdaceae.
Family 4. Nidulariaceae.
Family 5. Sclerodermacese.
Family 6. Sphaerobolacese. -
C. Phallomycetes.
Family i. Clathraceae.
Family 2. Phallaceae.
Fungi Imperfecti (Deuteromycetes).

ORDER I. SPH^ROPSIDALES, with 4 families.
ORDER II. MELANCONIALES, with i family.
ORDER III. HYPHOMYCETALES, with 4 families.
The above classification has been given in outline with the object
of presenting to students the information which is requested frequently
of the professor in the class room. A detailed presentation of the spe-
cial morphology, histology, embryology and taxonomy of each group
will be given in the pages which follow, omitting matters concerning
pathology and practice. A separate section of this treatise will be
devoted to the consideration of fungous diseases of plants and their
treatment.
CHAPTER II
SLIME MOULDS (MYXOMYCETES)

CLASS I. MYXOMYCETES
Considerable attention has been given in recent years to the sHme
moulds on account of their biologic interest, taxonomic relationship
and disease-producing forms. As organisms, they have been bandied
about. They have been claimed by zoologists and botanists alike,
for in certain stages of their life cycle they strongly suggest the protozoa,
such as the amoeba. Perhaps on account of this uncertainty one would
be justified in placing the slime moulds in the class Protista of Haeckel,
which group was intended to include all such primitive organisms
which naturalists have been unable to put satisfactorily either in the
animal, or the vegetable kingdoms, but which partake of the nature of
both the animal and the plant phylae. Hence we would have as a
tentative arrangement
Protista
/ \
Protozoa
/ \Protophyta
where the Protista represent the primitive stock of organisms which
have given rise to simple animals on the one hand, or primitive plants
on the other.
Fries and some of his predecessors considered that the slime moulds
were puffballs (Gasteromycetes) and the expression of this view is
suggested in the name Myxogastres given them by Fries in 1833.
Wallroth in 1836 viewing them as related to the fungi termed them
MYXOMYCETES. De Bary, the German botanist, in 1858, impressed
by their closer relationship with the animal world, called them Myce-
TOZOA. Zo'pi in 1885 describes them and Rostafinski,
as Die Pilzthiere
a pupil of De Bary, working under his supervision in an elaboration
of a monograph of these organisms, calls them Mycetozoa. We, there-
fore, are limited by strict priority to adopt the name Myxogastres
for them; but there are valid reasons why the name Myxomycetes
7
8 MYCOLOGY
should be used. One of the strongest arguments is thai if we consider

them as plants they belong to the phylum and hence this
of the fungi
name Myxomycetes aligns itself with Schizomycetes and Eumycetes

generally adopted for the other groups of fungi. It conduces to clarity
and simplification of classification to adopt the name of Wallroth for
the class of organisms incapable of an independent existence, being
destitute of chlorophyll and mainly saprophytic. The older name is
retained, however, as the name of the third order of Myxomycetes,
hence there should be little criticism of the view taken above. The
Myxomycetes (Mycetozoa, Schleimpilze, Pilztiere, Slime Moulds) are
chlorophylless organisms. Their vegetative condition is known as a
Plasmodium which is a naked streaming mass of protoplasm. Repro-
duction is by means of spores produced as exospores, or endospores,
the latter in sporangia, gethalia, or plasmodiocarps. The spores give
rise to amceboid cells or flagellate swarmers which unite later to form
the Plasmodium, or develop directly into the plasmodium.

ORDER I. ACRASIALES.—The members of this order live on
the excrements of animals and on the decaying parts of plants. They

commence development with the escape of an amoeboid body
their
from the walls of the spore and then move about by creeping move-
ments, never assuming ciHa for locomotion. The amoeboid cells pile
up on one another without coalescing to form what has been called an
aggregate plasmodium, and they remain distinct, and artificially sepa-
rable, though closely packed together until the fructification forms,

when they rise above the substratum and form bodies of definite
shape. Every one, or the majority of these definitely arranged amoe-
boid bodies, becomes a spore covered by a dehcate membrane and of an
average size of 5 to 10 m- These heaps of spores resemble the sporangia
of the true shme moulds, but there is no distinct sporangial wall,

the spores being held together by a structureless enveloping substance.
The plants of this group are saprophytes. Gutkdina rosea lives on
decaying wood in Europe. Dictyosteliuni mucoroides is frequent on
old dung, while Acrasis granulata is found on old yeast cakes. Poly-
sphondylium violacewn occurs in southern Europe on manure.
ORDER II. PHYTOMYXALES.—The shme moulds of this order
are parasites which live in the cells of higher plants. The plasmodium
is limited by the cell walls of the host plants, and has its origin in
amoeboid cells which enter and infest the host cells, resulting in a
SLIME MOULDS (mYXOMYCETES) 9
stimulation of the host to form gall-hke swelUngs. The whole Plas-

modium is later transformed by division into a greater or less number
of parts, which become surrounded by membranes to form spores.

The spores are free in the cells of Plasmodiophora, while in SorosphcBra
and in Tetramyxa they are clumped, and surrounded by a delicate
membrane. The order includes a single family:
—
Family i. Plasmodiophorace^. ^The characters of this family are
coincident with those of the class as given above. The family includes
four genera distinguished as follows:
A. Spores distinct from each other, irregularly aggregated and fiUing

the host cells.
{a) Spores regular in shape, spheric, (i) Plasmodiophora.

(b) Spores irregularly shaped, rod-like, or angular. (2) Phytomyxa.
B. Spores united into clumps inclosed by a delicate membrane.
{a) Spores united in groups of four each. (3) Tetramyxa.
(b) Spores in greater number, united into hollow spheres. (4)
Sorosphcera.
The genus Plasmodiophora comprises possibly three species found

in Europe and America. They are parasites in the parenchyma cells
of the cortex of the roots of the higher plants, where they produce
gall-like swellings. The plasmodium fills some of the living cells of
the host. The spores formed subsequently are spheric and lie free
within the host cells. The best known species is P. brassicce which
is the cause of a serious disease known as club foot, or finger and toes
(Fig. i). The symptoms of the disease, the relationship of host and
parasite, will be described in a subsequent section of this book. Two
other species have been described, viz., P. alni in the roots of the alder;
and P. eleagni in the roots of Eleagnus, the silverberry. Considerable
more study will have to be made of the organisms in the roots of the
alder and silverberry before we can definitely place the causal organ-
isms. Tentatively, we may adopt the generally accepted view of the
systematic relationship of the two responsible organisms until later
investigation either proves or disproves the nature of the parasites
attacking Alnus and Eleagnus.
The genus Phytomyxa is represented by two species which live as
parasites in the roots of Hving plants and cause tuber-like enlargements.
The Plasmodia fills the host cells, and later, the irregularly shaped
MYCOLUGY
—
Pig. I. Club-root of cabbage, Plasniodiophora brassicce. i, Turnip with club-
root; 2, section of cabbage root with parenchyma cells filled with slime mould; 3,
isolated parenchyma cell, (v) vacuole, (0 oil-drops in Plasmodium, (/>) Plasmodium;
4, lower cell with Plasmodium, upper cell with spores developing; 5, parenchyma
cell with ripe spores; 6, isolated ripe spores; 7, germinating spores; 8, myxamoeba.
{Figs. 2-8, after Woronin in Soraucr, Handbuch der PJlanzenkrankheiten, 1886, p. 72.)
SLIME MOULDS (mYXOMYCETES) . II
spores fill the infested host cells. Two species have been described.
The nature of Ph. leguminosanim is doubtful, as it may have been
•confused with one of the stages of the nodule-producing bacteria,
which are found in the roots of leguminous plants.
The parasitic slime mould, Tetramyxa, occurs as one described
species Tetramyxa parasitica, which lives in the stems and flower stalks
of water plants, as Ruppia rostellata, where it causes tubercles 0.5 to
I mm. in diameter. Each host cell contains numerous colorless spores
united into tetrads.
SorosphcBra is represented in Germany by S. veronica; found in the
stems and petioles of Veronica and V. chamce-
hederifolia, V. triphylla
drys. The cells of the galls are swollen and filled with numerous
spheric or ellipsoidal brown balls, 15 to 22 /x in diameter, formed of a
single layer of spores united into a hollow sphere and covered externally
by their pellicle.
ORDER III. MYXOGASTRALES.— This order includes the true
slime moulds which are non-parasitic, but live on decaying organic
material, such as old logs, leaf mould compost heaps,
in the forest,
spent tan bark and other organic debris in the woods, and fields,
along the roadsides. One form grows over the grass of lawns and
smothers the grass with its plasmodium and later by its sporangia and
spores. The plasmodium is a naked mass of protoplasm usually of a
reticulate structure and multinucleate. It arises by the union of the
myxamoeba which are developed from the flagellate myxomonads by
the loss of the vibratile flagella. Such a plasmodium is known as a
fusion Plasmodium.^ It usually assumes a reticulate, or net-like,
structure and currents of protoplasm are seen flowing along the strands
of greater or less thickness of which the plasmodium is composed. The
central portion of each current is denser and moves more rapidly than
the marginal clearer protoplasm. Perhaps we are justified in stating
that the outer protoplasm is the ectoplasm and the inner granular
cytoplasm containing food substances and other included substances
is the endoplasm. For some time the plasmodium may flow in a given
direction and later it may reverse its course, moving in an entirely
opposite direction. The color of the plasmodium difi"ers in different
species, as the following table will show. White or yellow seem to be
the more usual colors.
1 In Lahyrinlhiila Cicnkowskii parasitic in Vauchcria the plasmodium is filamen-
ous.
12 . MYCOLOGY
Yellow Fiiligo seplica.
Orange TricJiia scabra.
White Pliysai'um. cUipsoidcum.

Lead-colored Crihraria argillacea.
Pink Enteridiv.m splendens.
Ruby-red , Hemitrichia vesparum.
Red Tubifera ferruglnea.
Scarlet Cribrarla purpurea.
Brown Tubifera Casparyi.
Violet Cribraria violacea.
The movement of the plasmodium is associated with the incorpora-

tion of food. The yellow plasmodium of Badhamia utricularis has
been most carefully studied in its relation to a food supply. It can be
cultivated on such woody fungi as Stereum hirsutum, over which it
extends, devouring by enzyme action the more delicate hyphae.

Thus nourished, it will spread over the moist filter paper inside of the
covering bell jar until I have seen the plasmodium hanging down
like from the inner top of the bell jar. Such a captive
stalactites
Plasmodium has been fed by the writer pieces of mushroom
Agaricus campestris. Shaggymane, Coprinus comatus and beefsteak
have been placed on the surface of the protoplasm and in a few hours
these substances have been found in advanced stages of digestion.
Cheese is reluctantly invaded and is more refractory. The plas-
modium is responsive to changes in the moisture surroundings. It
moves toward a more abundant water supply. It is hydrotropic.

It moves against a current of water and is, therefore, rheotropic.
When highly illuminated, the plasmodium moves away from the
lighted surface. It is negatively heliotropic. If there is a sudden
change in the watery environment, the plasmodium will become
massed into a cake-like lump in which form it remains as a sclerotium,
macrocyst, or phlebomorph, if the substratum loses its water supply.
In the sclerotial condition, the writer has kept a plasmodium for
nine months on a plate of glass placed inside of a laboratory case

in an absolutely dry condition. It was started into activity at the
end of this period of rest on restoring free water to it again, and
by feeding it mushrooms, it was kept in its restored activity for
several weeks beneath a bell jar. The plasmodial stage may be pro-
longed for an indefinite period, if the environmental conditions of
temperature, Hght, moisture and food, are favorable. The writer has
3
SLIME MOULDS (m\ÔMYCETES) 1
kept a Plasmodium in a streaming condition for over a month be-

neath a bell jar. Physarum psitiacinuni, which inhabits the rotten
stumps of old trees, appears to pass a year as a plasmodium.
The early stages in the formation of the sporangium have been de-
scribed in Comatricha oUusata. When the fruiting period is reaclifd,
the watery- whiteplasmodium issues from the wood crannies and spreads
over an area perhaps half an inch across. The plasmodium is seen to
concentrate in thirty or forty centers and in an hour or two each
center has by rhythmic pulsation of the protoplasm risen into, a pear-
shaped body with a slender base and an enlarged upper portion. The
black hair-Uke stalk has grown to its full length in six hours and on
its summit is borne the young sporangium, which is a white viscid
globule of protoplasm. A pink flush now begins to appear in the

sporangium. The included nuclei are like those of the plasmodium
at first, but later as spore formation proceeds they divide mitotically.
The sporangia of the different slime moulds take various forms which
will be described in general in the systematic generic keys which
follow. They may be either symmetric or irregular in shape, sessile or
stalked. The irregular sessile forms, which simulate the net-like
appearance of the streaming protoplasm, are called plasmodiocarps.
When the fruit body is fiat and cake-like with separating walls imper-
fectly developed it forms an cethalium. The protoplasm which is
left on the substratum and dries down as a film-like residuum is known
as the hypothallus (Figs. 2 and 3).
The changes which take place in the formation of spores and
capilhtium have been minutely studied in a number of sUme moulds.
We owe much to R. A. Harper, E. W. Olive and B. O. Dodge in America
and to E. Jahn Germany for our knowledge of these processes. The
in
process in Didymium melanospermum, according to Harper,^ is as
follows: The spore plasm condenses so that it is finely granular in the
peripheral region and central region near the columella and foamy
vacuolar in the middle zone. The capillitium is already formed before
the condensation of the protoplasm has been accomplished. It con-
smooth threads which pass radially outward from the central
sists of
dome-shaped columellar cavity to the sporangial wall. The threads

of the capillitium are attached at their ends. The protoplasm is in
contact with these threads and at this stage the nuclei are scattered
1 Harper, R. A.: Amcr. Journ. Rot., i: 127-144, March, 1914.
—
14 MYCOLOGY
rather uniformly through the spore plasm and are of unequal size.
Vacuoles are formed in a still further condensation of the sporangial

-protoplasm and each of these apparent vacuoles is pierced by a capilli-
tial thread which runs through its central axis. Droplets of water are
formed along the capillitial thread as a still further evidence of water
extrusion. Cleavage planes now appear at the periphery of the mass
of sporangial protoplasm and progress inwardly toward the center.
The process of cleavage parallels the extrusion of water and the for-
mation of the blocks of protoplasm by these cleavage lines is assisted
Fig. 2. -A, B, Comatricha nigra. A, Sporangium, natural size; B, capillitium,

20/1; C, E, Stemonitis fusca; C, sporangium, natural size; D and E, capillitia, 5/1,
20/1; F, H, Enerthema papillatum, F, unripe; G, mature sporangium, lo/i; H, capil-
litium, 20/1. (C, D, after nature. A, F, G, H, after Rostafinski; B, E, after de Bary
in Die natiirlichen Pflatizenfamilien I. i, p. 26.)
by the presence of the vacuoles. The splitting up of the irregular blocks

of protoplasm, which have the nuclei irregularly distributed through
them, proceeds until the protoplasmic blocks arebinucleated,and before
this the nuclei are seen in various stages of division which proceeds
irregularly in Didymium, while in Fuligo the division of the nuclei is
simultaneous in a particular spore sack. The plasma membranes of

the capillitial openings are the source of cleavage furrows to even a
greater degree than the original surface plasma membrane of the spore
sack as a whole. In Fuligo in the final stages of spore formation the
spore plasm is condensed about the nuclei, but in Didymium, the ultimate
SLIME MOULDS (mYXOMYCETES) 1$
result of the progressive cleavage in f urrowin,^ is the formation of uninu-

cleated rounded spores. They he packed between the capillitial
threads.
Most genera moulds have a capiUitium (Figs. 2 and 3)
of slime
consisting of a system of threads, and as we have seen, it appears be-
fore the spores are formed. When the capiUitium extends from the
base of the sporangium, it is associated with a columella (Fig. 2). It
differs widely in the dififerent genera of the groups. In some genera, as

Trichia and Arcyria, the capiUitium consists of free threads, or elaters.
In those genera in which calcium carbonate is present in the sporangia,
it is found in the capiUitium usually when several threads meet forming
then the so-caUed hme knots. In Dictydimn, purplish-red granules
are imbedded in and are known
the threads of the false capiUitium
as dictydin granules. The formation of the capiUitium in certain
myxomycetes has been investigated by Harper and Dodge.^ They
find that the capiUitium is formed by the deposit of materials in the
vacuoles from which the capiUitial thread is formed and that radiating
threads run out from the larger granules which are deposited by the
process of intraprotoplasmic secretion. These radiating fibrUs sug-
gest rather strongly that they are cytoplasmic streams which are
bringing materials for the formation of the capillitial wall and its thick-
enings which are laid down sometimes as spirals, suggesting that the
process is comparable to the ordinary processes of cell-waU formation,
but along internal plasma membranes, rather than external. The

relation of the fibrils to the capillitial granules is best seen where a
capiUitial vacuole runs longitudinally. Strasburger's earher observa-
tions are confirmed by the recent work on capiUitial formation, when
he described the capiUitium of Trichia fallax as originating in vacuolar
spaces in the cytoplasm which elongate and take on the tubular form
of young capillitial threads, while the formation of the wall and spiral
thickenings are due to the deposition of granules as intraprotoplasmic
secretions consisting of microsomes of the membranogenous type.
Where the capiUitial threads are solid they may be called stereone-
mata; where hollow, coelonemata.
The spores are discharged from the sporangia, and if they find a
suitable medium which to grow,- such as free water, they give rise to
in
swarm cells, as amoeboid bodies, or myxamoebse. These soon acquire a
Ânnals of Botany, xxviii: 1-18, January, rQi4.
6
I MYCOLOGY
.flagellum at the anterior end and creep in a linear form with the flagellum
extended in advance, or swim about in the water with a dancing move-
ment occasioned by the lashing of the flagellum. They have a single
nucleus and a contractile vacuole. To a large extent they feed on
bacteria which are swallowed by pseudopodia which project from the
posterior end of the cell. The swarm cells increase rapidly by biparti-
tion. When this takes place, the flagellum is first withdrawn and the
main cell assumes a globular form; it then elongates and a constriction
occurs at right angles to the long axis. The nucleus divides by karyo-
kinesis and in the course of a few minutes the halves of the nuclear
and retreat to the opposite ends of the constricted cell
plate separate
which now divides into two, each new cell acquiring a flagellum.
Sometimes the swarm cells become encysted to form the so-called
microcysts, or zoocysts.
The spores of Ceratiomyxa, which are borne on the outside of
column-like sporophores, are white in color. The surface of the
sporophore is divided into lozenge-shaped areas each with a projecting
stalk bearing a single spore. The nucleus. of these spores, according
to Jahn, twice divide by karyokinesis, and finally, when the spore
germinates, eight amoeboid bodies are liberated, each of which develops
a flagellum and the cluster swims away by the lashing of the flagella.
Finally, these cells separate. myxomycetes have spores
All other
which in germination produce only one myxamoeba.
Spores of Reticularia which had been dry for eight months germi-
nated in thirty-five minutes at a temperature of 21°. Spores exposed
to a temperature of 37° for only five minutes germinated in eleven
minutes. The spores of Stemonitis flaccida germinated in one hour,
those of AmauroclKBte in two and one-half hours, those of Didymium in
four to five hours, while it took the spores of Stemonitis ferruginea in
wood decoction three to five days to germinate.
Some remarkable discoveries have been made with regard to an
alternation of generations in the slime moulds connected with a so-
called sexual act. Jahn, Kranzlin and Olive have worked upon this
problem. The generation in all the Myxomycetes, including Ceratio-
myxa, with the double chromosome number (8)' (diploid condition) in
the nuclei is of short duration. The nuclei of the swarm bodies, amoe-
boid bodies and the plasmodium have the single number (4) of chromo-
somes. Union of the nuclei to form fusion nuclei with double
—
the number (8) of chromosomes immediately precedes the formation

of the sporangia. The reduction division, which results in the forma-
tion of spores, is preceded by synapsis, cUakinesis and heterotypic
nuclear division. Small nuclei and large nuclei are seen. The large
nuclei are probably fusion nuclei. The small nuclei probably
disintegrate.
To the order Myxogastrales belong the majority of the Myxo-
MYCETES (Figs. 2 and 3). Many are found on decaying wood as Dic-
tydium cernumn with black spore contents, Arcyria nutans and A.
Fig. 3. A, B, Leocarpus fragilis. A, Sporangium, natural size; B, capillitium

200/1; C. Craterium leucocephalum sporangia, 6/1; D, Physarum sinuosum spor-
angium, 6/1; E, F, Tilmadoche miitabilis; E, sporangia, 20/1; F, capillitium, 200/1.
(.4, C, D, after nature; B, E, F, after Rostafinski in Die natiirlichen Pflanzenfamilien
I. I, p. 32.)
punicea have net-like capilHtia, the former with yellow, the latter
with a red one. Lycogala epidendrum has a cinnabar-red plasmodium
and a brownish-gray aethalium. Trichia varia, T. chrysosperma, He-
miarcyria clavata have yellow sporangia and golden-yellow spirally
sculptured elaters, Reticularia lycoperdon has a large brown cake-like
aethalium. The yellow plasmodium of Fuligo septica sometimes covers
spent tan bark and is known as "flowers of tan. " It is one of the most
generally distributed of slime moulds and the writer has found its
sethaha on the bark of street trees and even on the bricks of the street
pavements, as yellow-brown, cake-like fructifications crumbling readily
8
1 MYCOLOGY
into a powder. The Plasmodium of a species of Chondriodcrma lives at

the edge of melting snow fields, or even on the snow itself. The organ-
ism of malaria frequently called Plasmodium malaria; is not a slime
mould, but rightly belongs to the group of HdmosporidicB, a division
of the Protozoa.
The sHme moulds are cosmopolitan. Many of the same forms have
been found in North and South America, the West Indies, Europe,
Cape of Good Hope, AustraHa, New Zealand and Japan. The writer
has used a manual of the Myxomycetes of Buitenzorg, Java, in the
identification of species found near Philadelphia. About 214 species are
represented in the British Museum collection.
Laboratory Exercise. —The wjiter has found in his experience as a
teacher that time may be profitably spent by a class in mycology in the
identification of the common slime moulds. The sporangia, aethalia
and plasmodiocarps of the different kinds can be kept separately in
different small pasteboard boxes and material out of these boxes can
be distributed to the members of the class. The dried material is first
treated with 70 per cent, alcohol to remove the air, and then the treated
material is mounted for permanent preservation in glycerine jelly.
The absorption of water by the glycerine jelly is prevented by a ring

of asphalt. The "Guide to the British Mycetozoa exhibited in the
Department of Botany, British Museum Natural History," ist
Edition, 1895, 2d Edition, 1905, 3d Edition, 1909, has been used in
classes at the University of Pennsylvania with much success. After
the generic name has been determined. Lister's "British Mycetozoa"
or MacBride's "North American Slime Moulds" can be used to find
the name of the species.
BIBLIOGRAPHY
CoNARD, H. S.: Spore Formation in Lycogala exigiium Morg. Iowa Acad. Sci.,
17: 83, 1910.

Cooke, M. C: The Myxomycetes of Great Britain Arranged According to the
Method of Rostafinski, 96 pp., 24 pis., London, Williams & Norgate, 1877.
Cooke, M. C: The Myxomycetes of the United States Arranged According to the
Method of Rostafinski. Annals Lyceum, Nat. Hist., New York, 11: 378-409,
1877.
Cook, O. F.: Methods of Collecting and Preserving Myxomycetes. Botanical
Gazette, 16: 263, 1891.
DE Bary, Anton: Comparative Morphology and Biology of the Fungi, Mycetozoa
and Bacteria. Oxford at the Clarendon Press, 1887, especially pp. 420-453.
— 9
Harper, R. A.: Cell and Nuclear Division in Fiiligo varians. Botanical Gazette,
30: 217, 1900.
Harper, R. A.: IVogressive Cleavage in Didymium. Science, new ser. 27: 341,
1908.
Harper, R. A.: Cleavage in Didymium melanospermum (Pers.) Macbr. Amer.
Journ. Bot., i: 127-143, March, 1914, with 2 plates.
Harper, R. A. and Dodge, B. O. The Formation of the Capillitium in Certain
:
Myxomycetes. Annals of Botany, xxviii: 1-18, January, 1914, with 2 plates.

Harshberger, J. W.: Observations upon the Feeding Plasmodia of Fuligo septica.
Botanical Gazette, 31: 198-203, 1901.
—— — Distribution of Nuclei in the Feeding Plasmodia of Fuligo septica. Journ. of
Mycology, 8: 158-160, 1902.
• — —A Grass-killing Slime Mould (Physarum cinereum). Proc. Amer. Philos.
Soc, 45: 271-273, 1906.
Jahn, E.: Myxomyceten Studien. Ber. Deutsch. Bot. Gesellsch. I. Dictydium
umbilicatum, 19: 97-115, 1901; II. Arten aus Blumenau, 20: 268-280, 1902;
III. Kernteilung und Geisselbildung bei den Schwarmen von Stemonitis flaccida,
22: 84-92, 1904; IV. Die Keimung der Sporen, 23: 489-497, 1905; V. Listerella
paradoxa, 24: 538-541, 1906; VT. Kernverschmelzungen und Reduktionsteilun-
gen, 25: 23-26, 1907; VII. Ceratiomyxa, 26": 342-352, 1908; VIII. Der Sexualakt,
29: 231-247, 1911.
Kranzlin, H.: Zur Entwicklungsgeschicte der Sporangien bei den Trichlen und
Arcyrien. Archiv Protistenkunde, ix: 170-194, 1907.
Lister, A: Notes on the Plasmodium of Badhamia utricularisand Brefeldia maxima.
Annals of Botanj^, 2: 1-24, 1888.
Lister, A.: On the Division of Nuclei in the Mycetozoa. Linn. Soc. Journ., xxxix:
529, 1893.
Lister, A.: A Monograph of the Mycetozoa, 224 pp., 78 pis., p. 894.
Lister, A: Guide to the British Mycetozoa Exhibited in the Department of Botany,
British Museum of Natural History, ist Edition, 1895; 2d Edition, 1905;
3d Edition, 1909.
MacBride, T. H.: The North American Slime Moulds, being a list of all Species
hitherto described from North America including Central America, pp. xvii +
269: 16 pis., MacmiUan Co., 1899.
MacBride, T. H. On Studying Slime Moulds. Journ. Applied Microscopy, 2:
^
:
585-587, 1899.
MacBride, T. H.: The Slime Moulds. Rhodora, 2: 75-81, 1900.
Masses, G.: A Monograph of the Myxogastres, 336 pp., 12 pis., London, Methuen
& Co., 1892.
Olive, Edgar W.: Cytological Studies on Ceratiomyxa. Trans. Wise. Acad. Sci.
Arts and Letters, XV : 753-773.
Monograph of the Acrasieae. Proc. Boston. Soc. Nat. Hist., xxx: 451, 1902.
Evidences of Sexual Reproduction in the Slime Moulds. Science, new ser.
xxv: 266, 1907.
Penzig, O: Die Myxomyceten der Flora von Buitenzorg, Leiden, 1898.
20 MYCOLOGY
Rex, G. a.: The Myxomycetes, Their Collection and Preservation. Botanical

Gazette, lo: 290, 188';.
ScHROETER, J.: Myxogasteres in Engler and Prantl; Die naturlichen Pflanzenfam-

ilien, i: Abth. i, pp. 1-35, 1889-92.
Schwartz, E. J. The Plasmodiophoraccce and Their Relationship to the Mycetozoa
:
and the Chytrideae. Annals of Botany, xxviii: 227, 1914.

Strasburger, E.: Zur Ent\vickelungegeschichte der Sporangien von Trichia fallax.
Bot. Zeitung, xliii: 305-16; 321-3, May 16, 1884 and May 23, 1884.
Sturgis, W. C: The Myxomycetes of Colorado, No. i. Science, ser. xii. No. I,
pp. 1-43; general ser. No. 30, September, 1907; No. II. Science, ser. xii, No.
12, pp. 435-454, April, 1913; Colorado College Publications.
Sturgis, W. C: A Guide to the Botanical Literature of the Myxomycetes from
1875 to 1912. Science, ser. xii. No. 11, pp. 385-434. June-September, 1912,
Colorado College Publication.
ZoPF, W.: Die Pilzthiere oder Schleimpilze, i-vi -f 1-174 pp.. Figs. 1-51, Breslau.
CHAPTER III
THE BACTERIA IN GENERAL

CLASS II. SCHIZpMYCETES
The name Schizomycetes comes from two Greek roots (ax'i-ôi,
I split + ijLVKr]s, a fungus) which combined are equivalent to the term

splitting fungi, or fission fungi in allusion to the manner in which the
bacterial cells divide. The Germans call them Spaltpilze, which is
the German way of expressing the same thing. The name bacteria
is American science used in a general sense to include all of the
in
Schizomycetes without reference in particular to the genus Bacterium.
In popular use, such as newspaper articles, these lowly plants are
described as germs, microbes, or microorganisms. These English
synonyms are, however, inexact, having different shades of meaning
and are used in different ways in common speech, as consultation with
any large dictionary of our language will show. There is no ambiguity,
if we speak of all the Schizomycetes as bacteria, or bacterial organisms.
These plants are generally unicellular, or the single cells are united
into a coenobium. These coenobia are filamentous, sheet-Uke, or in
groups, seldom arranged in fructification-like masses of definite form,
as is the case with the Myxobaderia. All cells of the coenobium are
alike and only in the highest developed forms do we find a differentiation
into basal cells and filament cells. The heterocyst, found in the blue-
green algae, is totally absent. The cells of bacteria are the smallest
of plant cells; for example: Micrococcus progrediens has a diameter
of o.i5ju and Spirillum parvum has a thickness of o.i to 0.3^1, but yet
smaller are the ultramicroscopic organisms, which have come into
prominence recently as the cause of certain diseases. The smallest
bacteria stand at the borderline o,f what is with the best lenses and
optimum illumination the practical limit of microscopic vision. On
the other hand, with the application of the ultraviolet light of short
wave length in microphotography, it has been possible to obtain an
image of small objects whose enlargement has been 4000-fold. It
has been possible with the ultramicroscope of Siedentopf and Zsig-
MYCOLOGY
mondy to demonstrate small particles whose size is only many million

times that of a miUimeter. The accompanying figure (Fig. 4) adopted
from Fuhrmann' represents the relative size of the spheric bacteria
and the rod-shaped organisms, while the breadth of the largest known
bacterial cell, that of Beggiatoa mirabilis, which approaches that of a
human hair in thickness, is represented in the larger area where the
width of the cell is twice its length.
4. — Diagram representing the relative sizes of spheric and rod-shaped bacteria

{After Fuhrmann.)
,
THE BACTERIA IN GENERAL 23
and the numerous individual forms are united into slimy, skin-like
or lumpy masses known as zoogloea.
The interior of the cell shows no differentiation into nucleus and
cytoplasm, but the nuclein in certain forms seems to be scattered in
the plasma (Fig. 5). Considerable diversity of opinion exists as to the
nature of the cell substance of bacteria. The uniform staining of the
cell by ordinary methods suggests that the cell substance is all cyto-
plasm without nucleus. An opposite opinion is that the cell substance
is composed almost entirely of nuclear matter (chromatin) with perhaps
a thin layer of ectoplasm. Another view is that

of Zettnow (Zeitschr. f. Hyg., 1899: 18), who regards
the cell body of bacteria as composed largely or
almost wholly of chromatin mingled with varying
amounts of cytoplasm. This, however, can be said,
that it is fairly certain that bacteria contain both
chromatin and cytoplasm which vary in amount and
position in different cells (Fig. 5). The cell mem- Fig. 5.— i,c/!)'o-
brane is mostly colorless; seldom does it appear BTg^glai'oT"aibl'
greenish or rose-red, as in the purple bacteria. Bacteria with a cen-
When colonies of bacteria are colored, the coloring
ch^'romatln °g'rtTns
matter is an excretion product. which are considered
Locomotion. —
The movement of many bacteria is
equiv^rienTofâ
a true movement from place to place, not merely a nucleus. (From
Brownian movement. It is accomplished in nearly f^l"'sfLTE7uion,
all cases by the presence of cilia, or.flagella, which p. 91- After
^"^^'^''^^^
by some are considered to arise directly from the
cell membrane," by other investigators to arise from the ectoplasm
within; its origin in some way associated with a blepharoplast. Which-

ever view is the correct one, the motile filaments can in some large
spirilla be seen in the living unstained organism, but generally it re-
quires special methods of treatment and staining to make them out.
Great differences exist as to their distribution. Some forms, as the
cholera bacillus, have a single flagellum at one pole {monotrichous) ;
others, as many have a flagellum at each pole {amphitrichous)

spirilla, ;
others, as certain large spirilla, have a tuft at one pole {lophotrichous) ;
while others have cilia covering the whole cell, as the typhoid organism
{peritrichous) . Many organisms are without cilia, or flagella {atrichous)
and hence are non-motile.
—
24 MYCOLOGY
Cell Division mid Reproduction. — As with other plant cells in general

it may be said that growth is not conditioned on cell division. Growth
is the enlargement of the cell, not merely a swollen condition, and this
increase in size within definite limits for each species, which can be
is
determined by statistic study. As long as division is not preceded

by nuclear division, the term fission is applicable. Certain students
of the group claim that there is a division of the nuclear substance
(Fig. 6), and Fuhrmaftn actually figures division of the nuclear mate-
rial in such forms as Bacillus nitri,
''^-
^ Micrococcus butyricus, Spirillum
®
i_^ / ^ W volutans and the potato bacillus.
^ L ^' Possibly then division of the
^. nuclear substance precedes that
-O of cell division, and if that phenom-
enon is found general, the term
fission is no longer apphcable.
Fig. 6. Fig. 7.
Fig. 6. Bacterium gammari. a, b, c. Cells with typical nucleus of nucleoplasm,

surrounded by a nuclear membrane and by one or two karyosomes also showing
karyokinesis; d, a filamentous bacterium from intestine of an annelid worm, Bryo-
drilus chlorii, each cell with a nucleus. {From Marshall. Microbiology, Second Edi-
tion, p. 89, after Vejdowsky.)
Fig. 7. —Cells of Bacillus megatherimn. i. Polar granules as nuclei; 2, increase
in size of nucleus at time of sporulation; 3,
same; 4, change in size of nucleus which
issurrounded by a membrane and becomes a spore. (Fro?n Marshall, Microbiology,
Second edition, p. 90, after Penan)
Cell division may take place quite rapidly under favorable conditions.
Bacillus siihtilis divides in thirty minutes; Vibrio cholera, every twenty
minutes. The young cells attain full size in a short space of time.
Bacteriologists have estimated, that if bacterial multiplication was
unchecked and the division of each cell was accomplished inside of an
hour that in two days the descendants of a single cell would number
281,500,000,000, and that in three days the offspring of a single cell
would weigh 148,356 hundredweights. Lack of food, accumulation of
bacterial products injurious to the organisms that formed them explain
why their rapid multiplication is kept in check.
fformr library
Ml r Qtnt0 Cnllpift
The spores formed by the bacteria are of two kinds, arthrospores

and endospores. Arthrospores are whole vegetative cells which by a
thickening of their walls become resting spores. Some bacteriologists
would not include arthrospores as true spores. The true spores are
formed in the cells and differ from the cells in resisting greater heat
and by other definite structural and physiologic quaUties. (Fig. 7.)
The shape of the cell may be altered with the formation of one or two
spores within (endospores). In the hav bacillus, the spore occupies the
center of the cell and is smaller than the original mother cell, hence the
shape of the parent cell is not altered. Bacterium pants and Bacillus
amylobacter become swollen in the middle when the spore forms so that
the mother cell becomes spindle-shaped. The bacillus of lockjaw de-
velops a spore at one end of the cell, which becomes drumstick-shaped,
hence the German name trommelschlagel for such forms and the generic
name Plectridium now given to cells that produce terminal spores.
Bacillus amylobacter may develop one terminal spore, or two spores,
one at each end of the cell, so that the mother cell becomes dumbbell-
shaped. Bacillus inflatus may develop two spores also.
Spores may germinate at the poles, as in Bacillus BiitscJili and B.
amylobacter ; at the equator, as in Bacillus subtilis and B. loxosporus, or
obliquely, as in Bacillus loxosus. In germination resting spores absorb
water, and become more or less swollen, when the spore membrane
is dissolved and the germ tube protrudes.
The classification of bacteria according to their special activities,

or the products formed by these activities, is useful in presenting
another phase of the subject to the mycologic student. The fact is
noteworthy that we can group the various organisms into the photo-
genic (light-producing), chromogenic (color-producing), thiogenic
(sulphur-producing), zymogenic (ferment-producing), pathogenic (dis-
and thermogenic (heat-
ease-producing), saprogenic (decay-producing)
producing) without reference to their morphology, or genetic rela-
tionship. It is useful to be able to discuss the light, heat, color, etc.,
produced by these organisms as distinct phenomena worthy of experi-
mental treatment.
Photogenic Bacteria. —The phosphorescence associated with decaying
haddocks, mackerel and other sea fishes, the faint glow seen on badly
preserved meats (beef, mutton, veal) and sausages are produced by
photogenic bacteria. Most success is obtained by using sea fishes in
26 MYCOLOGY
experimenting with the phosphorescent bacteria, for these organisms

2 to 3 per cent, of sodium chloride,
require in their culture media from
besides the usual saltsand peptone, the medium should contain some
The number of
other source of carbon, such as sugar, glycerine, etc.
known photogenic bacteria is considerable. Migula names twenty-
five species and Molisch twenty-six. A few need only be mentioned
here, viz.: Bacterium phosphor escens Fischer; Bacillus photogenus
Molisch; B. luminescens Molisch; Microspira glutinosa (Fischer)
Migula; M. luminosa (Beijerinck) Migula; Pseudomonas javanica
(Eijkmann) Migula. The results of numerous experiments are that
the production of light by bacteria is an exclusively aerobic phenome-
non, for in the absence of oxygen, they are non-luminous. The light
issometimes strong enough that jars containing luminous bacteria can
be photographed by the light emitted by the organisms within the jar.
—
Chromogenic Bacteria. Most bacteria are colorless and even in such
forms in which color is associated with their growth on culture media,
the organisms are colorless. The bacillus which causes the "bleeding
host," Bacillus prodigiosus, is colorless with the pigment in the form of
granules scattered about between the bacterial cells. In other cases,
the pigments and fluorescent substances are diffused in the culture
medium outside the living cells. Hence, we may call such bacteria as
chromoparous. The chromophorous species are those in which the
protoplasm is actually colored. Such are some sulphur bacteria
Chromatium and Thiocystis, and finally, there are some forms as Bacillus
violaceus in which pigment is lodged in the cell wall, when we may call
them parachromatophorous. Practically all of the colors of the spectrum
are represented in the color productions of bacteria: violet {Bacillus
violaceus), indigo {B. janthinus), blue {B. pyocyaneus), green {B.
fluor escens), yellow {Sarcina lutea), orange {Sarcina aurantiaca) and red
{B. prodigiosus). The erythrobacteria, or colored sulphur bacteria,
are unique in the power carbon dioxide in the presence
of assimilating
of sunlight by the activity of bacteriopurpurin (a red coloring matter)
which behaves Hke the chlorophyll of green plants.
Thermogenic Bacteria. — Such substances as hay, silage, manure and
cotton waste frequently become heated, the temperature inside the
mass being raised to 60° or 7o°C. This spontaneous heating is due to
the respiratory activity of the thermogenic bacteria of Cohn (aerobic),
which set up fermentation and putrefaction. The horticulturist uses

manure, especially horse manure, in the construction of hot l)eds for

the cultivation and forcing of young plants. In silos, the highest
temperature recorded during the fermentation of the ensiled material
was 7o°C. but the best silage is secured by keeping the temperature
below 5o°C. Sometimes spontaneous heating increases to the point
this
of actual ignition (spontaneous combustion) and it may occasionally
happen that such substances, as baled cotton, may be set on fire in this
way, for Cohn found in damp cotton waste a Micrococcus which, when
furnished with a plentiful supply of air, raised the temperature of the
decaying mass to 67°C.
Aerobic and Anaerobic Organisms. —Another useful division of
bacteria is into those which are aerobic, requiring oxygen for their
growth, and anaerobic, those which are indifferent to the presence of
oxygen. The process of respiration in the aerobes is the same as in
all Contrasted with the obligatory aerobes, we
ordinary organisms.
have those which thrive only in the absence of oxygen (obligatory
anaerobes). The growth
some of the latter is inhibited by small
of
traces of oxygen (Bacillus and some butyric organisms). One
tetani
of the classic experiments in biology was devised by Engelmann
(Botanische Zeitung, 1881 and 1882) to detect minute traces of free
oxygen. It is a well-known fact that in the process of photosynthesis,
or carbon fixation, by green plants that free oxygen is formed. Experi-
ments have shown that not all the rays of the spectrum are equally
effective in causing this chemic change. The red rays between Fraun-
hofer's lines B and C are most effective and after them those just
beyond the F line. It is these rays that are most active in the evolution
of oxygen. Engelmann reasoned, that if a green alga was placed under
the microscope and illuminated from below by a spectrum, so that the
algal filament paralleled the band of spectrum colors, that if aerobic
organisms were introduced into water beneath the cover glass, these
aerobic organisms would congregate in greatest numbers along the
green alga at those points illuminated by the rays most effective in
oxygen evolution by the plant. His anticipations were realized for
he found a grouping of the aerobic bacteria in the neighborhood of the
B and C Fraunhofer lines and beyond the F line, where theory told him
to expect the greatest photosynthetic activity. Such minute quan-
tities of oxygen must be formed by a filamentous green alga, that this
experiment becomes a microchemic test for the gas.
CHAPTER IV
CLASSIFICATION OF BACTERIA
Classification According to Nutrition. —An illuminating classification

of bacteria has been based on their mode
where three biologic
of life,
groups may be recognized: the prototrophic, the metatrophic and the

paratrophic bacteria. The prototrophic bacteria, which include the
nitrifying bacteria, bacteria of root nodules, sulphur and iron bacteria
and erythrobacteria, are those which either require no organic com-
pounds for their nutrition, or which given a small amount of organic
carbon can derive all of their nitrogen from the atmosphere, or which
with a minimum of organic matter can derive energy by breaking up
inorganic bodies.
The sulphur bacteria live in sulphur springs where hydrogen sul-
phide (HoS) isformed by putrefaction of dead animals and plants.
The sulphur bacteria in such places form a white furry growth on the
rotting vegetation. Here the H2S is attacked and water and sulphur
are formed, H2S +O= H2O -\- S. The sulphur is deposited in the
amorphous granules, which impart
living cells of the bacteria as yellow
to the organism a yellow color. To explain the facts observed, we need
assume only that the protoplasm increases the oxidizing power of the
atmospheric oxygen and renders it active. The conversion of H2S
into water and S gives 71 calories and the further oxidation of the freed
sulphur into sulphuric acid 2109 calories. The fact that the sulphur
bacteria can live without organic compounds together with their inability
to live without sulphur indicates that it is the oxidation of the sulphur
alone which takes the place of respiration in other organisms.
The ferrobacteria live in stagnant pools in marshy places. On
such pools of water, we find a greasy scum of ferric hydroxide Fe(0H)3
together with organic matter and some phosphate of iron. The ferric
compounds are reduced by the action of reducing substances formed by

putrefaction to the ferrous state which are dissolved by carbon dioxide
CO2 and unite also with it to form ferrous carbonate. The atmospheric
oxygen can convert this carbonate back to ferric hydroxide, but Wino-
CLASSIFICATION OF BACTERIA 29
gradsky has shown that the process is assisted by the iron bacteria
and the ferric hydroxide is deposited as a tube about such organisms
as Leptothrix ochracea. These tubes, or sheaths, are deposited later
as bog iron ore.
The nitrifying bacteria are found in the soils of our gardens, fields
and meadows and in virgin soil derived from places the world over.
Winogradsky has discovered that the conversion of ammonia into
nitric acid takes place in two steps and that bacteria are effective in
both of these operations. One set of bacteria belonging to the genera

Nitrosococcus and Nitrosomonas oxidize the
ammonia to nitrous acid, or its nitrite, and the
conversion of this nitrous acid (nitrite) to nitric
acid, or its nitrate, is accomplished by Nitro-
bader. Nitrosococcus is a non-motile spheric
cell, 3^t in diameter, soil from South
found in
America and Australia, while Nitrosomonas
europcea found in all soils from Europe, Africa
and Japan is a short ellipsoidal motile iorm 0.9
to iju wide and 1.2 to i.8/x long with a short
cilium. Nitrosomonas javanensis from Java is
almost spheric, 0.5 to 0.6/^, with a cilium 30/x
long, which is the longest known among bac-

teria. Nitrobacter are minute non-motile rods
/
(o.5M
.
X , \ rr.1
These organisms are ofr ii.
•
the i,ean. Glycine his pida, with

Fig. 8. — Roots of soy
0.25M).
greatest importance in putting the nitrogen of tubercles. (After Conn,
,1 •!•- r !•!
the sou mto a form which can be absorbed by p
Agricultural
g^
uuuji
Bacteriology,
)
the roots of the cultivated plants.

The bacteria which produce the nodules (Fig. 8) on the roots of
leguminous plants are probably the same the world over and to them
Beyerinck has given the name of Bacillus radicicola, while Frank called
them Rhizobium leguminosarum (Fig. 10). When the seeds of clover, or
some other leguminous species are planted, and soon after the primary
root appears with its root hairs. Bacillus radicicola, attracted chemo-
tactically to the fine root hairs, penetrates the walls of these root hairs
by ferment action. So many bacilli enter the root hair cells that they
form slimy cords, almost hyphae-like, as they move into the middle
cortex cells of the root. Here in the cortex cells, the microorganisms
form nests or pockets, that are filled with the nodule-producing bacteria
30 MYCOLOGY
(Fig. 9). The presence of these bacteria causes the formation of swell-
ings, tubercles, or nodules on the roots of the leguminous plants. Here
Bacillus radicicola remains, utilizing free atmospheric nitrogen until
about the time of flowering of the host, when it begins to assume in-
volution forms, enlarging considerably and assuming S-shaped or
Y-shaped forms (Fig. i o) . Then they are gradually absorbed by the
""*>§»**'
Fig. 9. — Cells of root tubercle of Lupinus angustifolius magnified to show the

bacteria; four cells with nuclei. {After Moore, Geo. T., Yearbook U. S. Dept. Agric,
1902, 'pi. xxxix.)
green leguminous plants and their substance is transformed into a

form of nitrogenous substance, which is utilized by the leguminous
host, either as food, or stored as nitrogenous reserve supplies. The
nodule becomes emptied of its contents and remains as a hollow sac,
enough of the organisms being returned to the soil to seed it and provide
for infection of other leguminous crops that may follow. The growth
1
of these useful organisms in the soil is stimulated by aeration, by some

organic material, by proper soil drainage, by the application of lime
which overcomes soil The farmer becomes independent of

acidity.
the ordinary nitrogenous fertilizers, which are expensive, by plowing
under the leguminous crops, which on decay yield up to the soil the
nitrogenous substance largely accumulated by bacterial action where
it is available to that large class of nitrogen-consuming plants such as
the grasses, weeds, root crops, fruit crops and the like, which are de-
pendent on the soil nitrates for their nitrogen. The leguminous plants
as nitrogen-storing plants should, in an up-to-date rotation, be
alternated with the nitrogen-consuming crops.
C' '"b-^y
Fig. io. —Left, branching forms of bacteria from clover tubercle (X2000);
right, rod forms from fenugreek tubercle ( X 2000). {After Moore, Geo. T., Yearbook
U. S. Dept. Agric, 1902, pi. xxxix.)
Metatrophic Bacteria. —The metatrophic bacteria include the zymo-

genic, saprogenic and saprophile bacteria, which cannot live unless
they have organic substances at their disposal, both nitrogenous and
carbonaceous. They where organic substances and foodstuffs
flourish
are exposed to decay in impure water and in the waste from animal
bodies. Many of them produce profound fermentative changes
(zymogenic bacteria) in bodies. Others cause putrefaction and decay
(saprogenic bacteria), while others develop in media which have been
decomposed by saprogenic species and as saprophile organisms break
these substances up into simpler chemical form.
32 MYCOLOGY
P'ermentation is well exemplified in an old and well-known process,

the conversion of alcohol into acetic acid by a number of organisms
morphologically very similar. Hansen considers that there are three
different species concerned in the acetic fermentation, namely, Bacterium
aceticum, B. Pasteurianus and B. KiUzingianus, which are non-motile,
medium-sized rods often in chains and forming pellicles which appear
on the surface of the liquid, afterward sinking to form in the liquid a

deposit known as mother of vinegar. The changes which take place
in the conversion of alcohol to acetic acid may be expressed as follows:
CH3.CH2.OH +O= CH3.CHO -f H2O

Alcohol Aldehyde
CH3.CHO +O = CH3.COOH
[Aldehyde Acetic Acid
This is conducted in barrels with wood shavings, where the alcoholic

fluid trickling over the shavings coated with the bacteria, and in contact
with the changed to acetic acid.
air, is
Lactic acid fermentation is important to man, because upon the
changes in milk by the lactic acid organisms depends the manufacture

of a considerable number of valuable products of the dairy, such as
buttermilk and cheese. This fermentation is an aerobic process whose
optimum is found between 30° and 3S°C. There is a considerable
number of bacteria capable of converting milk sugar into lactic acid,
such as Vibrio cholera, Bacillus prodigiosus and others, but the true lactic
acid bacteria are those which are the cause of the souring of milk.
Formerly, they were all classed as Bacterium acidi lactici, but recent
investigations have shown that not one species but a considerable
number are at work, sometimes one form; sometimes another being
active. A common kind is a short non-motile rod, o.^^xX. i to 2/^,
facultatively anaerobic, known by such names as Bacterium acidi lactici,

B. aerogenes, and probably comprising several races of one species.
The true lactic acid fermentation is the change of lactose, or milk
sugar, into lactic acid. As lactose is not directly fermentable it must
"be converted into such simple sugars as glucose and galactose. The
following equation approximately represents the chemic change
involved.
C12H02O11 + H2O = CeHi.Ofi -f CeHisOe
Lactose Water Glucose Galactose
C6Hi20r. = 2C3Hfi03
Lactic Acid
CLASSIFICATION (W BACTERIA ^t^
Several other important fermentations are due to bacteria, as the

causal organisms, namely, the butyric, cellulose, and mucilaginous
fermentations. The retting of vegetable fibers, the manufacture of
indigo, the curing of tobacco are all dependent on bacterial fermentations.
The saprogenic organisms are concerned with decay, or putrefac-
tion. The decomposition of dead animal and plant bodies is far from
being a simple putrefactive process. Nitrogenous and non-nitrogenous
bodies are both concerned in the putrefactive changes and they are
broken down into simpler nitrogenous and non-nitrogenous compounds,
or even elements. Proteins are spUt up into albumoses and peptones,
aromatic compounds (indol and- skatol), amino compounds (leucin,
tyrosin, glycocol), fatty and aromatic acids and inorganic end products
(nitrogen, ammonia, hydrogen, methane, carbon dioxide and hydrogen
sulphide). Ptomaines and,'other poisonous bodies are formed known
as toxins, a name applied indiscriminately to all bacterial poisons.^
The activity of all these organisms in causing decomposition of
animal and plant products is important in preserving the circulation
of carbon and nitrogen in nature. Without such destructive changes,
the elements carbon and nitrogen would be combined in such a form
and plants. In the dissolution of these
as to be forever lost to animals,
complex bodies, the simpler chemic compounds are released and can
be used over again by living animals and plants. Much should be
made of the circulation of the elements in nature and the two chief
cycles are the carbon cycle and the nitrogen cycle with a sulphur and
phosphorus cycle as There are two main processes in organic
well.
life: the constructive (anabolism), and the destructive
processes
processes (katabolism) Construction is accomplished mainly by green
.
plants and the prototrophic bacteria. Destruction is the work of

animals, metatrophic and paratrophic organisms; which have to break
down organic matter to Uve. Thus the elements of the organic world
are kept in perpetual circulation.
Paratrophic Bacteria. —-These organisms occur only in the tissues
and vessels of living organisms and are, therefore, true parasites. Many
of them are responsible for animal and plant diseases and the special
types, as far, as they concern this book, namely, those which induce
^Consult Lathrop, Elbert C: The Organic Nitrogen Compounds of Soils and

Fertilizers. Journ. Franklin Inst. 1S3 : 169-206, Feb.; 303-321, Mch.; 465-
498, Apr., 191 7.
34 MYCOLOGY
diseases in plants will be considered at length in another section.

Most attention has been paid to diseases of animals and man due to
bacteria and the number works dealing with the subjects of
of special
bacteriology, pathology, immunity and disease would form a library.
Nearly every phase of the relationship of bacteria to animals and man
has been cultivated, and microbiology has been placed on a firm founda-
tion, as a subject of human inquiry. The field is too vast for one man
to cultivate it, and hence, we find a narrow specialism perhaps more

than in any of the other departments of biologic investigation.
An interesting phase of the relationship of parasite and host has
come recently into the scientific limelight. Dr. Erwin F. Smith in
the study of the organism which produces the crown gall of woody
plants,Pseudomonas tumefaciens (Fig. 143), finds that the growth and
formation of the tumors suggests the development of cancer in man.
He thinks the formation of tumors in plants away from the point of
infection suggests a similarity (Fig. 158).
SYSTEMATIC ACCOUNT OF THE BACTERIA

For the use of students who may not have access to larger works
on bacteria and who would like a short systematic account of the
bacteria the following synopsis is given.
ORDER I. EUBACTERIALES.— The organisms of this order
are unicellular, or in plate-like, spheric, or filamentous coenobia, if
imbedded in a slimy matrix, then not of a definite form.

—
Family i. Coccace^. Single spheric cells. Division in one,
two or three directions.
Streptococcus.- —Division always in one direction, coenobia, there-
fore, chain-like, cells without flagella. Pathogenic: 6'. erysipelatos,
specific germ of erysipelas to be distinguished with difficulty from S.
pyogenes. Not pathogenic: S. mesenterioides {Leuconostoc mesen-
terioides), occurring in mucilaginous masses in the molasses waste of
sugar factories, and its presence disastrous to the industry.
Micrococcus. —Division in two directions, coenobia, sheet-like,
without flagella. Pathogenic: Micrococcus pyogenes aureus (
=
Staphylococcus pyogenes aureus), the cause of pus formation and
purulent discharge from wounds, M. gonorrhxce (= Gonococcus gonor-
rhcece) specific germ of gonorrhoea. Not pathogenic: M. aurantiacus,
luteus, cinnabareus producing pigments.
Sarcina. —Division in three planes, coenobia in bales, or pockets,

no flagella. S. ventriculi, frequent in the stomach of men, but non-
pathogenic. S. aiirantiaca, flava, luiea are chromogenic. .S'. rosea
with red cell contents occurs in swamps, or colors the soil a rose-red
color.
Planococcus.— Division and coenobic formation as Micrococcus, in
flagellate. P. citreus produces a yellow color.
Planosarcina. — Division and coenobic formation as Sarcina, in
flagellate.
Family Bacteriace.e. —
2. longer or shorter
Cells cylindric,
straight, or at least never spirally twisted. Division always at right
angles to the long axis, and only after a preliminary elongation of the
cell. The rods may separate early in some species, in others they
remain united for a considerable time as longer or shorter filaments.
Endospores are frequent, rare, or wanting. Flagella may or may not
be present.
Bacterium (Ehrenberg char, emend.). — Cells as longer or shorter
cylindric rods, often forming filaments of considerable length.With-
out Endospore formation in many species, absent in others.
flagella.
Erwin F. Smith ("Bacteria in Relation to Plant Diseases": 168 to

171) believes that bacteriologists should substitute Bacterium for
Pseudomonas as the older generic name, and he would establish a new
generic name Aplanobacter for the non-motile forms generally referred
to Bacterium. This distinction is not adopted in this text-book.
Pathogenic: Bacterium (Aplanobacter) Rathayi the cause of Rathay's
disease of the orchard grass; B. michiganense the cause of the Grand
Rapids (Mich.) tomato disease; B. anthracis the first organism deter-
mined to be the cause of disease, causing anthrax or splenic fever;
B. mallei specific in glanders in men and horses; 5. pneumonice, the cause
of pneumonia; B. tuberculosis responsible for tuberculosis (consumption,
phthisis) in man and animals. It can be distinguished by its staining
reactions. If stained with carbol fuchsin and then treated with dilute
nitric acid (1:5), the stain remains fast, while with other organisms,
washed out.
the stain will be After this treatment the tissues can be
treated with methylene blue for differential staining. B. leprcB, the
organism of leprosy; B. influenza, the cause of influenza, or grippe; B.
diptheritidis, the causal bacterium of diphtheria; B. pestis, specific in
the disease known as the plague, which as the Black Death devastated
36 MYCOLOGY
London in 1665 in which 70,000 persons perished. It is carried by

infested rats.
N on- pathogenic: B. acelicum sets up in alcohoHc sokition the acetic
acid fermentation and its films later form mother of vinegar. B.
acidi lactici ferments sweet milk transforming it into sour milk where
the acidity is due to lactic acid. B. phosphoreiim is a phosphorescent
fresh- water organism.
Bacillus (Cohn char, emend.). — Cells straight, rod-shaped to ovoid,
long sometimes united into filaments. Motile by wavy,
or short,
bent flagella scattered over the whole surface of the cell. Formation
of endospores frequent. Motility may be active for a time, and then
is lost. Pathogenic: B. muscB causes the Trinidad banana disease;
B. tracheiphilus is responsible for the wilt of cucurbitaceous plants;
B. amylovorus, the pear-blight organism; B. carotovorus, specific in
soft rot of carrot; B. aroidew, an organism which causes soft rot of the
calla; B. tetani, the causal microbe in tetanus, or lockjaw, is found in
the soil and may enter the skin or superficial muscles of man through a
pin prick, or rusty nail point; B. typhi, the typhoid bacillus. Non-
pathogenic: Bacillus subtilis, the hay bacillus found in hay and
infusion,
is the cause of decay. B. coli in the alimentary canal of animals and
men and in the water polluted by sewage. B. butyricus produces
butyric acid fermentation and the coagulation of casein. B. radicicola
(= Rhizobium leguminosarum) lives in the roots of leguminous plants
and forms the root tubercles or nodules (Figs. 8, 9, 10). B. amylobacter
(= Clostridium butyricum) ferments cellulose, dissolves casein and is
useful in the retting of plants for fiber production. B. prodigiosus
is found on many food substances imparting to them a dark red color.
B. calfactor appears in hay infusions, where it produces a rise of tem-
perature. B. putrificus, a widely distributed organism. Many bacilli
that occur in the ocean are luminous.

Pseudomonas. —Cylindric bacteria, sometimes long, sometimes short,
occasionally in threads. Locomotion accomplished by polar flagella,
the number of which may vary from one to ten, most frequently
one flagellum is present, or three to six. Endospores are formed, but
are rare. The following are the causes of diseases in cultivated plants:
Pseudomonas campestris is responsible for the black rot of cabbage and
other cruciferous plants. Ps. hyacinthi causes the yellow disease of
hyacinths. Ps. vascularum is associated as the causal bacterium in
.
Cobb's disease of sugar cane. Ps. pyocyanea causes blue pus. Ps.
putida occurs in water, where it develops a green fluorescent pigment.
Ps. syncyanea produces in milk a blue coloring matter (blue milk).
Ps. etiropcea belongs to the group of organisms which cause nitrification.
Family 3. Spirillace^.^ — Spirally wound or bent cells with occa-
sional endospore formation, usually motile. Cell division transverse
to the long axis of the cell.
Spirosoma. —-Spirally bent, rigid cells usually rather large and with-
out flagella. Unicellular free or enveloped in a gelatinous capsule.
Only a few species are known.
Microspira. —-Comma-shaped, or sausage-shaped, single, or united
cells, motile by means of a single, wavy, polar flagellum (rarely two or
three flagella), rarely longer tha*n the cell. Endospores unknown.
Usually united with the next genus.
Spirillum. — Rigid rod-shaped cells of varying thicknesses, lengths
and pitch of spiral turns, hence, either as long screws, or loosely wound.
Flagella occur at one or both ends of the cells as polar tufts varying in
number from five to twenty. In some species, endospore formation
has been observed. Sp. comma is the cause of asiatic cholera and is
found in cultures often in long spirally wound filaments. There are

nmny non-pathogenic spirilla in water from rivers and ponds as S.
danubicum in the Danube, Sp. berolinense in Spree water, Sp. ruftim
in stagnant water. Sp. rufum forms blood-red slimy masses between
decaying algae.
SpirochcBta. —Thin, flexible, snake-like, motile cells usually quite

long without observed flagella and endospores, and unsegmented.
Spirochceta Obermeieri is the cause of relapsing fever (f ebris recurrans)
S. {Treponema) pallida is the organism of syphilis. S. dentium is found
associated with the teeth in man.
Family 4. Phycobacteriace^ (Chlamydobacteriace^). Cylin- —
dric cells united into sheath-surrounded threads and reproducing by
motile or non-motile conidia, which arise from the vegetative cells
without a resting stage.

Streptothrix (== Chlamydothrix, Leptothrix, Gallionella).
Non-motile —
unbranched threads possessing a sheath of varying
cylindric cells in
thickness. Septa vague. Reproduction is accomplished by roundish,
non-motile conidia arising from the vegetative cells. S. fluitans in
water.
38 MYCOLOGY
Crenothrix. —The cells are arranged in unbranched threads attached

at one end and enlarging toward the distal extremity. Filaments
covered by a rather thick sheath. The reproductive cells are non-
motile conidia, which on discharge immediately germinate. Crenothrix
polyspora in springs and water pipes, where it forms attached slimy
growths. The sheaths in iron waters are impregnated with iron
oxidhydrate.
Phragniidiothrix. — Cylindric cells with delicate, scarcely visible
sheath. The cells of the filament are at first in one plane which later
divide in three directions to form clumps or packets of cells. Later

the single cells round off and become free. Ph. nrnltiseptata with fila-
ments 3 to 12/X broad and looyu long attached to the bodies of crustaceae.
Cladothrix {SphceroHlus in part). —The
and often tufted
fixed
filaments form delicate sheaths. The and by inter-

cells are cylindric
calary growth may break laterally through the sheath to form false
dichotomous branches. Reproduction is accomplished by motile
swarm spores (gonidia) which bear a tuft of flagella a little to one side
of a pole. Cladothrix dichotoma occurs frequently in stagnant water,
attached and forming furry growths. The following species occur in
the soil: C. rufula, C. profundus, C. intestinalis, C. fungiformis, while
C. intrica has been isolated from sea water and sea mud.
Family Thiobacteriace^ (Beggiatoace.e). Cells with sul-
5. —
phur inclusions, unpigmented, or colored rose, red or violet by bacterio-
purpurin; never green. The plants are generally filamentous with
division transverse to the long axis.
Thiothrix. —Unequally thick attached filaments encased in a
delicate, scarcely visible sheath. Rod-shaped conidia are formed at
the ends of the threads. Th. nivea is found in sulphur springs and in
stagnant water.
Beggiatoa. — Sheathless, free-filamentous bacteria, motile by means
of an undulating membrane. Cells with included sulphur granules.
Spore formation unknown. B. alba is found in dirty water, drain
water from sugar factories and attached to decayed plants in sulphur
springs. B. mirabilis forms white growths on dead marine algae.
The colored sulphur bacteria, sometimes placed in the family
Rhodobacteriace^, belong here. They have rose, red or violet cell
contents due to the presence of bacteriopurpurin (see ante). The im-
portaut genera according to Erwin F. Smith (''Bacteria in Relation to

Plant Diseases," I: 163) are Thiocystis, Thiocapsa, Thiosarcina,
Lamprocystis, Thiopedia, Amosbobader, Thiothece, Thiodictyon, Thiopoly-
coccus, as well, as the three genera Chromatium, Rhabdochromatium,
Thiospir ilium.
Family 6. Actinomycetace.'E (Position doubtful). —Radially ar-
ranged branched filaments in colonies, non-motile. Filaments divid-
ing into oidia-like reproductive cells.
Actinomyces chromogenes occurs in A. bovis is the cause of

soil.
lumpjaw in cattle man. The plant occurs in

and occasionally in
rosettes usually 30 to 40/i in diameter. The filaments which are often
curved sometimes spirally exhibit true branching and are interlaced
in a network. Recently Youngken (Amer. Jour. Pharm., September,
1 91 5) has described the foundation of the large swellings
(mycodomatia)
on the roots of the waxberry, Myrica carolinensis, and other species, as
due to a species of ray fungus, Actinomyces myricarum, that abun-
dantly fills infested cells in the cortex of the tubercular swellings. A.
thermophilus is found on hay and manure.
ORDER II. MYXOBACTERIALES.— Individual plants en-
closed in slimy masses which assume more or less regular fructifica-
tion-Hke shapes.
—
Family i. Myxobacteriace^.^ Erwin Baur and Roland Thaxter
have studied these forms most intimately. The plants of this family
consist of motile, rod-like microorganisms, with a gelatinous base and
forming false plasmodioid aggregations preceding a cyst-producing,
quiescent state in which the rods may be encysted in groups or con-
verted into spore-masses. The slightly reddish rods in the vegetative
stage are elongate, sometimes 15// long and vary httle in size in the
different genera and species. is by fission and the active
Cell division
rods show a slow sliding movement without organs of locomotion. The
vegetative phase in artificial cultures usually lasts about a week, or
even two weeks, and the formation of cysts which follows must be more
rapid in nature. These organisms are found in moist places on decay-
ing wood, dung, funguses and lichens, growing best, according to Baur,
at 3o°C. Three genera are included in this family.
—
Chondromyces. Rods producing free cysts within which they
remain unchanged. The cysts are various, sessile or developed on a
stalk (cystophore).
40 MYCOLOGY
Folvangiiim {= Myxobacter, Cyslobacter). — The rods form large

rounded cysts one or more of which are free inside a gelatinous stalked
matrix.
Myxococcus. — Slender rods which swarm together, after a vegetative
phase, to form well-defined, more or less sessile or stalked encysted

masses of coccus-like spores.
BIBLIOGRAPHY
Abbott, A. Principles of Bacteriology, 9th Edition: Lea & Febiger, 1915.

C: The
DE Bary, Comparative Morphology and Biology of the Fungi, Mycetozoa and
a.:
Bacteria. Oxford at the Clarendon Press, 1887.
Baur, Erwin: Myxobakterien Studien. Archiv fur Protistenkunde, v Bd., Heft I,
92-121, 1904.
Buchanan, Estelle D. and Robert Earle: Household Bacteriology. The
Macmillan Co., New York, 1914.
Chester, Frederick D.: A Manual of Determinative Bacteriology. The Mac-
millan Co., 1914.
Conn, H. W.: Bacteria, Yeasts, and Molds in the Home. Ginn & Co., Boston, 1903.
DuGGAR, Benjamin M.: Fungous Diseases of Plants. Ginn & Co., Boston, 1909.
Ellis, David: Outhnes of Bacteriology (Technical and Agricultural), Longmans,
Green & Co., 1909.
Engler, Adolf and Gilg, Ernest: Syllabus der Pflanzenfamilien. Berlin, 191 2,
Siebente Auflage, pp. 1-5.

Eyre, J. W. H.: The Elements of Bacteriological Technique. Philadelphia, W.
B. Saunders & Co., 1902.
Fischer, Alfred, transl. by Jones, A. Coppen: The Structure and Functions of
Bacteria. Oxford at the Clarendon Press, 1900.
Fuhrmann, Dr. Franz: Vorlesungen iiber technische Mykologie. Jena, Gustav
Fischer, 19 13.
Hiss, Philip H. and Zinsser, Hans: A Text-book of Bacteriology. D. Apple ton
& Co., 1915.
Jordan, Edwin O.: A Text-book of General Bacteriology, 3d Edition. Phila-
delphia, W. B. Saunders & Co., 191 3.
KisSKALT, K.: Bakteriologie Zweite Auilage. Erster Teil von Prakticum der
Bakteriologie und Protozoologie. Jena, Gustav Fischer, 1909.
KtJSTER, Dr. Ernst: Anleitung zur Kultur der Mikroorganismen. Zweite Auflage,
Leipzig und Berlin, 191 3.
Lafar, Dr. Franz: Technical Mycology. The Utilization of Microorganisms in
the Arts and Manufactures. London, Charles Griffin & Co., vol. i, 1898.
Lipman, Jacob G.: Bacteria in Relation to Country Life. The Macmillan Co.,
New York, 1908.
Marshall, Charles E. and Others: Microbiology for Agricultural and Domestic
Science Students. Philadelphia, P. Blakiston's Son & Co., 1911.
Meyer, Dr. Arthur: Practicum der botanischen Bakterienkunde. Jena, Gustav

Fischer, 1903.
MuiR, Robert and Ritchie, James: Manual of Bacteriology. The Macmillaii
Co., 1913.
Newman, George: Bacteria. Especially As They Are Related to the Economy
of Nature to Industrial Processes and to Public Health. G. P. Putnam's Sons,
New York, 1899.
Park, William H. and Williams, Anna W.: Pathogenic Microorganisms.
Lea & Febiger, Philadelphia, 1914.
Perchâl, John: Agricultural Bacteriology Theoretical and Practical. Duck-
worth & Co., London, 1910.
Prescott, Samuel C. and Winslow, Charles Edward A.: Elements of Water
Bacteriology, 3d Edition. John Wiley & Sons, New York, 1913.
QuEHL, A.: Untersuchung liber Myxobakterien. Zentralblatt fur Bakteriologie,
II Abt., xvi Bd., 1906.
Smith, Erw7N F.: Bacteria in Relation to Plant Diseases, vol.
i, 1905; vol. ii, 1911;
vol. iii, Publication No. 27, Carnegie Institution of Washington.

1914.
Th.^xter, Roland: On the Myxobacteriaceae, a New Order of Schizomycetes.
Bot. Gaz., xvii, 1892; Further Observations on the Myxobacteriacese. Loc.
cit., xxiii, 1897; Notes on the Myxobacteriaceae. Loc. cit., xxxvii, 1904.
Wettstein, Richard R. von: Handbuch der Systematischen Botanik Zweite
Auflage, 191 1 : 69-83.
CHAPTER V
CHARACTERISTICS OF THE TRUE FUNGI
CLASS III. EUMYCETES
The hyphomycetes {v(t)ri, a web + hvktjs, a mushroom)
true fungi or
are thallophytes in which the thallus, as the Greek derivation implies,
consists of a system of threads {hypha) which form a cobwebby struc-
ture known as the mycelium (Fig. 1 1). A single thread of the mycelium
is an hypha (plural hyphae) and a hypha may be unicellular, or multi-
cellular. All true fungi are colorless, that is they are chlorophylless;
and although they may have other pigments present, yet in the absence of
chlorophyll, they are dependent plants.
As dependent plants, they must get
their organic food from extraneous
sources, and as all organic matter is
either dead, or living, a natural classi-

fication of fungi into saprophytes and
parasites can be made. A saprophyte
Fig. II. —
Gray mould, Miicoy,
{aairpos, rotten
.
+ j
4>^t6v, a plant)
•.
is any
j
i •
i
•
i
•
r r
showing mycelium and the sporan- Organism which dcrives its chief food
gia on upright sporangiophores. supply from dead, or dead and decaying
^^.^^^ ^^ plant organic material, while
a parasite {irapaaiTos, one who lives at another's expense) is an
organism, which exists at the expense of living animals, or plants
(Fig. 12). But some saprophytes may change their mode of nutri-
tion and become parasitic; such saprophytes are called facultative
parasites, while those which retain their saprophytism under all condi-
tions are obligate saprophytes. Again some parasites can adjust their
methods of nutrition, so that they can become saprophytes. Such
parasites are called facultative saprophytes, while those organisms
which are always parasitic are obligate parasites. These distinctions
are useful, but it should be emphasized that there is no absolute border-
line between one condition and the other. There are imperceptible
42
—
CHARACTERISTICS 07 THE TRUE FUNGI 43
gradations which preclude an absolute pronouncement as to whether

a plant is a saprophyte, or a parasite.^ Botanists generally concede
that the true fungi have been derived from filamentous algal ancestors
and the groups of algae from which the principal forms of fungi have
Fig. 12. Russula nigricans parasitized by Nyctalis aslerophora. {After Brefeld.)
been derived are fairly well known. For example, it is believed that
such fungi as belong to the order OOMYCETALES have been derived
1 Massee, George: On the Origin of Parasitism in Fungi. Annals of Botany,
xviii: 319.
Ward, H. M.: Recent Researches on the Parasitism of Fungi. Annals of Bot-

any, xix: I.
Bancroft, C. K.: Researches on the Life History of Parasitic Fungi. Annals

of Botany, xxiv: 359, 1910.
44 MYCOLOGY
from a green alga like Vaucheria. With our present knowledge, it is
impossible toname any one existing alga as the progenitor of a definite

fungous form, but we are safe in assuming in a general way that certain
phyla of fungi have been derived from certain phyla of algae,by the
loss of chlorophyll and in the loss of an independent existence. Another
view, which is open to argument, is that certain of the prototrophic
—
Fig. 13. Development of Mucor miicedo. a, b, c, d, Stages in the formation of
zygospore; /, sporangium; g, mature sporangiospores; h, one germinating. {After
Schneider, Pharmaceutical Bacteriology, p. 142.)
filamentous bacteria to which attention has been previously called

have been the direct progenitors of certain of the filamentous fungi,
but on account of the character of the reproductive organs in the lower
true fungi their derivation from green algae is the more probable, and
mycologists even speak of the algal fungi referring especially to aquatic
genera, such as Saprolegnia, which like their algal ancestors not only
retain the general morphologic features of the algae, but also live in an
CHARACTERISTICS OF THE TRUE FUNGI 45
aquatic medium, and the success of the process of fertilization depends

on the presence of free water. Such fungi form a subclass of EUMY-
CETES, the PHYCOMYCETES.
The vegetative organs of fungi are concerned with the absorption
of food, the assimilation of the food and in the nutrition of the organs
of fructification which together form the reproductive system. That
the student may appreciate the morphology of the vegetative organs of
the fungi, three examples from widely divergent orders will be chosen
by way of illustration. A common mould is Mucor mucedo which
appears on horse manure. If a spore of this fungus is placed in a nutri-
tive medium, its wall breaks and there protrudes a germ tube rich in
protoplasmic contents (Fig. 13, h). This germ tube grows in length
into an hypha without the development of partition walls dividing it
into shorter cells. This hypha branches and rebranches in its growth
over the nutrient substratum spreading in all directions, if unimpeded
by other organisms growing on the same food substance. The ultimate
branches of this mycelium, which is throughout unicellular, are much
attenuated, fine hyphse representing the end ramifications of larger and
coarser hyphae nearer the point of origin of the whole mycelium (Fig.
13). The finest hyphae usually enter the substratum, while the coarser,
stronger hyphae form a cobwebby mass over its surface. We can
distinguish therefore the feeding hyphae, which are rhizoidal hyphae,
and the aerial hyphae in which probably the metabolic changes are
most active where the mycelium is in open contact with the air. Later,
when the mycelium is well established on the nutrient substratum,
erect vertical hyphae appear at indefinite points on the larger aerial
hyphae. These are the fruiting hyphae, or sporangiophores, which
ultimately cut off a terminal cell which becomes the sporangium, or case,
in which the reproductive cells or spores are formed, while the end of
the sporangiophore projects into the interior of the sporangium as a
columella (Fig. 13, /).
The common green mould, Penicillium glaucum, may be taken as the
second illustration (Fig. 14). If we sow a spore on nutrient agar in a
Petri dish after a few hours the spore swells and there emerges a germ
tube which at first is undivided by a partition wall. Later, as the older
hyphae branch to form new ramifications, cross-partitions are formed
which divide the mycelium into short cells, so that in that respect the
mycelium of PeniciUhim differs from that of Mucor. The hyphal
46 MYCOLOGY
branches are coarser in Penicillium and do not form the fine-pointed

ends found in Mucor. The presence of transverse walls in the fungi is
thought of sufficient importance to make a subclass known as the

MYCOMYCETES to contain all of the true fungi EUMYCETES
which have a mycelium which is multicellular in contradistinction
to those which have unicellular mycelia and that form the subclass
PHYCOMYCETES. From this spreading myceHum of transversely
septated hyphae in Penicillium arise hyphae which branch at the

extremity into a number of erect branches from the ends of which
are cut off in sequence a series of small round cells, the spores, which if
undisturbed remain connected in a chain, so that the
fructification roughly resembles a small broom, or
whisk. The large vertical hypha is a conidiophore,
and as the spores are pinched, or abstricted off from
the secondary branches as single cells, they are
known as conidiospores {kovls, dust + airopa a seed)
(Fig. 14 and Figs. 243 to 263 inclusive).
The third example, which we will use to describe
in general terms the vegetative organs of the fungi,

is the honey-colored toadstool, Armillaria mellea
(Fig. 15). The toadstools, or fruit bodies, often form
Fig. 14. — Con- dense clumps around the base of some dead or dving
mo°n ""Te'en-mouS; tree, or

almost cover an old stump on which ihey
Penicillium glau- grow. The Cap is of a houcy-colored brown, about
llZnr'ôi ^coSSo- two inches across, and

the stem may be six inches
spores. (After Conn, long and paler than the cap. Microscopic sections
that are closely bound together to form the stem and

cap. If we examine the base of the stalk, we find that it arises from
a dark-colored cord-like strand which has been termed a rhizomorph
because of its resemblance to a root (Fig. 15, II and IV). These
rhizomorphs constitute the mycelium and they either ramify through
the soil, or else are found beneath the bark of the dead tree, where
they unite to form open-meshed nets of a dark brown color. These
rhizomorphs are strands of hyphae that run longitudinally. The
hyphal cells are bound together in a cord-like cable which is peculiar
in that it shows apical growth, constantly elongating at its extremity,
as it grows beneath the bark, or penetrates the soil (Fig. 15)
—
Fig. 15. Details of the mycelium of Armillaria niellea. I, Piece of mycelium
on slide; II, piece of old mycelium {Rhizomorpha sublerranea); III, rhizomorph pro-
ducing fruit bodies; IV, apex of rhizomorph capable of growth; (a) peripheral hyphs;
{b) pseudo-epidermis; (c) growing point; {d, e, h) pith; (/?) hollow center. (7 and
IV after Brefeld; III, after Hartig in Zopf, Die Pilse, 1890, p. 25.)
48 MYCOLOGY
its it grows beneath the bark, or penetrates the soil (Fig.

extremity, as
15). Such a compound thallus differs strikingly from the filamentous
thalluses of the two previously described fungi. The union of the
hyphal cells in some of these fleshy fungi may be so intimate as to con-
stitute a pseudoparenchyma, and this close union of the cells may be
made still more intimate by clamp connections where two adjoining cells
are bound together endwise by a clamp-like protuberance of one of the
cells attached to the end of the other adjoining cell. When the pseudo-
parenchyma is external, it rnay serve for the protection of the internally
disposed hyphae, and be looked upon as protective tissue. Mechanic
tissues for the support of fungi are not unknown in some of the groups,
as in some of the polypori; where there are clamp connections, trans-
verse septa and thickened cell walls. A few of the higher fleshy fungi
have conducting hyphae, which are larger and more tubular than the
surrounding hyphae, and which conduct later, oil and other substances.
Those which conduct a milky juice, as in some species of Russula and
Lactarius, may be termed laticiferous hyphs. There are some fungi in
which the hyphal form of thallus is not present. The yeasts are either
single ellipsoidal cells, or these cells are loosely connected together in
a chain of bed-like cells. These chains are due to the budding or
sprouting method of cell multiplication where a bud, gemma, or sprout,
grows out from the mother cell as a daughter cell. buds
It in turn
producing a granddaughter and so
cell forth. Such a method of
reproduction is known as gemmation.
In the parasitic fungi, the hyphae run either into the cells, through
the cells (intracellular), or between the cells (intercellular). Where
the hyphae are intercellular, short branches may be formed which
penetrate the host These short branches take various forms and
cells.
are known an haustorium (Figs. 36 and 67).

as haustoria; a single one as
Occasionally in the mildews, the mycelium may be superficial and
hence epiphytic, while the mycelia which are internal are endophytic.
These are useful terms when describing the parasitic habits of fungi.
Some of the groups of fungi have mycelia that form resting bodies
of hyphae. These are the most compact of all forms of mycelia and
—
are known as sclerotes {sclerotium ia), which in many cases assume
tuberous forms. They are resting states of the mycelia and act as
stores of reserve material. These are some of the principal forms of
the vegetative thallus of the fungi. Further details will be given in
he discussion which follows. Some sudden epidemics of rust fungi
have been ascribed by Eriksson to the presence of the protoplasm of

the rust mixed with the protophism of the host. To this included
fungous protoplasm he gave the name mycoplasm.
Some fungi are symbiotic, that is, they are found in intimate re-
lation with chlorophyll-containing plants and obtain from them food
of a carbonaceous character, but without apparently injuring the
green symbiont. When they live with algae, they commonly form
lichens; or if in connection with the roots of trees, orchids; and in
prothallia they form what is known as mycorhiza (Fig. 16).

The spores or reproductive cells of fungi may be of two kinds:
non-sexual spores and sexual spores. The non-sexual spores are cells
which are formed vegetatively. They are cells which take special
Fig. 16. — Ectotrophic mycorhizas. At left hyphal mantle on root of hickory

Carya ovata in cross section; at right root tip of an oak, Quercus, with fungous mantle.
{From Gager, after W. B. McDougall.)
forms in the different groups of fungi and are produced as special cells
in a purely vegetative manner. They represent a special part of the

thallus given over to reproduction. Upon the formation of these
spores, which may germinate at once or live for some time as resting
spores, the rapid multiplication of the fungi depends. It is the innu-
merable quantity of these non-sexual spores upon which an epidemic
of some particular fungous disease may depend. Only the most general
characters of the various kinds of spores can be discussed in an intro-
duction of this kind. The special kinds will receive due attention as
we proceed. Spores which are cut off, or pinched off, in concatenation
from the end of a vertical hypha, are known as conidios pores. In the
rusts such conidiospores become and in the mushrooms
nredospores,
basidiospores. Where the non-sexual spores are formed in a spore case,
4
50 MYCOLOGY
or sporangium, they may be termed sporangiospores (Fig. 13,/). Fre-

quently spores are formed by a modification of certain cells of the hy])lial
branch. These spores are usually thick-walled, as in the smuts, and
become known as chlamydos pores. Where the whole hypha is divided
up into a chain of spores one after the other in close order, such spores
are called oidiospores. Special receptacles are associated with the
formation of the non-sexual spores. These are found in the sac fungi,
ASCOMYCETALES, where the depressed conceptacle becomes a pycnidium,
or conidial fruit, and the spores which it contains are pycnidiospores,
pycnospores, pycnoconidia or the stylospores of Tulasne. This form of
conidial fruit is surrounded by a firm wall or peridium. The pycnidia
may be depressed in the tissues of a host plant or elevated above its
surface, as the case may

In some fungi the conidiophores, in-
be.
stead of being separate, are arranged in parallel order, side by side,
at an early stage, and thus are united into a fascicle to which the name
coremium has been applied.
The principal sexually produced spores in the fungi are zygospores,
oospores and ascospores. The first two forms are found. in the sub-
class PHYCOMYCETES.
Their formation proceeds in such a manner that the zygospores are
produced isogamously, that is, by the union of two similar cells, while
the oospores are heterogamous, that is, they are produced by a union of
an egg cell and a sperm cell. Hence, we distinguish two orders of
the PHYCOMYCETES, namely, the ZYGOMYCETALES and the
OOMYCETALES, the first showing isogamy and the latter heterogamy.
Details will be given when these orders are considered in detail. Until
recently, was believed that sexuality did not exist in the sac fungi,
it
ASCOMYCETALES, but recent research has shown that the nuclei

of two adjoining cells unite and this is followed by the formation of a
spore sac, or ascus, containing sac spores, or ascospores. The formation
of the asci is usually associated with the production of definite fruit
bodies. It is found in any of the other
doubtful whether sexuality is
groups of fungi. Curious nuclear fusions in the rusts have been sug-
gested as a sexual union, but it is safer to await future discoveries
before adopting such a position. However, there are fungi in which
and many of these belong to the
sexual organs seem to be lost entirely
most highly developed forms where the thallus and fructifications are
of a complex type. The whole trend of evolution in the fungi is for
1
the reduction in size and importance of the sexual organs, until they
have disappeared completely. This may be a result of the perfect
manner in which the dififerent specific types are reproduced and multi-
plied by the various kinds of non-sexual spores found in the different
fungous groups.
CHAPTER VI
HISTOLOGY AND CHEMISTRY OF FUNGI

Histology. —
Naked cells which are destitute of a cell wall and con-
sist of naked protoplasm occur as motile cells in only two unimportant
groups of the OOMYCETALES. The cell wall of fungi does not
appear from the results of numerous workers upon its chemistry
to be of the same nature in the different groups of them. A general
term which has been in current use and which was first suggested
by A. de Bary is that of fungous cellulose, but that term, as far
as indicating the chemic character of the membrane is concerned, is
a misnomer. if we employ the term in
It has its correct application,
the sense of fungous membrane substance. We owe to C. van Wis-
selingh (1898) the examination of about a hundred species from nearly
all of the orders and most of the families of EUMYCETES. Wissehngh
could detect the presence of cellulose with certainty only in two families,
the Saprolegniace^ and the Peronosporace^. This carbohydrate
could not be detected either in the ZYGOMYCETALES or in any
of the MYCOMYCETES examined, and especially was it found to be
absent in the yeast Saccharomyces cerevisia. The researches of
Winterstein, Gilson and Wisselingh proved that chitin formerly sup-
posed to be of animal origin was found in the membranes of fungi.
With the exception of the two families mentioned above, the bacteria
and the yeasts, chitin has been detected in all other species of fungi
examined, e.g., Mucor mucedo, M. racemosus, Rhizopus nigricans,
Penicillinm glaucum, Trichothecium roseum, in the sclerotia of Botrytis
cinerea and Claviceps purpurea. We do not know at present of the
simultaneous occurrence of cellulose and chitin in the same cell wall.
E. Winterstein has found true hemicellulose in certain fungi and other

chemic substances have been reported such as carbohydrates of the
pentosan group, pectose, callose, etc.
The outer layers of the wall, in some fungi (Tremellace^) may be

mucilaginous, so that it is resolved into a soft gelatinous mass. Lignifi-
cation has been reported in the large pileated fungi though whether
52
HISTOLOGY AND CHEMISTRY OF FUNGI 53
the presence of lignin is proved thereby must remain an open ques-

tion. Deposits and incrustations of calcium oxalate crystals are found
in themembranes of fungi, as the spicules in the sporangial wall of
Mucor mucedo.
The cell contents, or protoplasm, of fungi may be divided into
cytoplasm with its inclusions and nucleoplasm. The cells contain
either a single nucleus (Erysiphe), two, as in Exoascus, or several,
as in the mycelial cells of Penicillium glaticum and Peziza convexula.
The hyphae of many contain numerous, sometimes over hundreds
of nuclei (PHYCOMYCETES). The structure of the nucleus in
basidia as described by Wager agrees with that of the higher flowering
plants. It has a nuclear membrane, nucleolus and nuclear network of
threads coiled in a loose knot. Chromatin granules occur. The nucleus
undergoes division either by fission, or by karyokinesis, as first observed
by Sadebeck. Chromosomes are formed from the chromatin bodies
when the nucleus begins to divide. A reduction of chromosomes has
been observed by Stevens. Fats and oils are present in fungous cells
and are found in the form of drops or globules. Glycogen has been de-
tected in the spore sacs of the ASCOMYCETALES. Volutin is a
name given by Meyer to a reserve substance which contained C, H, O, N
and P atoms. Mannite, trehalose and glucose have been found in
many fungi by Bourquelot. Special substances of a poisonous nature
such as ergotin, muscarin, phalhn are of special significance in cer-
tain fungi.
Colors. — Full details regarding the coloring matters in fungi will
be found in Zopf's "Die Pilze in morphologischer, physiologischer,
biologischer und systematischer Beziehung," 1890. Clear bright
colors are present in such species as Peziza aurantia, P. coccinea.
Russula virescens, has a cap with shade of green lighter, or darker, in
individual specimens. Russula emetica is red. Blue is the predominat-
ing color in the genus Leptonia. Armillaria mellea has a honey-
brown, or yellow color. The violet color of Cortinarius violaceus is
well known. The color in a number of fleshy fungi changes when the
fruit bodies arebroken, injured or exposed to the air. This change of
color due to an oxidizing enzyme. The flesh of a number of species
is
of Boletus changes from white or yellow to a deep indigo-blue when
broken, or abraded. The deliquescence of species of the genus

Coprhius, when the color changes from white to black with the melting
54 MYCOLOGY
down of Lhe whole fruit body has been proved to be a process of auto-
digêstion. When the hyphae are colored, the color is confined generally
to the cell wall, although Biffen states that in some hyphae the color
is located in the contents, the wall remaining colorless. Spores are
colored frequently as in Ascobolus which grows on manure. The spores
at first colorless change through pale lilac to clear deep amethyst. The
coloring matter is confined to the spore walls, but in some cases the
contents are colored, while the wall is colorless, as in many a^ciospores.
Physiology or Fungi
The research of recent years in the nutrition of fungi has shown

that nine chemic elements are necessary for the structure and complete
development These elements are carbon, hydrogen,
of the true fungi.
oxygen, nitrogen, sulphur, potassium (or rubidium),
phosphorus,
magnesium and iron. Analysis of the ash. constituents of fungi shows
that phosphoric acid and potassium are the chief ones, the latter form-
ing seldom less than one-quarter and sometimes one-half of the total.
Phosphorus is present in the ash to the extent of 15 to 60 per cent,
and is eagerly absorbed by growing fungi, as is shown by Dcedalea
quercina, which in its growth completely extracted the phosphoric
acid from decayed wood. Winogradsky, Meyer, H. Molisch and W.
Benecke have shown that magnesium is indispensable to fungi. Be-
necke has demonstrated a considerable difference in development shown
by two, otherwise equal, specimens, the one grown without magnesium
and the other in a medium containing 0.0025 mg- of crystallized magnes-
ium sulphate per 25 c.c. and Guenther has proved that 0.005 "^g- of
magnesium sulphate was necessary to induce a sowing of Rhizopus
nigricans to grow at all.
As to iron, as an indispensable element before the matter was put
to the test, it was thought that fungi being chlorophylless did not
require iron like the green plants in which iron was concerned in the
formation of chlorophyll. The experiments of Hans Molisch tend to
prove the essential importance of iron in the nutrition of the true fungi
for in presumably iron-free cultures, the spores of Aspergillus niger
did not develop beyond the formation of a sickly mycelium. Similar
results were obtained with sowings of pressed yeast cells, spores of
Mucor racemosus and a species of Penicillum. Iron in addition to
HISTOLOGY AND CHEMISTKY OF FUNGI 55
being a nutritive material also acts as a stimulant. The position of

sulphur, as an important nutritive element, is doubtful. It is inferred
that because this element forms an important constituent of the albu-
minoids, that it is, but there are no re-
therefore, essential to fungi,
liable experiments which prove that to be Awaiting more detailed
so.
investigations, sulphur has been included in the above list of nutri-

tive elements. The source of the C, H, and O which form such an
important part of the food of fungi is the dead or Hving bodies of other
plants and animals, principally plants in which are found sugars,
and other bodies of organic origin.
starch, cellulose, mannite, citric acid,
The source of nitrogen is similarly from soluble nitrogenous bodies,
peptones, propylamin, asparagin and others, but few if any of the
higher fungi can utihze free atmospheric nitrogen, as can the bacteria
which form the nodules on the roots of leguminous plants, described in
a former section of this book. The various culture media on which
bacteriologists and mycologists cultivate successfully a large series of
bacteria and fungi will be considered in a subsequent chapter. Modern
research along the lines of technique has demonstrated many im-
portant points about the growth and nutrition of the higher fungi
and these will be discussed, as we proceed to the end of the book.
The chemic investigation of the fungi began with the refinements
in the technique of modern organic chemistry and much has been pub-
lished on the subject, so that there is a bibliography too voluminous to
give. Much of the most important chemic work on fungi published
prior to 1890 will be found in Zopf's "Handbook." No general work
of this kind has recently appeared, so that we must depend on recent
original papers on the chemistry of fungi, and
on the statements
in part
of Zopf's great book. The following inorganic elements have been
found in fungi: chlorine, sulphur, phosphorus, sihcon, potassium,
sodium, lithium, calcium, magnesium, aluminium, manganese and
iron. Manganese has been found in the cap of Lactarius piperatus.
Aluminium has been reported as occurring in the ash of lichens. The
mean of a number of analyses^ of mushroom {Agaricus campestris),
truffle (Tuber), Morchella esculenta, two other species of MorcheUa,
species of Boletus, a,nd Polyporus officinalis is as follows: potassium 45
per cent., phosphoric acid 40 per cent., magnesia 2 per cent., sodium
1.4 per cent., calcium 1.5 per cent., iron oxide i per cent., silicic acid
'ZoPF, Wilhelm: Die Pilze: ii8.
56 MYCOLOGY
I per cent., sul])huric acid 8 per cent., chlorine i per cent. The organic
compounds carbohydrate group found in fungi are cellulose,
of the
grape sugar, glycogen and kinds of gums, mannit, inosit, and several
other less important ones. The organic acids include oxalic, malic,
acetic, citric, formic, lactic, helvelhc, and propionic acid, as well as other
less well-known acids.
Fats and oils are often present as reserve substance in many repro-
ductive spores, as in oospores, zygospores, ascospores, and the like.
Large quantities are also often present in the mycelium, as in Lactarius

deliciosus, which contain 6 per cent. (5.86 per cent.). Fat is, as a rule,
not entirely absent from any species of fungus. Fliickiger gives the
fat content of the sclerotium of Clavkeps purpurea as 35 per cent. The
mushroom Agariciis campestris has 0,18 per cent, and Helvella esculent a
1.65 per cent.
Resin occurs in fungi in the form of excretions, partly as infiltra-
tion of the cell walls, partly as contents of the living cells. The intense
orange-yellow color of the caps and stipe of the Agaricus (Pholiota)
spectabilis, according to Zopf, as also the pale yellow of the gills and
the flesh of cap and stipe together with the ochre-yellow color of
the masses of spores is due to the presence of a resin acid which is
present as a hyphal cell content. Pigments of various kinds classified

by Zopf are also found. Besides the important substances mentioned
above, chemists have found coniferin, muscarin, trimethylamin (spores of
Tilletia caries), ergotin, cholin, phallin, cholesterin. Several of these
will be discussed in connection with the poisonous or non-poisonous
character of certain of the fleshy fungi.
Enzymes {Jev^vjjios, leavened, from €u, in and ^vp-v, leaven, a term
first suggested by Kiihne for an unorganized ferment). —The study of
the ferments, or enzymes, of the fungi and higher plants has thrown a
flood of light upon their metabolic activity, for enzyme action is the
strategic center of vital activity. Pasteur emphasized the role of micro-
organisms as ferment producers, and that led to the classification of
ferments into organized and unorganized. Since Buchner discovered
zymase, ferments have been divided into endocellular and extracellular.
Endocellular enzymes as those which cannot diffuse out of the cell,
such as zymase, while extracellular enzymes are those which are capable
of diffusion out of the cell, such as invertase. Hepburn defines an
enzyme as a soluble organic compound of biologic origin functioning
HISTOLOGY AND CHEMISTRY OF FUNGI 5 7
as a thermolabile catalyst in solution. In connection with this defini-

tion, it is important to know that a catalytic agent is one which alters
the rate of a reaction without itself entering into the final product
(Ostwald, 1902), or which does not appear to take any immediate part
in the reaction, remains unaltered at the end of the reaction and can
be recovered again from the reaction product unaltered in quantity
and quality.
Enzymes differ from ordinary inorganic catalysts in their sensitive-
ness to heat and light. They C, and most of
are destroyed at 100°
them cannot be heated above 60° C.^ The velocity of the
safely
reaction increases with a rise of temperature up to an optimum and
as the temperature is increased above the optimum the enzyme is
permanently inactivated. Enzymes retain activity even after ex-
posure to action of liquid air. Light in its ordinary form in the pres-
ence of oxygen and ultraviolet light independent of oxygen are de-
structive to enzymes. Again, enzymes possess most of the important
properties of colloidal solutions, such as their non-diffusibility. They
are soluble in water, in dilute salt solutions, or in glycerin. They
exhibit the phenomenon of adsorption.
An important discovery has recently been made which has thrown
considerable light on the activity of enzymes, and that has been the
stimulation exercised by certain substances which have been called
activators and the by other substances, which have
inhibition exercised
been called paralyzers. The activators are in some cases simple chem-
ical substances, such as acids, alkalis and salts, or they are complex
bodies of unknown chemic character, but they have this in common that
they can be separated from the enzyme by dialysis, and are not de-
stroyed by heating. An enzyme may be rendered inactive by the
removal of its activator, but it can be restored to activity by mixing
again with this substance. In the case of some enzymes, the inactive
substance, as it is a cell may be called a zymogen, or profer-
formed in
ment, but when associated with the activator the active enzyme is
developed. An activator is inorganic. A kinase is a more or less
complex organic body which activates a proferment.
Substances which reduce, or destroy, the activity of enzymes are
called paralyzers, which may be formed as products of enzymatic
'Haas, Paul, and Hill, T. G.: An Introduction to the Chemistry of Plant
Products. 1913: 340-341.
58 MYCOLOGY
aclivity or be foreign substances. Acetic and lactic acids formed by

enzyme activity will destroy the ferments producing them unless
neutralized. Among foreign substances which act as paralyzers
may be mentioned formaldehyde, mercuric chloride, alcohol, chloro-
form and hydrocyanic acid. Anti-enzymes are a class of substances,
which are antagonistic to the action of enzymes. The distribution of
the enzymes in the various groups of fungi including the slime moulds,
bacteria and true fungi have been investigated by a number of zymolo-
gists. For example, Monilia sitophila may form maltase, trehalase,
raffinase, invertase, cytase, diastase, lipase, tyrosinase and trypsin.
Dox^ has demonstrated in moulds, the following: protease, nuclease,
amidase, lipase, emulsin, amylase, inulase, raffinase, sucrase, maltase,
lactase, histozyme, catalase and phytase, and he has found that these
enzymes are formed regardless of the chemic character of the substratum.
Without going into all the details of the occurrence of enzymes in the
fungi, the following classification of the principal enzymes found in the
various groups may prove useful to the student.
Classification or Enzymes in Fungi
I. HYDROLYTIC ENZYMES.
(a) Carbohydrate-splitting enzymes (carbohydrases) :
Amylase, or Diastase, which hydrolyzes starch to dextrin and

maltose. The Koji fungus, Aspergillus oryzece (Taka-diastase).
Cytase, which hydrolyzes hemicellulose to galactose and mannose
in Botrytis.
Inulase, which hydrolyzes inuhn to levulose.
Invertase, which hydrolyzes cane sugar to dextrose and levulose.
Saccharomyces, Fusarium, Aspergillus niger.
Lactase, which hydrolyzes lactose (milk sugar) to dextrose and
galactose. Kephir organism.
Maltase, which hydrolyzes maltose (malt sugar) to dextrose.
Saccharomyces octosporus.
Raffinase, which hydrolyzes raffinose to levulose and melitiose.
Aspergillus niger.
Trehalase, decomposing trehalose into a reducing sugar. Poly-
porus sulphiireus.
1 Dox, A. W.: Enzyme Studies of Lower Fungi. Plant World, 15: 40, February
1912.
HISTOLOGY AND CIIEMISTKY OF FUNGI 59
(b) Protein-splitting enzymes (proteases):

Pepsin, which hydrolyzes proteins to albumoses and peptones.
Trypsin, which hydrolyzes proteins to peptides and amino-
acids in A maniia muscaria and Boletus edulis.
(c) Urea-splitting enzymes (ureases):

Urease obtained from Micrococcus urcce, which hydrolyzes urea
into ammonia and carbon dioxide.
(d) Nuclease, which spHts nucleic acid.
(e) Fat-splitting enzymes (esterases and lipases):

Lipase in Penicillium glaucum and Aspergillus niger, also
Empusa. Phycomyces, which break up fatty oils.
(/) Glucoside-splitting enzymes:

Emulsin, which hydrolyzes amygdalin to glucose, hydrocyanic
acid and benzaldehyde. Also such other glucosides as saUcin,
populin, coniferin which fungi are able to utilize.
2. FERMENTING ENZYMES.
(a) Alcoholic fermentation of glucose, levulose, mannose, etc., by
zymase in yeasts.
{b) Lactic acid fermentation of lactose by lactic acid bacteria,
(c) Butyric fermentation of lactose by the butyric acid bacteria.
3. Clotting Enzymes (Coagulation, CurdUng).

Rennin (Chymosin), which curdles milk. Bacillus mesentericus
vulgatus.
4. OXIDIZING ENZYMES.
(a) Oxidases, which oxidize alcohols to acids, e.g., the action of
Mycoderma aceti, etc.
{b) Tyrosinase. Russula nigricans and species of Boletus, Lacta-

rius, etc.
(c) Peroxidases, which set free oxygen from hydrogen peroxide,
causing this substance to blue guaiacum resin.
{d) Catalase, which decomposes hydrogen peroxide with the
evolution of molecular oxygen.
In concluding this brief study of the enzymes it may be stated that

they can be detected by chemic, bacteriologic, serologic and histologic
6o MYCOLOGY
means. Details of the occurrence of the above enzymes will be found

in the books noted in the footnote below.'
CHEMOTAXIS
The attraction or repulsion of motile microorganisms by chemical
stimulants known as chemotaxis is found in the activity of the zoospores
of the OOMYCETALES and in the growth of the hyphae of fungi in gen-
eral toward or away from the stimulus. To these phenomena the names
of positive and negative chemotropism have been given. The thorough
investigations of M. Miyoshi with Aspergillus niger, Mucor mucedo,
Penicillium glaucum, Phycomyces nitens, Rhizopus nigricans have shown
that the following substances act as powerful stimulants: ammonium
phosphate, asparagin, dextrin, saccharose and glucose. The threshold
value (marginal limit) or minimum quantity capable of producing a
chemotactic effect was ascertained by Miyoshi as o.oi per cent, in
the case of glucose acting on Mucor mucedo. On gradually increasing

the dose, a second limit is reached where repulsion occurs. The
entrance of fungi into leaves and the growth of hyphae along certain
lines inside of the host tissue and the formation of haustoria are per-
haps all indications of chemotropic response.
iRayliss, W. M.: The Nature of Enzyme Action (Monograph on Biochem-

istry). Longmans, Green & Co., 1914.
Green, J. Reynolds: The Soluble Ferments and Fermentation. Cambridge
at the University Press, 1899.
Haas, Paul and Hill, T. G. An Introduction to the Chemistry of Plant Prod-
:
ucts. London, Longmans, Green & Co., 1913.

Harden, Arthur: Alcoholic Fermentation (Monograph on Biochemistry).
London, Longmans, Green & Co., 1914.
Lafar, Franz, transl. by Salter, Charles, T. C: Technical Mycology, ii,
Pt. I : 61-65.
Marshall, Charles
E. and others: Microbiology. Philadelphia, P. Blakiston's
Son & Co., 1911
Oppenheimer, Carl: Die Fermente und ihre Wirkungen. Leipzig, 1903.
Vernon, H. M. Intracellular Enzymes. London, John Murray, 1908.
CHAPTER VII
GENERAL PHYSIOLOGY OF FUNGI

The influence of light on the development of the EUMYCETES
has been investigated by a number of workers. The influence of light
on the direction of growth is known as phototropism. On account of
the contradictory evidence of earlier investigations, Friedr. Oltmanns
experimented with Phycomyces nitens using a powerful electric arc light.
He found that Phycomyces behaved positively phototropic under weak
illumination, but negatively so under a powerful light. It remained
aphototropic with an intermediate illumination, and in young sporangial
hyphse with gray sporangia, a given degree of illumination caused
attraction, while with older sporangiophores with blackened sporangia
repulsion was noticed. The germination of the spores of such fungi, as
Penicilliwn glaucum, Trichothecium roseum, Fusariiim heterosporium,
Rhizopus nigricans, does not seem to be affected by light; while
von Wettstein found that light retarded the germination of the spores
of Rhodomyces Kochii. The evidence as to the influence of light on
the vegetative development seems to be contradictory. J. Schmitz
found that Spharia carpophila grew better in the dark than in daylight.
G. Winter found Peziza Fuckeliana to cease growth in the dark and the
fungus perishes if light be long excluded. Mac DougaP experimented
with Coprinus stercorarius. He found that it developed a much greater
length than the normal in darkness, but the fruit bodies remained in
a rudimentary or incomplete stage. After growth had proceeded in
this manner some time the illumination of the body was followed
for
by the production of fruit bodies in a manner demonstrating most
conclusively that the action in question was due to a purely stimula-
tive action of light, since the rays did not participate in any synthesis
of material.
The rate of cell reproduction does not seem to be influenced by the
presence or absence of light. In many fungi, the formation of a
1 Mac Dougal, D. T. : The Influence of Light and Darkness upon Growth and
Development. Memoirs of the New York Botanical Garden, ii (1903: 279).
61
62 MYCOLOGY
fructification does not seem to be affected by the light conditions,

but here the evidence is contradictory, some fructifications being
formed better in light than in the dark and vice versa. Kolkwitz after
eliminating various sources of error of earlier experimenters found
that in his cultures of Aspergillus niger and Oidium ladis that con-
siderable acceleration of respiration is experienced with a brief illumina-
tion by a powerful electric arc. Koernicke^ finds that Roentgen rays
inhibit the growth of fungi with prolonged action.
Luminosity of Fungi. —The luminosity of wood and decaying logs
in the forest is associated with the mycelia of certain fungi. The
phenomenon is connected frequently with gill-bearing fungi, such
as Agaricus, Armillaria mellea, Pleurotus olearius, and as determined by
Molisch with the two ascomycetous fungi. Xylaria hypoxylon and A^.
Cookei. In order to prevent any error arising in the experiments

through the presence of luminous bacteria, Molisch^ grew Armillaria
mellea, Xylaria hypoxylon, X. Cookei, Mycelium X. in pure cultures,
the latter succeeding well on bread. He found that under such con-
ditions the plants became phosphorescent. Such phosphorescence
is connected with a supply of oxygen and is not due to the separation
of some luminous substances, but is intracellular in its origin.

Liberation of Spores.—The spores of the gill fungi (HYMENOMY-
CETES) are very adhesive, when freshly set free. As a result of
this, special arrangements are found for the liberation of the spores
from the surfaces of the gills and the hymenial tubes. Paraphyses
between the special conidiophores known as basidia serve to increase
the spaces between the spores, preventing contact and allowing a
freer fall of the spores. The arrangement of the gills is such as to
economically increase the spore-bearing surface, and, therefore, the
total number of spores that a fruit body can produce. By various
growth movements of the cap and fruit stalk, the spore-bearing sur-
face is placed in the best possible position for the liberation of spores.
The spores liberated from the gills on the under surface of a pileus
placed over a horizontal sheet of paper fall vertically downward and
form a spore print, which consists of radiating lines corresponding to
the inter-lamellar spaces. The number of spores set free by large
fruit bodies is prodigious. A specimen of the mushroom Agaricus
1 KoERNiCKE, Max: Ber. d. deutsch. Bot. Ges., 1904: 22, 14H.
2 Molisch, Hans: Leuchtende Pflanze, 1904: 25-46.
GENERAL PHYSIOLOGY OF FUNGI 63
{PsalUota) campestris with a diameter of 8 cm. produced 1,800,000,000

spores, one of Coprimis annatus 5,000,000,000 and one of Polyporiis
sqnamosus 11,000,000,000 spores. Buller has estimated that a large
fruit body of the giant puffball Lycopcrdon bovisla (40 X 28 X 20 cm.)
Fig. 17. — Diagram of the discharge of spores from a fruit-body of Polyslictus

versicolor as seenby a beam of light. A stream of spores is carried round within
the beaker very slowly by convection currents and recorded. Reduced about 2/3.
{After Buller: Researches on Fungi, 1909: 97.)
contained 7,000,000,000,000 spores, or as many as 4000 mushrooms

of the size above mentioned.
Spores dropping from any fruit body which is suspended in a
closed glass chamber can be seen in clouds, or individually, without the
64 MYCOLOGY
microscope by concentrating a beam of light upon them (Fig. 17).
This is a simple method of examining the discharge of spores from the

mushroom. It can be used conveniently with the xerophytic fruit
bodies of Lcnzitcs betuUna, Polystictus versicolor, Schizophyllum com-

mune at any time in the laboratory by keeping them dry for months
and reviving them by placing them in a jar with wet cotton. They
quickly revive and begin to shed their spores in six hours and this
discharge continues for some days.
Ordinarily, spore discharge from any fruit body is a continuous
process, but if placed in hydrogen, or carbon dioxide, the liberation of
spores ceases quickly, demonstrating that oxygen is necessary. Ether
and chloroform act similarly to the gases above mentioned. The
X A B
—
second and the spores of the common mushroom shortly after leaving
the cap fall at the rate of i mm. per second approximately.
The violent discharge of the spores prevents the adhesive spores
from massing together and from sticking fast to the gill surface. At
first the spore is shot out horizontally, then under the influence of
gravity, it describes a sharp curve and then falls vertically. The
path described by the falling spore has been appropriately called a
Fig. 19. Amanitopsis vagineta. Relations of spores to the fruit-body. A,

Transverse section through two gills, h, basidia projecting, the arrows show spore
parts (sporabola), Magn. 15; B, vertical section of hymenium and subhymenium, c,
paraphyses, a-c, basidia stages, Magn. 370; C, isolated basidium with two basidios-
pores; D, discharged spore; E, basidium, Mayer, mo. (After B idler, 1909: 165. )j
sporabola (Fig. 19). There are two distinct types of fruit bodies as to
spore production and spore liberation. These are the Coprinus comatus
and the mushroom types. The deliquescence, or melting of the fruit
bodies of the Coprini is a process of auto-digestion and it assists mechan-
ically in the discharge of the spores. Spore discharge precedes deliques-
cence. The spores are set free from below upward and by auto-diges-
tion those parts of the gills are removed from which the spores have
5
66 MYCOLOGY
been shed, thus permitting the opening out of the cap and the freer
discharge of the remaining spores. The discharged spores are conveyed
by the wind (Fig. 20). The mushroom type is the usual kind where
the spores are discharged without deliquescence.
The spores of Bulgaria, Gyromilra, Peziza and others of the
AscoMYCETALES are scattered by the wind, but those of Ascpbolus
immersus and Saccobolus are dispersed by herbivores. The spores of
Peziza repanda, according to Buller, are shot up into the air to a
height of 2 to 3 cm. and leave the spore sac (ascus) together, but
Fig. 20. — Semidiagrammatic sketch in a field with horse mushroom, Agaricus

{Psalliola ) arvensis, showing Hberation and discharge of spores horizontally
and from velum. Reduced to y<i. {After Buller, Researches on Fungi, 1909: 218.)
separate as they leave the ascus mouth. Puffing is due probably to a

stimulus given'j to the* protoplasm in contact with the ascus Hd, and
it is observed when poisonous substances are applied such as iodine
mercuric chloride, silver nitrate, copper sulphate, sulphuric and acetic
acids are used. With^some of these forms the ascus may be considered
as a squirting apparatus by which a jet of spores leaves its mouth.
The writer^ noted the puffing of the spores in Peziza hadia when the
large saucer-shaped fruit bodies were held in the hand. At intervals
of several minutes the puffing took place.
Ascobolus immersus as a coprophilous (dung-inhabiting) fungus has
1 Harshberger, J. W.: Journ. of Mycol., 8: 158, October, 1902.
special adaptations: (i) the protrusion of the ripe asci beyond the general
surface of the fruit body; (2) the diurnal periodicity in the ripening of
successive groups of asci; (3) the positive heliotropism of the asci; (4)
the considerable distance to which the spores are ejected (sometimes
30 cm.) with which is associated; (5) the large size of the asci and spores;
and (6) the clinging of the eight spores together, while describing their
trajectory through the air. The forcible explosion of the sporangio-
phore of Pilobolus crystallimis by which the whole sporangium is dis-
charged a considerable distance into the air is due to the tension exerted
by gases and water vapor within the swollen sporangiophore.
The escape of biciliate zoospores (swarm spores) in such genera of
aquatic fungi as Achlya and Saprolegnia is through a terminal pore in
the zoosporangium. It appears that the discharge is associated with
the motility of the cilia. In the moulds (Mucorace^), the sporangial
wall which is coated with minute particles of calcium oxalate becomes
soluble in water at maturity and the intersporal substance swells up
assisting in the liberation of the spores. The entire inner peridium
about the size of a pin's head is forcibly ejected in the gasteromycetous
fungus, S phcBrobolus stellatus, and this is due to the unequal tension of
the different peridial layers.
The disposal of spores and conidia is facilitated by water in the
case of the motile zoospores of such fungi as Achlya prolifera, Phytoph-
thora injestans and Saprolegnia ferax, where cilia come into play.
Many spores are no doubt carried passively by water currents. Wind
is, however, one of the chief agents in the distribution of fungous spores,
such as those of the puffballs, the rusts and the moulds, although the
is probably exaggerated.
distance that such spores are carried Flies,
which feed upon the strong-smeUing slime in which the minute spores
of such fleshy fungi as Mutinus caniniis, Icthyphallus impudicus are
imbedded, assist in the carriage of such spores and those of ergot
{Claviceps purpurea) in the Sphacelia stage, where viscid drops exude
that are attractive to and although some flies arekilled by it, yet
flies,
sufficient escape to carry the spores. Slugs and snails by crawling

alternately over diseased and healthy plants, probably disseminate
spores. That birds serve as distributors of spores is indicated by the
studies of Heald with the chestnut blight fungus, Endothia parasitica,
in which he found that a single downy woodpecker carried as many as
657,000 pycnospores. Certain subterranean fungi such as truffles are

68 MYCOLOGY
eaten by rodents attracted by the strong smell that they possess and
probably the mammal is instrumental in the spread of the spores.
Many of the coprophilous fungi have spores which pass through the
alimentary canals of different animals without being destroyed and
germinating in the dung, or manure from such animals, they propagate
the species. Pilobolns crystallinus is one of them. The sporangia, which
are shot off from the sporangiophore, adhere to blades of grass, which
are eaten by horses, and later the fungus makes its appearance on horse
manure. The spores have passed through the horse apparently unaf-
fected and more readily germinable. Man with his agricultural imple-
ments is concerned with the spread of fungous spores. Giissow states
that a threshing machine, which has been used for threshing smutted
wheat, is infested so fully with spores that any grain subsequently
threshed, unless the machine is sterilized properly after use, will
become liable to infection.
CHAPTER VIII
ECOLOGY OF FUNGI
As fungi are either saprophytes or parasites, their Kfe history is
bound up with the substratum on which the saprophytes are found and
with the host plant upon which the parasite hves, yet there are many
diverse forms of saprophytic fungi and the greatest variety of fungous
parasites. Of special interest in connection with the ecology of fungi
are the organsby which various fungi are tided over periods of drought,
inclement seasons, or during the winter's cold. These organs are
compacted masses of hyphse of a rounded, globular, or ellipsoidal form
ranging in size from those that are almost microscopic to those which
are the size of a small canteloupe. These tuber-like masses of hyphae
in a resting state are known as sclerotia (Gr. ayXupos, hard). They
are found in a great many fungi, as commonly in the ergot, Claviceps pur-
purea, a.nd the lettuce drop, Sclerotinia libertiana, which forms sclerotia
that may reach a length of 3 cm. in exceptional cases. These sclerotia
are obtained readily in culture tubes with beerwort agar, or glucose agar,
as culture media. From the sclerotium later arises the stalked fruit
body, or apothecium. Cordyceps militaris is a fungus which attacks
the larva of insects. Itsmycelium penetrates the insect's body and

later in the Isaria form produces aerial hyphas which cut off conidio-
spores. The growth of the mycelium is such as to penetrate to all parts
of the larva filUng it up as if it were stuffed with cotton.
The mass of hyphae is converted into asclerotiumand the larval body
is mummified, but still retaining its original external form. Later, the
next spring, a stiff-stalked stroma arises with an enlarged extremity in
which the perithecia with their asci- and ascospores are formed. Later
the needle-shaped ascospores are set free and by cutting off conidio-
spores reproduce the disease. Cordyceps (Torrubia) ophioglossoides is
parasitic upon an underground truffle, Elaphomyces muricatus, Fig.

21). The stroma is erect, yellow and club-shaped at the extremity.
and ascospores are borne in the swollen part of the
Perithecia, asci-
stroma. The fungus which discharges its spores above ground finds the
69
—
70 MYCOLOGY
Fig. 21. A Cordyceps tnilitaris; B, Cordyceps Hugelii on a caterpillar; D, Cordy-

ceps spharocephala on a wasp; E, Cordyceps cinerea on a beetle; F-K, Cordyceps
ophioglossoides, F on a deer truffle; G, ascus; H, conidiophore; J, conidiospores; K,
germinating spore. See Die naliirlichcn Pflanzenfamilicn I. i, p. 368.
1
ECOLOGY OF FUNGI 7
underground truffle in Llie following manner. When the spores germi-

nate, they give rise to hyphaiwhich grow over a densely cespitose, com-
mon moss, Mniiim horniim, which develops a large number of feeding
rhizoids, that penetrate the soil to the depth at which Elaphomyces
grows. The mycelium of Cordyceps not only covers the aerial portions
of the moss, but follows the rhizoids underground until they reach the
underground truffle over which the moss may happen to grow. Bot-
anists searching for Elaphomyces always know where to look for it by
the presence of the Cordyceps hyphae, on the moss Mnium hornum.
There is a black beetle, a native of France^ with a pale, velvety abdo-
men, known as Bulboceras gallicus, about as large as a cherry stone. By
rubbing the end of the abdomen against the edge of the wing cases it
produces a gentle chirping sound. The male has a horn on his head.
This insect burrows in the soil among the trees of the pine forests and is
nocturnal in its habits. It descends vertically into the soil in search
of the underground truffle-like fungus, Hydnocystis arenaria, upon
which the insect rabassier feeds. The fungous fruit body is about the
size of a cherry with a reddish exterior covered with shagreen-like warts.
The beetle, which feeds upon Hydnocystis arenaria and Tuher Requenii
one of the truffles, locates the fungi by a subtle sense of smell. The
human truffle hunter finds these underground by the burrows which
the beetles make in digging for their chief source of food and he usually
finds groups of these fleshy funguses directly beneath the openings of
the beetle holes.
Rozites gongylophora is a gill fungus which is raised as a fodder by
leaf-cutting ants in their subterranean passageways in the tropics of
South Brazil.
On a visit to the Berlin Botanical Garden in 1898, the writer noted
the following remarkable examples of sclerotia-bearing fungi: Poly-
porus sapurema A. Moller (Fig. 92, Teil I, Abt. i**, Die naturlichen
Pflanzenfamilien, p. 171). The sclerotium is over 30 cm. in diameter
and weighs at least 20 kg.It is furrowed and roughened and leather
colored. A specimen from Blumenau, Brazil, developed in the Victoria
house of the Berlin Garden four largepilei in August and September,
1897. Polyporus found in Australia produces a sclerotium

mylittce
{Mylitta australis Fr.), which as "native bread" is used as food
by the natives. Polyporus tuhcraster, which grows in the mountains
1 Fabre, F. H.: Social Life in the Insect World, 1912 217-237.
72 MYCOLOGY
of Italy, develops a large edible sclerotium called by the natives

pietra fungosa. The sclerotium of Polystictus socer (Fig. 94 A, Teil I,
Abt. I**, Die naturUchen Pflanzenfamilien, p. 177), known as
Pachyma malacense, is cm. long, brownish red
of variable shape, 8 to 10
externally with a white interior. found in the Malay Archipelago.

It is
These are a few of the true sclerotia which probably includes the
"tuckahoe" of the North American Indian, Pachyma cocos.
Living on limbs, twigs and the leaves of the beech in the deep
shade of the forest is found a scale insect {Schizonema imbricator) ,^
which is covered by a woolly coat consisting largely of a waxy secretion
from the body. This woolly material is quite abundant and where the
insects live in masses together the entire limb, or leaf surface has a
downy white appearance. The abdomen
of the insect moves con-
stantly with a jerky motion and the cottony material is, therefore,
constantly agitated. The insects secrete a honey dew so copiously
that it runs down to the leaves beneath and to the ground. Upon this
honey dew and the dead bodies of the scale insect, a pyrenomycetous
fungus, Scorias spongiosa, lives. It grows as a spongy mycelium con-
sisting of much-branched, rigid, septate hyphge with the strands glued

together by a mucilage. Pyriform perithecia, long-necked spermogonia
and pycnidia are formed from the mycelium, which is saprophytic
on the products of the insect's body.
The anther smut of the caryophyllaceous flowers occurs in America
and Europe on Cerastium viscosum, Saponaria officinalis and Silene
inflaia, and on species of Dianthus, Lychnis, Melandrium, Stellaria,
etc. The spores of this smut replace the pollen grains in the anthers
of these plants and when the flowers open a violet smut dust is dis-
charged from the anthers instead of the pollen. Female flowers of
Melandrium attacked by the fungus show a marked morphologic
differentiation in the development of mature stamens out of staminal
rudiments. These anthers are invaded by the fungus and in them the
parasite fructifies.
The formation of galls is a marked feature of the ecology of fungi.
One form of these malformations is seen in the witches' broom (hexen
besen) which are due to the attack of a number of species of Exoascus
on different forest trees; The branchlets are clustered into broom-like
masses with leaves that are somewhat altered in shape and fall earlier
^Harshberger, J. W.: Joum. of Mycol., 8: 160, October, 1902.
ECOLOGY or FUNGI 73
than those on normal twigs. Witches' brooms are found on such conif-
erous trees as the white cedar in New Jersey and are due to Gymno-
FiG. 22. — Black knot of plum, Plowrighlia morbosa, on beach plum, Priinus marilima
Nantucket, August 17, 1915.
Sporangium Ellisii, a rust fungus. These malformations occur on the

hackberry, but on this tree they are due to the attack of a mite Phy-
toptus followed by a fungus. Plum pockets are a form of gall in which
74 MYCOLOGY
the fruit is enlarged by the attack of the fungus at the expense of the
stone which fails to develop. The hollow
on the plum are due to
galls
Exoascus pruni. The on our red cedar trees in
so-called cedar apples
the spring are caused by the attack of an annual rust fungus, Gym-
nosporangium juniper i-virgini ana, and from the surface of these
apples two-celled spores arise. The white rust of cruciferous plants,
Cystopus candidus, produces blisters on the leaves and stems of shep-
herd's purse. The black knot of the plum is a tumor-like swelling of
the branches of plum trees due to the attack of an ascomycetous fungus,
Plowrightia morbosa (Fig. 22). Large swellings on oak trees the size of
a man's head and over are caused by a fungus, Diachana strumosa, and
some of these swellings may be the size of a large pumpkin. Galls due
to insects are frequent on plants, but a discussion of them is extralimital.
According to conditions of environment, we may briefly treat of
fungi as hygrophytic, mesophytic and xerophytic forms. The hygro-

phytic forms include the aquatic fungi, such as Achlya^ Mono-
ble pilaris, Saprolegnia live and carry on
and other genera which
their reproduction in water. group would belong
Perhaps to this
a fungus of the genus Cyttaria, which was found by Darwin in the

beech (Nothofagus) forests of southern Patagonia. The beech trees
grow in cold, wet valleys completely barricaded by great moulder-
ing trunks of former beech trees on which the globular, bright yellow
fructification occurs and which is eaten by the Fuegians.
The mesophytic forms include many of the common fleshy gill
fungi that live in our woodsand forests, appearing in surprisingly great

numbers after a spell of wet weather. Here we might include species
of Amanita, Boletus, Russula, and Clavaria and others which are not
infrequent, while in our meadows occur mushroom and coprini. Three
conditions seem favorable to their growth: abundant leaf mould,
warmth and abundant moisture.
The habitats of the fleshy fungi are of general interest. Collybia
platyphylla develops its fruit bodies on the shaded side of decaying
logs. The fairy-ring fungus, Marasmius oreades (Fig. 23) produces its
sporophores in lawns in the form of rings long known as fairy rings.
Frequently grassy spots are enclosed by the circle of toadstools which
are several feet in diameter. The fruit bodies of Pholiota adiposa
(Fig. 24) grow from wounds in living trees.
In forest operations the slash, when scattered, rots more rapidly
than when piled. This is due to the fact that two types of fungi
ECOLOGY OF FUNGI 75
are active in rotting the brush, one set entering the Hmbs and branches
above the ground and the other gaining access to the brush actually
in contact with the soil. Brush is rotted at the top when piled
with one group of fungi and at the bottom by another group, while
the middle of the pile, not in contact with the soil and yet protected
from the sunlight, apparently will not rot to any extent until the
Fig. 23.— Fairy ring formed by Marasmius oreades, an edible toadstool. (From
Wiley, Foods and Their Adulteration. After Coville, Circular 13, Division of
Botany.'
pile disintegrates suflSiciently to expose these central layers to the

soilmoisture on the one hand, or to the sunlight on the other.
Four fungi cause rotting of oak slash in Arkansas, viz., Stereum
rameale, S. umbrinum, S. versiforme and S. fasciatum. Two fungi are
responsible for the decay of short-leaf pine slash. They are Lenzites
sepiaria and Polyslictus abiet'mus.^
The xerophytic forms are those which have corky or leathery fruit
^Lona, W Investigation of the Rotting of Slash in Arkansas. U. S. Dept.
II.:
Agric. Bull. 496, Feb. 16, 191 7; also Humphrey, C. J.: Timber Storage Conditions in
the Eastern and Southern States with Reference to Decay Problems Bull, 510, U.
S. Dept. Agric, May 17, 1917.
76 MYCOLOGY
bodies growing on sticks and logs where they can dry up without any
loss of vitaHty. They revive after a rainfall and resume the function
of discharging spores and the discharged spores are capable of germina-
—
ECOLOGY OF FUNGI 77
Fig. 25. Schizophyllum commune, a xerophyte. A and B, fruit-bodies seen

from above growing on wood, natural size. C and D, two fruit-bodies seen from
below and in section; about twice magnified; £, section through pileus in wet weather
showing gills split down their median planes; F, section of a dry pileus; E and F
about 12 times natural size, (after Buller, Researches on Fungi, 1909: 114.)
78 MYCOLOGY
existence (Fig. 25). "The gills are partially or completely divided

down median planes into two vertical plates. While desiccation
their
is proceeding, the two plates of each of the longer and deeper gills bend
apart and spread themselves over the shorter and shallower gills.
When desiccation is complete, the whole hymenium is hidden from
external view and the fruit body is covered both above and below with a
layer of hairs (Fig. 25). The closing up of the fruit bodies at the
beginning of a period of drought serves to protect the hymenium. A
fruit body which retains its vitality even when dry for two years will
revive again in a few hours and spores are discharged" (Fig. 25).
As it is not the purpose of this book to consider the so-called Hchens
in the classification which follows as distinct entities in which the
lichen fungus and the lichen alga are in symbiosis forming a
lichen thallus, important to describe the ecology of the actual
it is
relationship of the two plants to each other, as a matter of botanic

interest. Danilov, Elenkin, Peirce and Fink have shown that the dual
hypothesis, or that of mutuahstic symbiosis, is untenable. A lichen
is a fungus belonging to the orders ASCOMYCETALES, or BASIDIO-
MYCETALES, which lives during all or part of its life in parasitic
relation with an algal host and also sustains a relation with an organic
or an inorganic substratum. Having squarely assumed this position
as to the true nature of what currently passes for a lichen, it is interest-
ing to note that there are ten algae known as Hchen hosts: Chlorococcum
(Cystococcus) humkola, Pahnella botryoides, Trentepohlia (Chroolepus)
umbrina, Pleurococcus vulgaris, Dactylococcus infusionmn, Nostoc lichen-
oides (?), Rivularia nitida, Polycoccus pimctijormis, Gleocapsa polyderma-
tica and Sirosiphon pulvinatus. It is important to note, that although
the larger number of the above are blue-green algae, yet the two species
of green algae. Chlorococcum humicola and Trentepohlia umbrina form
the hosts of many more lichens than all the others combined. Hence
the student of these plants can study the algicolous fungi, mainly
ASCOMYCETELES, a few BASIDIOMYCETALES, those parasitic
upon algje, as the lichens, while the non-algicolous fungi can be over-
looked by the lichenologists. We can do no better than quote Bruce
Fink,^ who sums up the main arguments against mutualism and the
1 Fink, Bruce: The Nature and Classification of Lichens. I. Views and Argu-
ments, Mycologia, iii: 231-269, September, 1911; II. The Lichen and its Algal Host,
Mycologia, iv: 97-166, May, 1913.

:
ECOLOGY OF FUNGI 79
advocation of the fungal nature of lichens, as follows: "Lichens com-

monly grow where there are free algae of the same species as those
by these Uchens. The spores of the lichens germinate and
parasitized
attack the free alga; as other fungi attack their hosts. Lichens perform
like other fungion culture media and may be made to produce their
reproductive organs on these media. Lichen spores also attack the
algal hosts,when the spores and the algae are introduced into cultures
together; and the resulting lichen is normal and sometimes fructifies
in the cultures. Algal hosts extracted from lichen thalli grow in cul-
tures like free algae of the same species grown on similar culture
media. The researches of Elenkin and Danilov prove that lichen
hyphae absorb food from the algal host cells, which are killed by
severe parasitism, or more probably by parasitism and saprophytism
combined. The relation of the lichen to its substratum proves that
higher lichens can take comparatively little food from it and must
depend more than lower lichens upon the algal hosts; and this shows
that the parasitism of the lichen upon the algal host has become more
severe in the evolution of the higher lichens. Finally, the algae para-
sitized by lichens are in a disadvantageous position with reference to
carbon assimilation.
"Lichens are like other fungi with respect to vegetative structure
and fruiting bodies. The bridges which connect lichens with other fungi
are not few, but many. Since it is thoroughly demonstrated that the
lichen is parasitic, or partly parasitic and partly saprophytic on the
alga, there is no longer even a poor excuse for a 'consortium' or an
'individualism' hypothesis.
"The parasitism of lichens on algae is peculiar in that the unicellular
or the filamentous hosts are enclosed usually by the parasite, which
carry more or less food to the host. The host inside of the parasite is
placed in a disadvantageous position with reference to carbon assimila-

tion and may depend, for its carbon supply, more or less upon material
brought from the substratum by the parasite. Some algal individuals,
not yet parasitized, may be found in most lichen thalli."
Lichen thalli are of three kinds: crustaceous, foliose and fruticose.
The arrangement of the layers of the lichen fungus and its algal host
varies in different lichens, but in Stkta the following are met in a ver-
tical section of the thallus (Fig. 26)
(a) Tegumentary layer.
8o MYCOLOGY
Fig. 26.— A foHaceous lichen, Parmelia perlata. i, Plant slightly reduced in

size; a, apothecia; b, lobe of thallus; c, patches of soredia; 2, longitudinal section of
apothecium and cross-section of thallus; a, ascus; b, c, hypothecium; d, upper gonidial
(upper algal) layer; e, medullary layer;/, lower gonidial layer; g, lower cortical layer;
I, 3, cross-section of vegetative thallus. (From Gager. After Schneider.)
ECOLOGY OF FUNGI 8l
(b) Upper cortical layer.

(c) Algal layer (gonidial layer).
(d) Medullary layer.
(e)Lower cortical layer.
The tegumentary layer consists of several rows of flattened hyphal
cells extending at right angles to the underlying cortical cells which
consisting of hyphal cells are pseudoparenchymatous, resembling the
parenchyma tissue of higher plants. The algal layer contains the
gonidia, or green plants, which act as hosts to the fungous hyphge. The
medullary layer which is thicker than the others consists of much
elongated hyphae forming a loosely interwoven tissue with large air
spaces. The lower pseudoparenchymatous and from its
cortical layer is
lower surface rhizoids are developed. The apothecia and perithecia

are the fruit bodies of the ascomycetous fungi which form the lichens.
A vertical section through an apothecium of Sticta shows the following
layers: (a) the epithecium, (b) the thecium consisting of the spore
sacs (asci) and paraphyses, (c) the hypothecium or hyphal structure
immediately below the thecium, (d) upper algal layer, (e) medullary
layer, (/) .lower algal layer, (g) cortical layer (Fig. 26).
Some of the fruticose lichens have a central core-like strand of hyphae
running through the medullary region which serves as supporting
mechanic tissue as in Usnea barbata. The soredia are vegetative repro-
ductive bodies consisting of from one to many algae surrounded by
continuous hyphal tissue and are common upon the upper surface and
margins of most of the higher lichen thalli. Among the Basidio-
LiCHENES basidia are formed with basidiospores on sterigmata as in
Cora, Dictyonema, Laudatea.
CHAPTER IX
FOSSIL FUNGI AND GEOGRAPHIC DISTRIBUTION
Fungi in the Fossil State.^ —All the known fossil fungi numbering
over 400 species have been figured and described by Meschinelli in his
"Fungorum fossilium omnium Iconographia " published in 1898.
Zeiller in discussing the chronologic sequence of the groups of fungi
states that representatives of the families Chytrideace^, Mucora-
CE^ and Peronosporace^ have been found in the tissues of the
higher plants preserved in rocks of lower Carboniferous and Permian
ages. Many different plants extending from the Carboniferous period
upward show various forms of the ASCOMYCETALES on leaves and
in the tissues especially those of the stems. The fleshy fungi of the
famines Agaricace^ and Polyporace^ have been found in deposits
of tertiary age. Weiss has announced the discovery of a mycorrhiza
in the root of a probable Lycopodiaceous plant of the lower Carbonif-
erous strata. Where Polyporus and Lenzites occur, as in the brown
coals, silicified woods occur which have been half destroyed by their
mycelia.
GEOGRAPHIC DISTRIBUTION OF FUNGI
This important and interesting subject can be presented in the barest
outline. The modern teaching of geography emphasizes home geog-
raphy as a fundamental study. In following this suggestion in the
investigation of the local fungi, it will be found that we must deal
with distinct habitats, such as leaf mold, sandy soil, wet soil, decayed
logs, tree stumps, living trees, living herbs and the like. The black
mould, Rhizopus nigricans, is one of the commonest of fungi. It occurs
on bread and other organic substrata, such as sweet potatoes, whenever
the conditions are suitable for its growth. If horse manure is covered
with a bell jar with wet paper inside, there develops first the gray
mould, Mucor mucedo. This is accompanied or followed by Pilobolus
^Seward, A. C: Fossil Plants, 1898: 207-222.
Weiss, F. E.: A Mycorrhiza from the Lower Coal Measures. Annals of Botany,
xviii: 255 with 2 plates.
82
FOSSIL FUNGI AND GEOGRAPHIC DISTRIBUTION 83
crystalliniis, and this in turn by the white flecks of Oospora scabies.

Coprinus stcrcoraritis usually completes this series of coprophilous
fungi generally found on horse dung. Sometimes the Mucor is para-
sitized by Piplocepkalis and sometimes by Chatocladium. Peziza
coccinea is attached to dead twigs buried in the forest leaf mould, and
as it rises to the surface, it develops a long stipe with a crimson-red
saucer-shaped apothecium at its extremity. Russula emetica, R.
virescens, species of Clavaria and Boletus are regularly found beneath
deciduous trees growing out of the forest litter. The pufifball, Sclero-
derma vulgare, is found on the tops of old stumps in gregarious clusters.
Polyporus sulphur ens grows out of partly dead chestnut and oak trunks;
while the hymenophores of Armillaria mellea are found clustered about
the bases of trees beneath the bark of which the rhizomorphs will be
found growing. A species of Hydnmn was found a few feet above the
ground on a beech tree and Fistulina hepalica attached to tree trunks,
where the swollen base gradually blends with the straighter hole above.
Amanita muscaria and A. phalloides grow in solitary splendor at the
edges of woods and copses, while the habitat of the mushroom in open
fields is quite distinctive.
The earth-star, Geaster hygrometricus, grows more frequently in

sandy soil, where it spreads out its peridial segments.
The habitat of the local species of the lichen fungi is of interest.
The brown-fruited cup stumps
cladonia, Cladonia pyxidata, grows on
and on the earth, while the scarlet-crested cladonia, Cladonia cristatella,
is found on dead wood. The Iceland moss, Cetraria islandica, grows
on the ground as also the reindeer-lichen, Cladonia rangiferina, in ex-
tensive masses. Another earth-inhabiting form is Peltigera canina.
The trunks of trees are marked by the presence of Parmelia perlata
and the fruticose bearded lichen, Usnea barhata. Smooth bark appears
covered with runic character traced by the fruit bodies of Graphis
scripta. The rock-dwelling lichens include Physcia parietina and the
rock tripe (tripe de roche), Umbilicaria which grows on the outcrops
of Octorara schists at the Gulph.
The distribution of the chestnut blight fungus, Endothia parasitica,
is ofmore than local interest, although the agitation to control it
started near Philadelphia. Apparently the fungus was introduced from

China, where it has been found recently, with nursery stock into Long
Island. From the neighborhood of New York City, it spread northeast,
84 MYCOLOGY
northwest, west and southwest.^ Now it is found in Connecticut,

New York, throughout New Jersey, and as far west as the Alleghany
mountains in Pennsylvania. In isolated areas, it occurs in Virginia
and West Virginia, endangering the future of the chestnut tree in
America (Fig. 27).
Wherever the cultivation of the higher plants extends, the fungi
pecuUar to these plants will be found, as the wheat rust, Puccinia
Fig. 27. —
Map of the eastern United States showing distribution of chestnut
blight disease in ipii. Horizontal lines indicate area with approximately all the
trees dead; vertical lines approximate area where infection is complete; dots indicate
advanced points of infection. {From Gager, after Metcalf, U. S. Farmers' Bull. 467.)
graminis, in Europe, America and Australia. The damping-off fungus^

Pythium de Baryanum, which is death to seedlings, has been studied by
German, English and American botanists, as a reference to the litera-
ture will show. The downy mildew, of the grape, Plasmopara viticola,
apparently of eastern American origin, is found now in, Europe and
Cahfornia, where it has become a serious pest.
The black knot, Plowrightia morbosa, was apparently at one time
confined largely to the Atlantic seaboard and was particularly abundant
in New England and New York. It has now spread across the northern
' Cf. Stevens, Neil E.: Some Factors influencing the Prevalence of Endothia
gyrosa. Bull. Torr. Bot. flub, 44 127-144. March, 191 7.
:
FOSSIL FUNGI AND GEOGKAPHIC DISTRIBUTION 85
United States to the Pacific coast. Such diseases as the sooty mold of
orange, Meliola camellicB, and the brown rot of the lemon, Pythiacystis
citriophthora, arc confined to these last plants and to the regions where
the citrus fruits grow. The anthracnose of the sycamore, Gnomonia
veneta, is parasitic upon the leaves and shoots of the sycamore or plane
tree, Platanus occidentalism causing its leaves to dry up, as if bitten by
early frosts. seems to be more prevalent in the bottom of valleys,
It
where the plane tree grows along streams, as here we find cold-air
drainage. Sometimes after the first crop of leaves is lost, a second
crop appears. Wherever the sycamore grows, Gnomonia may be ex-
pected. The so-called fly-cholera fungus, Empusa muscce, is parasitic
in flies and is present on these insects in Europe, even in the far north,
in North America and South America (Argentina). The coprophilous
fungus, Basidioboliis ranarum occurs on the dung of frogs in Europe
and America. Taphrina ccsrulescens does not seem to be choice about
its hosts, occurring as spots on the leaves of Quercus cerris, puhescens,
sessiliflora in middle and southern Europe and on Quercus alba, aquatica,
coccinea, laurifolia, rubra, velutina in North America. The hairy

earth-tongue, Geoglossmn hirsutum, is truly cosmopoHtan, as it has been
reported from all over Europd, North America, Java, Mauritius and
Australia. The genus Cyttaria with eight ascospores in each ascus in-
cludes six species. C. Darwinii and C. Berterii were discovered by
Darwin in Patagonia. C. Gunnii occurs in Tasmania and C. Harioti in
Terra del Fuego. None of the species, therefore, are found outside of
the southern hemisphere (Fig. 28). The genus Hypomyces includes
species which live parasitically, or saprophytically, on other fleshy
fungi. H. ochraceus lives on species of Russula in Germany, England
and North America; H. chrysospermus occurs on species of Boletus in
Europe; H. aurantius on Polyporace^ and Thelephorace^ in
Europe; H. lateritius on Lactarius in Europe and North America; H.
vlolaceus with its tender small stroma and violet-colored fruit body lives
on a slime mould Fuligo septica in northern Europe; H. viridis is found
on species of Lactarius and Russula in northern Europe and North
America; //. cervinus grows on Helvellace^ and large Pezizace^ in
Europe; H. fulgens appears on the bark of pine trees in Finland and
Sweden; H. Stuhlmanni is confined to Poly poms bukabensis in Central
Africa; H. chrysostomiis is reported from Ceylon and H. flavescens on
a Polyporus in North America. Hypomyces lactijluorum planes down
86 MYCOLOGY
the gill surfaces of the Lactarius sp. on which it grows, converting an

otherwise grayish-white fruit body into a cinnabar-red one. It is
found in the woods about Philadelphia, Pa. The fungi belonging to

the family Laboulbeniace^
are included in 28 genera and approxi-
mately 152 and have been made known largely through the
species,
studies of Prof. Roland Thaxter of Harvard University. A few species
are found in Europe, in the tropics of Africa, America and Asia, but
North America is extraordinarily rich in specific forms. They occur
on dipterous, neuropterous and coleopterous insects, especially those
which live in damp places or in the water. The corn-smut Ustilago
maydis is a parasite confined exclusively to the maize plant, Zea mays,
and to the closely related if not identically the same grass the teosinte;
EuchlcBna mexicana as pointed out some years ago by the writer^ as
proof of the common origin of these two grasses. Wherever maize is
cultivated the smut is found associated with it.
The rusts (Uredine^) are arnong the most specialized of fungi in
their parasitic habits, some species being confined to one or two hosts.
They ascend with their host plants above the snow line on high moun-
tains and toward the poles wherever flowering plants and ferns grow.
Whole genera are confined, however, to certain regions. Thus the
genus Ravenelia which lives on mimosaceous and caesalpinaceous plants
extends north to the 40° north latitude. Many rust fungi are iden-
tically the North America, north and middle Europe, and of
same in
the 500 species known from North America and 400 European rusts
approximately 150 species are common to both countries. Only a few
Mediterranean species are found in North America, as Uromyces gly-
cyrrhizcB and Puccinia Mesneriana. A less number of species are com-
mon to North and South America. It is noteworthy that Puccinia
malvacearum introduced into Spain from Chile in 1869 has in the forty-
six years which have elapsed since its introduction into Europe spread
over the world.
The genus Exohasidium includes 18 species of fungi which cause the
formation of fleshy galls chiefly on plants of the family Ericace^.
Tabulated the principal species are:
Exobasidkim laccinii on Vaccinium; Europe, Siberia, America.

Exohasidium rhododendri on Rhododendron, Europe, America.
1 Harshberger J. W.: Cont. Bot. Lab. Univ. of Pa., 1901: 234.
FOSSIL FLTNGI AND GEOGRAPHIC DISTRIBUTION 87
on Ledum, Finland.
Exohasidiiivi Icdi
Exobasidium andromcdcc on Andromeda, Europe, North America.
Exobasidimn azalew on Azalea, North America.
Exobasidium anlarclicum on Lcbetanlhus, Patagonia.
Exobasidium gaylussacice on Gaylussaeia, Brazil.
Exobasidium leucothoes on Leucolho'e, Brazil.
Exobasidium lauri on Laurus, Italy, Portugal, Canaries.
Exobasidium Warmingii on Saxifraga aizoon, Greenland, Tyrol, North Italy.
In closing this consideration of the geographic distribution of the

fungi, the interest which attaches to it as a study may be best empha-
sized by giving in tabular form the distribution of the species belonging
to a single family. The family Clathraceê includes eleven genera of
highly specialized morphology.
Family Clathrace^.
1. Clathrus cancellatiis , Mediterranean Region, South England,
North America.
Clathrus columnatus, North and South America.
2. Blunienavia rhacodes, Brazil.
3. Ileodidyon cibarium, Australia, New Zealand, South America.
4. Clathrella chrysomycelina, Tropic South America.
Clathrella pusilla, Australia, New Caledonia.
Clathrella kamerunensis Cameroon.
,
Clathrella Preiissii, Cameroon.
Clathrella crispa, Central and Tropic South America.
5. Simhliim periphragmoides, Tonkin, Java, Ceylon, East Indies,

Mauritius.
Simblum sphcerocephalum, North and South America.
6. Colus Miilleri, Australia.
Colus hirudinosus, Mediterranean Region.
Colus GarcicB, Tropic South America.
Colus Gardncri, Ceylon.
7. Lysurus mokusin, China.
8. Anthurus borealis, North America.
Anthurus Clarazianus, Argentina.
Anthurus Woodii, Natal.
Anthurus Mullerianus, Australia.
Anthurus cruciatus, Tropic South America.
88 MYCOLOGY
9. Aseroe rubra, New Zealand, Australia, Java, Ceylon, Tonkin,

South America.
10. Calathiscus sepia, East Indies.
Calathiscus Puiggarii, South Brazil.
II Kalchbrennera corallocephala, Africa, Cape, Natal, Angola,
Cameroon, Zambezi Region.
CHAPTER X
PHYLOGENY OF THE FUNGI
One most consistent attempts at representing the phylogeny
of the
been made by Dr. O. Brefeld through his researches
of the fungi has
which were pubHshed in collected form in " Untersuchungen aus dem
Gesammtgebiet der Mykologie. " As these volumes are bulky ones, a
student of Brefeld, Dr. F. von Tavel, has given a useful summary of
the chief points in his teacher's system in a book pubhshed in 1892,
entitled " Vergleichende Morphologie der Pilze." The phylogeny of the
higher fungi, according to Brefeld, based on the assumption, that
is
there is an entire absence of sexual organs in all of those groups above

the PHYCOMYCETES, but this view has been rendered untenable
owing to the discovery of undoubted sexual organs among the ASCOMY-
CETALES and the discovery of nuclear fusions in some of the rusts,
suggesting a sexual condition. However this may be, Brefeld and von
Tavel hold that the PHYCOMYCETES are algal-like fungi and prob-
ably derived from algal ancestors.
The OOMYCETALES are not linked directly with any of the higher
fungi, but the ZYGOMYCETALES through the former with sporangia
and conidia have probably given rise to the HEMIASCI and directly
through them to the ASCOMYCETALES. The forms of ZYGOMY-
CETALES with conidia above are phylogenetically connected in the
Brefeldian system though the HEMIBASIDII with the BASIDIO-
MYCETALES. This in brief is an outline of the phylogenetic views
of Brefeld, as expressed in a useful ground plan of the natural system
of hyphal fungi by von Tavel.
The question is asked naturally, whether the origin of the fungi has
been monophyletic, that is from a single ancestral form, or polyphyletic,
from a number of distinct ancestors? This question can be answered
only after an examination of the evidence. There are two orders of the
PHYCOMYCETES, or algal fungi, namely, ZYGOMYCETALES and
OOMYCETALES. As to the origin of these forms, the monophyletic
view would have us derive the ZYGOMYCETALES from the OOMY-
go MYCOLOGY
CETALES, which have been derived in all probabiUty from an alga

like Vaucheria with oogonia and antheridia, where the male sexual
organs are smaller than the female. To derive the ZYGOMYCE-
TALES from such a group would necessitate that the sexual organs
become of equal size.
Entomophthora is a connecting form where the sexual organs ap-
proach each other in size. This genus is then connected by insensible
differences with the heterogamic hermaphroditic moulds where there is
an appreciable difference in the size of the two cells that conjugate, the
and
larger being the female, the smaller the male, as in Absidia spinosa
These are directly connected with the
Zygorhynchus heterogamus.
homogamic hermophrodite moulds and these with the homogamic
heterothallic forms. The polyphyletic -view necessitates the deriva-
OOMYCETALES from a Vaucheria-like ancestor, and the
tion of the
ZYGOMYCETALES from a Zygnema-like ancestor, where conjugation
of similar cells (gametes) is found. The polyphyletic origin of the
fungi emphasized by the adherents to the doctrine of the origin of the
is
AscoMYCETALES from red alg£e, as there are three points of contact:

first, sac fungi with highly developed trichogyne (sterilized archicarp) of
the Collema type with red algae-like certain existing forms; second, sac
fungi with highly developed trichogyne of the Poly stigma type; third,
sac fungi with simple generalized copulating gametes of the Gymnoascus
type. We are, however, not in the position to name any known red
alga as the progenitor of the sac fungi, and it is far more reasonable to
search for one in another fungous line, where, in the hght of present-day
knowledge, there are known forms with sexual organs very much like the
sexual organs of simple, known forms of the Ascomycetales. We are
not now in a position to name any known phycomycete as a probable
ancestor, though the likehhood is that the original stock possessed phy-
comycetous characters, thus attributing a monophyletic origin to them.

One of the most instructive forms suggesting a mode of transition from
the PHYCOMYCETES to the ASCOMYCETALES, is Dipodascus.
Its sexual organs are strikingly like those of certain MucorAce^ or
Peronosporace^ young stages. The sexual organs can be
in their
recognized as antheridium and oogonium either from tlie same thread
(homothaUic) or from different threads (heterothallic). After absorption
of the wall between the gametes, the fertilized oogonium (or zygote)
grows out into an elongate stout ascus, or zygogametangium with the
PHYLOGENY OF THE FUNGI 91
production of numerous spores.^ Eremascus also represents such a con-

necting form. From Eremascus by reduction forms like Endomyces
arose which in two diverging series connects various ascomycetous
fungal forms. One series shows sprout conidia, the other oidia. The
yeast series, the Exoascus series are thus connected. Some would have
us derive the Laboulbeniace^ from red algal ancestors, but another
opposing view is that these unusual fungi have had a Monascus-like
ancestor. The other branch leads to the Basidiomycetales where the most
primitive forms have not typical basidia, as in the Hemibasidii, and
which are connected with such primitive types as are included in the
family. Entomophthorace^.^ The differentiation of types within
these large phyla will be dealt with as we proceed with a discussion of
the various groups of PHYCOMYCETES and MYCOMYCETES.
1 Atkinson, Geo. F.: Phylogeny and Relationships in the Ascomycetes. Annals
Missouri Botanical Garden, II: 315-376.
2 C/. Engler und Peantl; Die natiirlichen Pflanzenfamilien, I Teil Abt.:
60-63.
Masses, George: A Text-book of Fungi, 1906: 182-195.
CHAPTER XI
MOULD FUNGI
SUBCLASS PHYCOMYCETES
The fungi of this subclass are distinguished by their siphon-like
hyphge, because these hyphae are unicellular and multinucleate and sug-
gest the algae of the family Siphonace^
which Vaucheria belongs.
to
Hence the fungi of the subclass PHYCOMYCETES {4>vkos,
seaweed +
livKTis, a fungus) are usually designated as algal fungi. Although the
absence of transverse septa in the hyphae is used as a fundamental char-
acteristic, yet in the formation of the reproductive organs transverse
walls or septa cut these organs off from the rest of the vegetative
mycelium. Transverse septa are found regularly in some of the genera,
such as Dimargaris, Dispira, Protomyces and Mucor, so that the general
statement above is modified by such exceptions. A fungus, Leptomitus
lacteus, found in ditches and rivers shows a characteristic segmentation
of the hyphae, where through the deposit of a substance known as cellu-
lin the lumen of the hyphae is nearly closed, but at the point of constric-
tion, a smallpore remains through which the protoplasm passes.^
There are genera of the family Chytridiace^, such as Reessia and
Rozella in which the protoplasm during the vegetative state is not sur-
rounded by a cell wall, but is naked, and amoeboid in the host cells.
The fungi of this subclass are saprophytic or parasitic, aquatic, or
aerial, living endophytically as a rule. A few are parasitic on insects
and fishes. Two orders are distinguished, viz., the ZYGOMYCET-
ALES and the OOMYCETALES.
ORDER ZYGOMYCETALES
The fungi of this order show a strongly developed mycelium con-
sisting usually of unicellular, sometimes pluricellular, multinucleate
hyphas. These hyphae are distinguished in the typic forms as the rhiz-
oidal hyphae, aerial hyphae and reproductive hyphae. Vegetative re-
1 Massee, George: Text-book of Fungi, 1906: 242.
92
MOULD FUNGI 93
production is never through motile zoospores, but through immotile

spores produced in sporangia borne at the tips of the reproductive
hyphae known as sporangiophores, or by means of conidiospores,
chlamydospores (Mucor racemosus), oidiospores, or gemmae. Sexual
reproduction is by the conjugation
two similar or sHghtly dissimilar
of
gametes, and the formation of a resting cell, or sexually produced spore,
known as the zygote, or zygospore. Brefeld beheved that this group
gave rise to the higher groups of fungi and he showed an interesting
series of transition forms from those like Mucor with a typic terminal
sporangium (Fig. 13) with numerous sporangiospores (endospores)
through Thamnidium elegans with a large terminal sporangium (mega-
sporangium) and secondary lateral smaller sporangia (sporangioles,
microsporangia) and Thamnidium chcetocladioides (Fig. 32), where the
absent terminal megasporangium is represented by a spine-like sporangi-
phore, to Chcetodadium, where the number of endospores in the spor-
angioles is reduced to one inclosed within the sporangium, which be-
haves as a conidiospore; thence to Piptocephalis, where the monosporous
sporangiole has become virtually a conidium, or conidiospore. He re-
garded the ascus as potentially a sporangium, but recent discoveries
have shown this hypothetic view to be untenable, so that his views as
to the origin of the ASCOMYCETALES and the BASIDIOMYCE-
TALES from the ZYGOMYCETALES must be considered as not satis-
factorily proved.
Blakeslee, who has studied the sexual reproduction in the moulds,
finds that they may be divided into two groups, the homothallic (mon-
oecious) and the heterothallic (dioecious) forms. The homothallic
moulds are those in which the sexual gametes, which conjugate, arise
from the same mycelium, while the heterothallic forms are those in
which two distinct mycelia contribute the gametes which ultimately
unite sexually. The homothallic (hermaphroditic) moulds he divides
into the heterogamic hermaphrodites in an inequahty in
which there is
the size of the gametes (the large one being female and the small one
male), and the homogamic hermaphrodites in which the gametes are of
equal size. The heterogamic hermaphrodites include the following
fungi Syncephalis, Dicranophora fulva, A bsidia spinosa, Zygorhynchus
:
heterogamiis, Z. Mcelleri, Z. Vuillemini. The homogamic hermaph-

rodites comprise: Mortierella polycephala, Mucor genevensis, Spinel! us
fusiger and Sporodinia grandis (Fig. 28). The dioecious, or hetero-
94 MYCOLOGY
thallic species are all homogamic, that is, there is no difference in the
size of the two gametes which conjugate. This group includes such
Fig. 28. — Zygospore formation in Sporodinia grandis from material growing on toad-
stool. {Slide prepared by H. H. York, Cold Spring Harbor, July 29, 1915.)
)H=*^/
\=9c>i
^îj=f /'
Fig. 29. — Conjugation and development of zygospores between + and — races of

black mould, Rhizopus nigricans.
fungi as Absidia ccerulea, Mucor mucedo, and five other forms of Mucor,
Phycomyces nitens and Rhizopus nigricans (Fig. 29). Taking the con-
MOULD FUNGI 95
jugation in Mucor mucedo as an illustration of the method, we find that

the hyphae of two distinct mycelia, which may be designated as the
+ and — strains, give rise to lateral club-shaped branches. The tips
of these two branches (progametes) come into contact and a terminal
cell (gamete) is cut oflF from each branch respectively by a transverse
wall. The double partition wall is dissolved away by an enzyme, and

the two cells coalesce, their nuclei uniting in pairs. A zygospore, is
formed, as a resting spore (Figs. 28, 30 and 2,i)- It becomes covered
with a thick, warty brown coat. The zygote (zygospore) germinates
after a period of rest producing at once, because of the concentrated
foods it contains, a sporangiophore bearing a terminal sporangium with
sporangiospores. Sometimes the gametes fail to unite through some
check to the normal conjugation and the two gametes may then round
off and form and the size of these azygospores
thick- walled azygospores,
depends upon the gametes from which they develop. Blakes-
size of the
lee has discovered that for the production of zygospores in heterothallic

moulds the contact of the hyphae of two distinct mycelia designated
+ and — are essential. If two — races or two -|- races meet, there
is no result. In the homothallic moulds, the two conjugating gametes
may arise from the same mycelium. Where the -f- race of one species
of mould meets the — race of another species imperfect "hybrids" are
formed. The testing out, maleness or femaleness, of the different races
is made by growing in proximity different kinds of moulds,
possible
where a reaction occurs and imperfect hybrids are formed one race
must be plus and the other minus. Where the hermaphrodite forms are
grown, it is noticed that one gamete is larger and the other smaller, and
it is assumed, that the larger gamete is female and the smaller one male.
The race of dioecious Mucors, designated tentatively (+), shows a
sexual reaction with the smaller or male gamete, while the ( — ) or
vegetatively less vigorous race shows a reaction with the larger or
female gamete. It is inferred that the -f race of dioecious mucors is
female and the — race, male.

The immediate stimulus to the formation of the progametes prob-
ably lies in the contact of hyphae from different strains through the
osmotic activity of the hyphal contents. For this reason progametes
fail to form in relatively dry air. By suspending two small bags filled
with bread soaked in dilute orange juice and inoculated with mould
spores, any influence which the substratum might show is eliminated.
:
96 MYCOLOGY
Zygospores were formed in one week where the aerial radiating hyphae
had come into contact. By this experiment all influences exerted
through the solid culture media, or which were due to contact of vege-
tative mycelia, were eliminated.
The sporangia of Mucor mucedo areraised upon the ends of sporangio-
phores. When formed the sporangium consists of a wall beset
fully
with spicules of calcium oxalate, the spores separated from each other
by a slimy intersporal substance (zwischensubstanz), and a columella
which projects into the interior of the sporangium. The formation of
spores in Rhizopus nigricans and Phycomyces nitens has been studied by
Swingle,^ who finds that the columella is formed by the cutting upward
of a circular surface furrow or cleft, thus cleaving out the columella over
the end of which a plasma membrane is formed. The spore plasm of
Rhizopus divides into spores by furrows pushing progressively inward
from the surface and outward from the columella cleft both systems
branching, curving and intersecting to form multinucleated bits of pro-
toplasm (the spores) surrounded only by plasma membranes, which
become the spore walls and separated by spaces filled with the inter-
sporal substance (zwischen substanz). The endospores, or sporangio-
spores, of Rhizopus nigricans and Sporodinia grandis are multinucleate,
while those of Pilobolus are binucleate, according to Harper. The es-
cape of the mature sporangiospores takes place when a portion of the

sporangial wall is dissolved. The spores escape imbedded in the inter-
sporal slime, which dries up liberating the spores. Certain species of
Mucor are capable of fermenting grape juice, the power of fermentation
depending on the species. The following species produce alcoholic fer-
mentation (Lindner)
Quantity of alcohol
Species by volume,
per cent.
Mucor Jansscni 3.41

Mucor lamprosporus 3.71
Mucor javanicus 2 83
.
Mucor phimbeus 4.62

Mucor pirelloides i 06
•
Mucor racemosus 462

Mucor Rotixianus 5.25
Mucor griseo-cyanus 4 00
.
Mucor genevcnsis 5-2i
'Swingle, DeanB.: Formation of the Spores in the Sporangia of Rhizopus

nigricans and of Phycomyces nitens. Bull, jy, Bureau of Plant Industry, 1903.
MOULD FUNGI 97
Key to Families or the Order Zygomycetales

Non-sexual spores in sporangia, which in some genera are reduced
to conidioid bodies.
A. Non-sexual spores formed in sporangia in many cases accom-
panied by conidiospores.
(a) Sporangia (at least the main sporangia) with columella.
Conidiospores absent, or only sparingly found. Zygospores
naked, or only covered by curled outgrowths of the sus-
pensors. I. Mucorace^.
(b) Sporangia without columella; zygospore surrounded by a
thick covering of hyphae. II. Mortierellace^.
B. Non-sexual spores as conidiospores. Sporangia exceptionally
present.
(a) Conidiospores single. Zygospores formed directly by the
united gametes.
1. Sporangia present transitional to conidia; sporangia
monosporic and polysporic. III. Choanephorace^.
2. Sporangia never present; parasitic on other MUCOR-
ALES. IV. Ch^tocladiace^.
{b) Conidia in chains zygospore formed where the bent ends
of the gametes unite. V. Piptocephalidace^.
Non-sexual spores as true conidiospores borne singly at the end of

conidiophores. VI. Entomophorace^.
Family i. —
Mucorace^. The mycelium of the true moulds is
homogeneous, or it becomes heterogeneous through differentiation into

aerial and nutritive hyphae. Non-sexual reproduction by the forma-
tion of endospores in sporangia. The sporangia here may be simple or
branched. The sporangia are all alike, or there are as in Thamnidium
two different types known as megasporangia and microsporangia. The
larger sporangia have a columella, while the smaller ones are mostly
without a columella, but occasionally a columella is present. The
formation of conidiospores is unknown in the family. The zygospore
may arise by the fusion of two similar gametes formed from the same
mycelium (homogamic hermaphrodites) or by the union of two slightly
dissimilar gametes the product of the same myceUum (heterogamic her-
mophrodites), or it arises by the conjugation of similar gametes (+ and
— races) from two distinct mycelia (heterothallic and homogamic).
98 MYCOLOGY
The important genera of the family are Miicor, Rhizopus, Phycomy-

ces, Ahsidia, Sporodinia, Thamnidium, Dicranophora, Filaira and Pilo-
boliis.The genus Mucor, a key for the identification of the species will
be given at the end of the book, was estabhshed in 1729 by Micheli.
The genus may be divided into three groups of species. The first
division includes those species with unbranched sporangiophores, such
as Mucor mucedo. The second group comprises the moulds with clus-
tered branches of the sporangiophores, as Mucor corymbifer, M. erectus,
M. fragilis, M. pusillus, M. racemosus, and M. tenuis. The third sec-
tion is made up of species the sporangiophores of which show sympodial
Fig. 30. — Details of Chlamydomucor racemosus showing oidia, sporangia and zygo-
spore formation.
branching. Such are Mucor alternans, M. circinelloides, M. javanicus,

M. Rouxii and M. spinosus. (Also consult pages 695-702.)
The oldest known species, Mucor mucedo, was described fully for the
first time by O. Brefeld in 1872. Stiff sporangiophores, 30 to 401J,
thick, arisefrom the mycelium and are 2 to 15 Each

cm. in height.
bears a single globular sporangium 100 to 200/i in diameter and the
sporangial wall is beset with fine needles of calcium oxalate. The spores
are ellipsoidal 3 to 6/i by 6 to 12/i with faint yellowish cell contents. As
previously described, conjugation is between two similar gametes from
+ and — mycelia. Mucor racemosus, also known as Chlamydomucor
MOULD FUNGI 99
racemosus (Fig. 30), shows the clustered branching of the sporangio-

phore and in addition the hyphae are marked by the intercalary forma-
tion of chlamydospores. This mould produces sporangiophores 8 to 20/^
thick by 5 to 40 mm. in height, bearing brownish sporangia 20 to 70/x in
diameter. The globular colorless spores are 5 to ?>^x broad by 6 to lo/x
in length. This mould which grows on bread and decaying vegetable

matter, and if cultivated submerged in beer-wort, the hyphae swell ir-
regularly and a large number which divide

of transverse septa appear,
the hyphae into barrel-shaped portions. These cells or gemmae can be
separated readily, and when free, they become spheric and multiply by
budding, as in the true yeasts, and the submerged spores hiso bud and
constitute the so-called Mucor-yeast. At the surface of the liquid, they
develop the typic mould form. Miicor racemosus, according to Hansen,
is the only mould capable of inverting cane-sugar solution. It produces
in beerwort as much as 7 per cent, by volume of alcohol. Mucor erectus,
which grows on decaying potatoes, produces azygospores as well as zygo-
spores. It has the same appearance as the preceding and possesses an
active power of fermentation. In beer-wort of ordinary concentration,
it yields up to 8 per cent, by volume of alcohol, and in dextrin solutions
it induces alcoholic fermentation. Mucor spinescens, which grows on

Brazil nuts, has spiny projections on the rounded upper surface of the
columella. Mucor (Amylomyces) Rouxii occurs in the so-called
" Chinese yeast," which is in the form of small whitish cakes, consisting
of rice grains kneaded together with assorted spices. These cakes are
powdered and mixed with boiled rice upon which the mycelium grows,
converting the rice by slow degrees into a yellowish liquid which con-
tains glucose produced by the diastatic ferment of the fungus.
The black mould Rhizopus nigricans {Mucor stolonifer) grows on
bread and other organic substrata (Fig. 31). Several sporangiophores
arise from a single point of origin, namely, at the top of a mass of rooting
(rhizoidal) hyphae which constitute an adhesive organ or oppressorium.
Each erect stalk bears oblate spheroidal sporangia with distinct colu-
mella and sporangiospores, 6 to 17/i long. Arising from the base of the
clustered sporangiophore is a horizontal hyphae, which often attains a
length of 3 cm. and is known as the stolon, or stoloniferous hypha.
When the tip of this stolon comes into contact with the substratum a
new appressorium is formed from which arises a number of sporangio-
phores bearing sporangia (Fig. 31). This method of growth enables the
MYCOLOGY
black mould to spread rapidly andit sometimes chokes out other moulds
growing in on the same nutritive medium. In 1818,

competition with it
on account of this method of growth, it was named by Ehrenberg

Mucor stolonifer. Related to this fungus is one named Rhizopus ory-
zecB which grows in Ragi. The fungus Phycomyces nitens is found in
empty oil casks, on oil cakes and in concentrated fodder. It puts forth
sporangiophores 7 to 30 cm. long and 50 to 150/i in diameter which
stifif
bear at the summit black globular sporangia 0.25 to i.o mm. in diame-
ter, filled with yellow-brown, thick- walled endospores, 16 to 30/i
long and 8 to 15/x broad. Its zygospores are 300/x broad and their
Fig. 31. —
Black Mould, Rhizopus nigricans. A, Mature plant showing rhizoidal
hyphse {myc)\ stoloniferous hypha {st); sporangiophores (sph); sporangia (sp). B,
Younger cluster of sporangiophores and sporangia. (After Gager.)
borders are covered with many forked projecting hyphae known as

suspensoria. Recently H. Burgeff^ has studied the variability, sex-
uality and heredity of Phycomyces nitens and has brought his cultural
investigations into line with the recent developments of cytology and
genetics. His paper should be read by all students, who may be
interested in the extension of the methods of genetics into an
investigation of the lower plants.
The genus Absidia includes five species. In these fungi the suspen*-
sors are borne at the base of the two gamete cells which fuse to form the
zygospore, which when mature is covered by a basket-like covering of
1 BuRGEFF, H.: Untersuchungen liber Variabilitat,. Sexualitat und Erblichkeit
h&iPhycomyces nitens Kuntze. Flora, Band 108: 353-448; review by G. V. Ubisch

(Dahlem) in Botanisches Centralblatt, Band 128, Nr. 23: 630-632, 1915.
;
MOULD FUNGI
Straight appressoria, which hook together by their curved extremities,

thus giving additional protection to the zygospore. Sporodinia grandis,
the single species of another genus, lives on large fleshy fungi of the
Fig. 32. -Sporangia of i, Thamnidium elegans; 2, 3, 4, Thamnidium chcelodadioides

5, Chalocladium Jonesii. (After Brefeld.)
Agaricace^, Boletace^, Clavariace^ and Hy-

families (Fig. 28)
DNACE^. Its sporangiophores i to 3 cm. high are finally brown in color
and dichotomously branched. The .sporangia are spheric with a deli-
MYCOLOGY
cate sporangial wall, which soon disappears leaving the spores on a

hemispheric columella. These spores are ii to 70/x broad. The 300/x
broad zygospores are produced from similar branches of a dichotomously
branched zygosphore. The mycelium of the species of Thamnidium
enters the nutritive substratum. The large sporangia are terminal
while the smaller secondary sporangia are borne on lateral branches in
whorls below the terminal sporangium. This is typically seen in Th.
Fig. ^^.— Details of sporangia and sporangiophores of Pilobulus. i, P. tnicro-
sporus; 2, P. roridus; 3, 4, 5, P. anomalus; 6, zygospore of P. anomalus. {After

Brefeld.)
elegans (Fig. 32). A related species Th. Fresenii has an upright termi-
nal sporangiophore, which is either sterile, or ends in a large terminal
sporangium, while the smaller sporangia are as in Th. elegans. In
Th. amoenum, the lateral smaller sporangia are borne at the end of
coiled secondary sporangiophores. The secondary sporangia suffer
reduction in Th. ch(etocladioides (Fig. 32) which in addition to having a
straight terminal spine-like hypha in place of the terminal sporangia has
some of the lateral microsporangia replaced by sterile branches. The
MOULD FUNGI IO3
commonest is P. cryslaUinns which appears

species oi Pilobolus (Fig. 33)
on horse dung. few short feeding hyphae and an upright spor-
It has a
angiophore swollen at the extremity by gas and water vapor and, there-
fore, under tension. It bears at its extremity a flat rounded sporan-
gium filled with sporangiospores. An explosion of the sporangiophore
causes the whole sporangicum to be shot off a considerable distance.
Family 2. —
Mortierellace^. This family consists of two genera
Mortierella and Herpocladiella. The genus Mortierella, which is repre-
sented by a coprophilous species, M. Rostafinski, has a sporangium
borne on a sporangiophore which arises in a definite way from a snare of
hyphae that are knotted into a rounded mass at its base. In M. cande-
labrum, the sporangiophore branched candelabra-like. Brefeld men-
is
tions Mortierella and Rhizopiis as examples of the carposporangiate

ZYGOMYCETALES, where the sporangiophores appear always at
predetermined places on the mycelium, and not at indefinite points, as
in the majority of other moulds.
—
Family 3. Choanephorace^. Represented by a single genus
Choanephora and a single species infundibulifer on flowers of Hibiscus
in East Indies.
—
Family 4. Chcetocladiace^. This is a small family of one
genus {Chcetocladium) and two species, {Ch. Jonesii and Ch. Brefeldii)
(Fig. 32) which live parasitically on Mucor mucedo and Rhizopus nigri-
cans. The terminal sporangia of Thamnidium are never formed and
secondary sporangia are reduced to the unisporous condition suggesting
conidiospores with pointed branches between them.
Family 5. Piptocephalidac§.«. —Three genera Piptocephalis,
Syncephalis -and Syncephalastrum are recognized in Die Natiirlichen
PflanzenfamiHen. The eight species of Piptocephalis are parasitic
on the mycelia Mucor, Pilobolus and Chcetocladium species (Fig. 37).
of
The haustorial hypha flattens itself disc-like on the outer surface of
the host's hyphge and sends five rhizoida'l branches into the host cells.
An erect dichotomously branched conidiophore bears conidiospores
in globular clusters at the ends of its principal branches. Some
species of Syncephalis areparasitic on other fungi; but S. cordata
grows on manure, presumably as a saprophyte.
—
Family 6. Entomophthorace.e. The mycehum of the fungi of this
familyis more or less richly developed and fives endozoically in animals,
such as flies, mosquitoes, aphids, and seldom saprophytically as

I04 MYCOLOGY
Basidioboliis on the feces of frogs. Non-sexual reproductionsis mainly
by means of unicellular conidiospores which are discharged forcibly

from the ends of tubular conidiophores. Sexual reproduction is by the
conjugation of two gametes dissimilar in size, heterogamic and thus these
fungi connect the ZYGOMYCETALES with the OOMYCETALES
where oogamous reproduction is displayed. The zygospores formed
in conjugation are spheric, while the azygospores formed on the
mycelium without copulation are similar to the zygospores in struc-
ture and appearance. The family includes seven genera, includ-
FiG. 34. — Fly cholera fungus (Empusa musca). i, Fly enveloped in mycelium;
2,fungus between hairs of the fly; 3, conidiophores and conidiospores; 4, germina-

tion of spores; 5, formation of egg in Empusa sepulchralis. {After Thaxter.) See
Henri Coupin, Atlas des Champignons, Parasite set Pathogenes de 1' Homme et des
Animaux, 1909.
ing Empusa and Entomophthora, which may be chosen as types for

discussion.
The mycehum of Empusa musccB (Fig. 34) is parasitic in the
bodies of flies, destroying them in large numbers by an epidemic in the
fall, known The short hyphae frequently bud like
as fly-cholera.
yeast The conidiophores break through to the surface of the
cells.
insect's body, where the conidiospores 18 to 25/x broad by 20 to 30/^

long are forcibly discharged. These spores bore their way through the
chitinous covering of a healthy fly by means of a germ tube and the
MOULD FUNGI 105
hyphffi which enter the body of the tly bud hke yeast cells, which are
carried to all parts of the insect's body. Later the parasitic hyphse
arisefrom the gemmae. Resting spores are unknown.
Entomophthora is a genus of fungi inclusive of thirty species found
on various insects in Europe and North America. Entomophthora
sphcerosperma has a richly branched nutritive mycelium, which grows
through the body of insects. After the death of the host, the hyphae
break through the surface in connected strands part of which attach
the larva, or insect's dead body, to the substratum and part form a
thick white mantle over the surface.
The conidiophores are in branching bundles. The conidiospores
are elongated ellipsoidal, 5 to 8ju broad by 15 to 26ju long. Secondary
and tertiary conidia are found. The resting spores produced as azy-
gospores are spheric and 20 to 35^1^ broad with a smooth yellow wall.
It grows on larvae, especially frequent on the cabbage worm Pieris
brasskce in Europe and North America.
BIBLIOGRAPHY OF THE ZYGOMYCETALES

This is not intended to be a complete list of theworks dealing in whole or in
part with the mould fungi, but only a list of the works which may prove helpful to
the student of mycology.
Rainier, G.: Etude sur les Zygospores des Mucorinees, These pr^sentê a I'l^cole
de Pharmacie. Paris, pp. 136, pis. i-ii; Observations sur les Mucorinee.
Annales des Sciences naturelles, ser. 6, 1-15: 70-104, pis. 4-6. Sur les zygo-
spores des Mucorinees. Annales des Science naturelles, vi ser., I: 18, 1883;
Nouvelles observations sur les zygospores des Mucorinees, do., I: ig, 1884.
Blakeslee, Albert F. Sexual Reproduction in the Mucorineae. Proceedings

:
American Academy "Arts and Sciences, xl: 205-319 with 4' plates and bibli-
ography, The Biological Significance and Control of Sex. Science, new ser.
x.xv: 366-384, March 8, 1907; Papers on Mucors (a review). Botanical Gazette,
47: 418-423, May, 1909; Heterothallism in Bread Monld, Rhizopus nigricans.
Botanical Gazette, 43: 415-418, June, 1907; A Possible Means of Identifying
the Sex of (+) and ( — ) Races in the Mucors. Science, new ser. xxxvii: 880-
881,June 6, 1913; On the Occurrence of a Toxin in Juice Expressed from the
Bread Mould, Rhizopus nigricans. Biochemical Bulletin II: 542-544, July,
1913; Conjugation in the Heterogamic Genus Zygorkynchiis. Mycologisches
Centralblatt II: 241-244, 1913; Sexual Reactions between Hermaphroditic and
Dioecious Mucors. Biological Bull., xxix: 87-102, August, 1915; Zygospores
and Rhizopus for Class Use. Science, new ser. xlii: 768-770, Nov. 26, 1915.
Brefeld, O.: Botanische Untersuchungen liber Schimmelpilze, Heft i, Zygomy-
ceten, pp. 1-64, Taf, 1-6, 1872; Untersuchungen aus den Gesamtgebiete der
Mykologie, ix, 1891.
Io6 MYCOLOGY
Buchanan, Estelle 1)., and Buchanan, Robert E.: Household Bacteriology,

1914: 66-72.
DE Bary, a.: Comparative Morphology and Biology of the Fungi and Bacteria,
1887: 144-160.
Englee, a. and Gilg, Ernst.: Syllabus der Pflanzenfamilien, 7th Edition, 191 2:
37-38.
Gortner, Ross A. and Blakeslee, A. F. Observations on the Toxin of Rhizopus
:
nigricans. American Journal of Physiology, xxxiv: 354-367, July, 1914.

Jorgensen, Alfred: Microorganisms and Fermentation, 3d Edition, 1900: 97-
115-
Keene, Mary L.: Cytological Studies of the Zygospores of Spordinia grandis.
Annals Botany, xxxviii: 455, 1914.
of
Klocker, Alb.: Fermentation Organisms, 1903: 170-186.
Lafar, Franz: Technical Mycology, II, part i: 1-30, 1903.
Lendner, Alf.: Les Mucorinees de la Suisse. Materiaux pour la Flore Crypto-
gamique Suisse III, Ease, i: 1-177, 1908.
Schroter, J.: Mucorineae, Die natlirlichen Pflanzenfamilien, I, Teil i. Abt. 119-
142, 1897.
Stevens, F. L.: The Fungi which Cause Plant Disease, 1913: 101-108.
Swingle, Deane B.: Formation of the Spores in the Sporangia of Rhizopus nigri-
cans and of Phycomyces nitens. Bureau of Plant Industry No. 37, 1903 with
6 plates.
Underwood, Lucien M.: Moulds, Mildews and Mushrooms, 1899: 24-28.
VON Tavel, F.: Vergleichende Morphologic der Pilze, 1892: 25-40.
Wettstein, Richard R. von: Handbuch der systematischen Botanik, 1911: 160-
164.
CHAPTER XII
OOSPORE-PRODUCING ALGAL FUNGI

ORDER II. OOMYCETALES
The fungi of this order were derived probably from some ancestor,
or ancestors, which through the loss of chlorophyll became dependent
on extraneous supplies of organic food. If we look for such an ancestral
form among the algae, we find that it must have been related to Vau-
cheria, if not identic with that filamentous siphonaceous green alga
with reproductive organs, as oogonia and antheridia. Vaucheria is a
unicellular filamentous sparingly branched cell with a thin cell wall and
multinucleate. Hence it is sometimes called a coenocyte. Similarly,
the structural features of the more primitive Oomycetales are like
Vaucheria, but the absence of chlorophyll is distinctive. The forma-
tion of non-sexual sporangia with the formation of zoospores, or swarm
spores, known as zoosporangia is a feature of the fungi of this order.
As there is a pronounced difference between the male and female sexual
organs, oogamous reproduction is the rule. The oogonium is compara-
tively large and contains one or more oospheres, which are fertilized
by the sperm cell, which swim to it by cilia, creep to it, or are carried
into the oogonium through a fertilization tube. Sexual reproduction in
these fungi has been investigated cytologically by a number of students,
and they have found that the nuclear changes concomitant with fertili-
zation are characteristic. Albugo Candida, A. lepigoni, Peronospora
parasitica, Plasmopara, Pythium and Scleras para show a single large cen-
tral oosphere with a single nucleus, while the remaining nuclei pass from
the gonoplasm into the periplasm. A process is sent into the oogonium
from the antheridium and a single male nucleus passes into the oogo-
nium. A cell wall is developed about the oospheres and the male and
female nuclei unite, while the periplasm is used in the formation of the
spore wall (episporium). The ripe oospore has a single nucleus in
Peronospora parasitica, while in Albugo, it becomes multinucleate after
nuclear division. A central oosphere (gonoplasm) surrounded by peri-
107
Io8 MYCOLOGY
plasm occurs in Albugo blitl and .1. portulacce and the oosphere is
multinucleate and the' nuclei present fuse in pairs with a number of

sperm nuclei which enter from the antheridium. The oospore which
arises is multinucleate. This method is considered by mycologists to
be the primitive one as displayed in these two species, the uninucleate
oospheres of the first-named species having been derived from the multi-
nucleate. An intermediate position is occupied by Albugo tragopogonis,
where at first the oosphere is multinucleate but by the degeneration of
all but one female nucleus becomes uninucleate. Claussen^ finds that
Sa'prolegnia monoica develops both antheridia and oogonia, the latter
at first being filled with protoplasm and many nuclei which wander to
the periphery and undergo degeneration with a few nuclei left over.
These nuclei divide once mitotically. Around these daughter nuclei
the protoplasm collects to form the egg cells. Each egg has a single
nucleus near which is the coenocentrum of Davis, but which Claussen
thinks is a true centrosome. The simple or branched antheridia form
germ tubes which enter the wall of the oogonium and a single male
nucleus fuses with the nuclei of the egg cells to form the oospore.
Claussen contrasts the life cycle, as determined by his investigations,
with those of Trow in the following diagrammatic presentation:
Trow Diploid Haploid Diploid
/ Antheridium — Male nucleus \

/ \
Oospore, Mycelium Oospore
Multinucleate \ /
\Oogonium— Egg cell — Egg nucleus/^
Claussen Haploid Diploid.
Strasburger considers that the superfluous nuclei in the oogonia and

the antheridia are comparable with the superfluous egg nuclei of certain
of the brown seaweeds belonging to the family Fucace^. When the
oospores germinate, they either produce directly a mycelium, or give
rise to zoospores. The fungi of this order are essentially parasitic,
being found in this condition as endophytic parasites, or as endozoic
parasites on fishes and insects and Euglena.
1 Claussen, P.: Ueber Eintwicklung und Befructung bei Saprolegnia monoica.
Festschrift der Deutsch. Bot. Gesellsch., xxvi; 144-161 with 2 plates, 1908.
OOSPORE-PRODUCING ALGAL FUNGI IO9
Key to Families of the Order Oomycetales
A. Zoosporangia, oogonia and antheridia present; conidia absent.

(a) Mycelium well developed.
1. Antheridium forming motile spermatozoids, which enter the
oogonium. Family i. Monoblepharidace^.
2. Antheridium not forming spermatozoids, fertilization through
an antheridial tube, or beak. Family 2. Saprolegniace^.

(b) Mycelium poorly developed, sometimes represented by a single
cell.
1. Fruit body as a single cell or by division forming a sporangial

sorus; parasites on algae, protozoans, rarely on flowering
plants. Family 4. Chytridiace^.
2. Fruit body through division a chain of cells which develop
sometimes into zoosporangia, sometimes into antheridia and
oogonia. Family 5. Ancyclistace^.
B. Conidia present. Family 3. PERONOSPORACEiE.
The following descriptions of the above five families are presented

in order to introduce the student to the characters which fundament-
ally distinguish them. Therefore, all generic keys are omitted because
the introduction of them under each family would increase the size of
the book unduly.
Family i. Monoblepharidace^. —This family is represented by
the genera M onoblepharis and Gonapodya. The genus Monoblepharis
is represented by two species of which M. sphcBrica is the most com-
mon. an aquatic fungus found growing saprophytically on dead
It is
animal and plant parts under water. The hyphae of the mycehum
are tubular, branched and unicellular. The swarm spores (zoospores),
which are formed much as in Saprolegnia, have only a single flagellum.
The oogonia are either terminal in position or interstitial and there is
no differentiation of an outer periplasm, but the whole protoplasm
of the oogonium contracts to form an oosphere. Later a pore appears
at the apex of the oogonium through which the uniciliate spermatozoids
enter to fertilize the egg cell. The antheridium in M. sphcBrica appears
as a penultimate cell immediately below the oogonia. An opening is
formed at the top through which the spermatozoids escape. The
oosphere on fertilization becomes an oospore. Because of the aquatic
no MYCOLOGY
habit and formation of motile spermatozoids, Brefeld considers Mono-

blepharis to be the most primitive of the Oomycetales.
—
Family 2. Saprolegniace^. The members of this family, as
their name indicates, are saprophytes on both dead plants and animals
in water with the exception of the fungus which causes the salmon
disease and it is both a saprophyte and a facultative parasite. The
hyphae in the vegetative condition are relatively large, arising from
delicate rhizoids which penetrate the substratum. Swarm spores
which are biciliate are formed in terminal, long, tubular zoosporangia
opening by an apical pore through which the zoospores crowd their
way out into the water. Sometimes, as they escape, they collect into
ball-shaped masses which are caused to slowly about by the activity
roll
of the cilia. The female sexual organs, the oogonia, are terminal on
the branches of the thallus hyphae. Several oospheres without dis-
tinction of periplasm are formed inside of a single oogonium, and
sometimes, as many as thirty or forty are found. The antheridia,
which are club-shaped, are formed on slender branches of the mycelium
which also bear the oogonia, or which are distinct from those
which are oogonial bearers. These antheridia approach the oogonia
and an antheridial beak is formed which penetrates the wall of the
oogonium and comes into contact with the oospheres by growing from
one oosphere to another. Sometimes the antheridia, as in Saprolegnia
monilifera, are not produced at all and the oogonia develop partheno-
genetic oospores which germinate after a rest period of a few days to
several months. The series representing reduction in sexuality begins
with such forms as Saprolegnia monoica with an oogonium and an
antheridium which develops a fertihzing process through Achlya
polyandra, which forms antheridial branches which do not touch
the oogonia, to Saprolegnia monilifera without any trace of antheridia.
Androgynous forms are those in which the same hyphal branch
develops both antheridia and oogonia and the diclinous species
like Saprolegnia dioica and Achlya oblongata ar.e those in which the
antheridia and oogonia are borne on distinct branches.
Saprolegnia ferax usually attacks only fishes, tadpoles and the
spawn of frogs. It appears on aquarium-kept fishes on the sides of
the body at the tail end, or among the gills. In the latter place, if
abundant, it frequently causes asphyxiation and before this state is
final the fish turns over on its back and rises to the surface. In the
OOSPORE-PRODUCING ALGAL FUNGI III
experience of the writer, immersion of the diseased fish in strong brine

in many cases brings about a cure, if the growth of the fungus is not
too great, observed a sick bream in the lake of Fure So
Petersen'^
with a wound quite overgrown with Saprolegnia hyphge and he has
found frog eggs which were attacked, the hyphae growing in the jelly
around the eggs, penetrating into them. The fungus can be raised in
the laboratory on dead fishes by allowing tap water to slowly flow over
them in a jar. A few days are necessary to secure a copious growth.
Frogs which die under the ice in winter for lack of oxygen float to the
surface in the spring entirely covered by this fungus. It thrives
best in the early stages of decay, for as putrefaction advances bacteria
and an extent as to check the growth of the
infusoria increase to such
fungus. When air insects, fall into lake or pond water
such as gnats,
in great numbers, species of Saprolegnia, Achyla and Aphanomyces
appear in great numbers and seem to form a gray felt on the surface.
The vegetable materials on which the Saprolegniace^ mostly Hve
are branches and shoots of trees, except Salix, owing to presence of
salicin, which fall into the water. Second in importance are half-
rotten rhizomes of Calla, half-rotten leaves and leaf stalks of Nuphar
and Nyniphcea and other parts of aquatic plants which float on the
surface. Species of the genus Achlya are mostly associated with such
materials. Achlya polyandra have been repeatedly found by me on the
fruits ofOsage oranges which have fallen into the pond at the Univer-
sity of Pennsylvania. The most favorable environmental conditions
seem to be the absence of air about the hyphae, quiet, still, pure water,
that does not contain much iron and a relatively open light surface.
Low temperature conduces to the formation of oogonia, which also
keeps in check other competing organisms (Fig. 35).
—
Family 3. Peronosporace^. This family is rich in parasitic
forms which may be accounted as the cause of important diseases of
cultivated plants. The hyphae of the mycelia are irregularly and copi-
ously branched and are found mainly in the intercellular spaces of the
host tissue sending short branches called haustoria into the adjoining
living cells. These haustoria maybe glohulsLV (Albugo = Cystopus),
club-shaped (Peronospora corydalis), branched (Plasmopara) (Fig. 36),
or branched and snarled {Peronospora). Septa are absent except
1 Petersen, Henning E. : An Account of Danish Fresh-water Phycomycetes.
Annales Mycologici, viii, No. 5, 1910.
112 MYCOLOGY
when the reproductive organs are formed. Non-sexual spores, or

conidiospores, are borne on conidiophores which may remain within
the host (Albugo = Cystopus), or grow beyond the surface. They may
be either simple or branched. These conidiospores either germinate,
as in Phytophthora infestans and Peronospora nivea by means of zoo-
Fig. 35. —
I, Zoosporangium of Achlya racemosa; 2, escape of zoospores; 3, fly-
covered by mycelium; 4, zoospores of fungus; 5, Achlya ferax with zoosporangia and
zoospores; 6, Achlya proUfera, 24 hours after germination of zoospores. 7, Achlya
monoica, with antheridia and oogonia; 8, Achlya conlorta. {After Henri Coupin,
Atlas des Champignons Parasites el Pathogenes de I' Homme et des Animaux, pi. xviii,
1909.)
spores which escape or by the protoplasm escaping {plasmato parous), as

in Peronospora densa, or by germ tubes, which in some species {Perono-
spora lactuca) appear at the end of the spore (acroblasfic) , or at the
side of the conidiospore (pleuroblaslic) , as in Peronospora radii. The
oogonia and antheridia, which are also present, are formed in the
—
OOSPORE-PRODUCING ALGAL FUNGI 113
tissues of the host. The different kinds of nuclear fusion, which

accompany have been described previously. The oospore,
fertiUzation,
which is formed, acts as a zoosporangium in some cases for it gives
rise to numerous spores; or in other cases it produces a germ tube.
In most of the forms, the oogonium contains a mass of protoplasm

known as the oosphere. This is divisible into an outer clearer por-
FiG. 36. Plasmopora vilicola. A, Conidiophore with conidiospores (nearby

oospores); B, Haustoria; C, Swarmspore formation. A, 950/1; B. C, 600/1. {After
Millardcl in Die naliirlichen Pflanzenfamilien I. i, p. 115),
tion, the periplasm, and a denser more granular central portion, the
gonoplasm. After fertilization, the oospore develops a thick wall of
two layers, an extine and intine, and becomes a resting spore. It
accumulates fatty substances, which are utilized when the spore
germinates in the spring after a long winter's rest. The family has
had many revisions and in order to simplify matters Pythium and
Albugo (Fig. 37), which are placed in separate families by some
8
.
114 MYCOLOGY
authors, are placed in the family Peronosporace^. Details of the

important forms which cause plant diseases will be given in the third
part of this book. These fungi will be referred to under each genus
following the systematic generic key which is here given.
Generic Key of the Family. PERONOSPORACEiE
Mycelium of these fungi parasitic or saprophytic in plant tissues;

zoosporangia as distinct organs producing biciliate zoospores.
Zoospores formed out of protoplaem
which escapes out of the conidia. i. Pythium.
Zoospores formed within the zoosporangia.
2. Pythiacystis.
Zoospores elongate. 3. N emato sporangium
^_^ Mycelial hyphae branching non-septate
/ ^ usually coarse, of strictly parasitic habit.

Conidiophores short, thick, subepidermal,
conidia in chains. 4. Albugo
Conidiophores longer superficial, simple
or branched, conidia not in chains.
Conidiophores scorpioid cymosely
branched conidiospores developing
swarmspores. 5. Phylophthora.^
Conidiophores simple, or branched
monopodially; conidia sprouting as a
plasma, or by swarm spores. Con-
Fig. 37. — White rust idiophores regularly branched.
Cystopus (Albugo) portula Conidiophores simple erect with a
cecB, on purslane, Portulaca
oleracea (Cold Spring swollen end (basidia-like) bearing
Harbor, L. I., July 24, short sterigma-like branches of equal
1915-)
length. 6. Basidiophora.
Conidiophores with lateral branches developed normally of
unequal length. Conidiophores stout, with few branches,
oospore united to wall of oogonium. 7. Sclerospora.
Conidiophores slender, freely branched persistent; oospore
free. [
8. Plasmopara.
Conidiophores with forking branches; conidiospores sprout-
ing with a germ tube. Upper end of conidiospore with a
. .. .
OOSPORE-PRODUCING ALGAL FUNGI 115
papilla through which the germ tube grows (acroblastic).

g. Bremia.
Conidiospores without papilla; pleuroblastic. 10. Peroiwspora.
The most important species of these genera from the standpoint of

the plant pathologist are the following enumerated below with their
common English names where such have been given.
English name Host plant
Pythium de Baryantim Damping-off fungus.. . . Seedlings

Pythiacystis citriopthora Brown rot of lemon.. . . Lemon fruits
Albugo {Cystopus) Candida... White rust of crucifers. Cruciferous plants
Albugo (Cystopus) portulaca; White rust of purslane. Portulaca oleracea
Phytophthora cactorum Mildew of succulents.. Cacti, etc.
Phytophthora infestans Late blight of potato.. Potato
Phytophthora phaseoU (Fig. Downy mildew of beans. Lima-bean
44)-
Plasmopara cubensis Downy mildew of cu- Cucumber
cumber.
Plasmopara Halstedii Helianthus annuiis and
H. tuberosus
Plasmopara viticola ... Downy mildew of grape. Grape vine
Bremia ladiica Downy mildew of lettuce. Cynara, Cineraria, Lactuca
Peronospora effusa Mildew of spinach Spinach
Peronospora parasitica Downy mildew of crucifers! Cabbage
Peronospara Schleideniana. . . Onion mildew Onion
CHAPTER XIII
OOMYCETALES (CONTINUED)
—
Family 4. Chytridiace.e. This family according to some authors
is made to include six families which are here reduced to six subfamilies.
It includes fungi of short vegetative duration, which may be a few days
in length. The swarm spores quickly give rise to new generations.
The resting period is represented in the case of the endophytic para-
sites by the time which elapses between the growth of two successive
crops of the host plants. The majority of the species of the family
are true parasites, partly endobiotic, partly epibiotic, and a few

are saprophytes. Half of the plant parasites live in fresh-water
algae, nearly as many in flowering plants, some of which are in
aquatic plants, some in swamp plants. About ten species are found
on marine algse. All species are microscopically small, yet they
cause galls, dwarfing, dropsy and crusts of the host plants. The
mycelium absent or in the form of slender protoplasmic filaments,
is
occasionally as distinct one-celled hyphae. The cell, which produces

the fruit body, frequently serves as the chief nutritive organ. Later,
it divides to form zoospores. mycelium has weak develop-
The true
ment. The short germ tube merely serves as an organ by which the
parasite gains entrance to the host cell, and in the endophytic forms, it
disappears quickly, but in the epiphytic species, it serves as an haustor-
ium, sometimes with rhizoidal extensions. In the better-developed
forms of Cladochytrie^, the slender mycelium serves to carry the
fungus from cell to cell of the host. The sporangia are always zoo-
sporangia which develop swarm spores, or zoospores. They are thin-
walled and quickly mature, or they are thick-walled and form resting
sporangia. Sexual spores are formed in only a few types and the differ-
ence between antheridia and oogonia is morphologically little pro-

nounced. The swarm spores have as a rule a single flagellum, rarely
do they have no such locomotory appendages. The sexually produced
oospores have the appearance of resting sporangia with the empty
antheridium attached as an appendage. Few of these fungi attack our
116
OOMYCETALES II7
cultivated plants, but where the attempt is made to grow alga^ and
other water plants, the fungi of this family occasionally do considerable
damage.
As an example of the first subfamily Olpide.e, may be chosen
Olpidiuni endogenum, which lives in the cells of desmids and kills
them. The zoosporangium found in desmid cells are oblate spheroids
and develop a long tube which projects out of the desmid cell through
which the zoospores with a single cilia escape into the water. O. ento-
phytum is parasitic in such filamentous algse as Vaucheria, Clado-
phora, Spirogyra. Olpidiopsis saprolegnm lives in the elongated
cells of Saprolegma,YiXod\xcmg enlargements in the hyphae of the fun-
gous host. The swarm spore bores a hole in the cell wall of its host
and swells out into a zoosporangium which develops a tube through
which the biciliate swarm spores escape into the water.
The subfamily Synchytrie^ includes most of the fungi which
attack the higher plants. Such are Synchytrimn decipiens on the
hog peanut (Amphicarpea monoica); S.fulgens on the evening primrose
{Oenothera biennis); S. stellarice on Stellaria; S. succiscB on Succisa
pratensis; S. taraxaci on dandelion; S. vaccinii causing a gall on cranber-
ries, Pycnochytriiini globosum on violet, wild strawberry, blackberry and
maple seedlings. P. myosotidis occurs on certain members of the

borage and rose families.
Cladochytrium tenue of the subfamily Cladochytrie^ lives in the
subaquatic tissues of the sweet flag, Acorus calamus, flag Iris
pseudacorus and a grass, Glyceria aquatica. Its mycelium is widely
distributed in the cells ofits hosts. Spheric sporangia 18 fx wide and
sometimes formed as intercalary enlargements of the mycelium,
66/x are
or they are formed at the end of the hyphae, with a colorless supporting
cell. They give rise to a short tube-like mouth which breaks out of
the host cell. The zoospores are uniciliate.
Representing the Oochytrie^ is an interesting fungus first fully
investigated by Nowakowski, namely, Polyphagus euglencE, which
attacks the cells of Euglena, a unicellular animal. Its mycehum con-
sists of a. central enlarged portion from which run out in a number of
directions branches which end in extremely fine points which penetrate
the cells of Euglena. The enlarged central portion develops a swollen
tubular outgrowth into which its protoplasm wanders. The contents
of this outgrowth then divide into numerous uniciUate swarm spores
Il8 MYCOLOGY
which escape into the water. Under certain conditions a cyst appears
in place of a zoosporangium. This is thick-walled and of a yellow
colorand enters a period of rest. After the rest period, the membrane
of the cyst rupture and a sporangium appears. Cysts may arise
by a kind of sexual union where two unlike mycelia fuse and the
protoplasm of both flows out to form a cyst between the original cells.
Urophlyctis pulposa attacks leaves and stems of Chenopodium and
A triplex species. U. alfalfcE grows in the roots of the alfalfa in
South America and Germany.
—
Family 5. Ancyclistace^. This is a small family consisting
of fungi whose mycelium is very sHghtly developed and not easily
distinguished from the fruit body. In one subfamily Lagenide^,
the mycelium is entirely absent. In the Ancycliste^, there is a
rich development of the mycelium which forms lateral tube-like
branches, which penetrate other cells. The fruit bodies are sac-hke
and give rise to zoospores. Sexual organs are present as antheridia
and oogonia, the contents of the former passing over completely into
the latter. The oospore, which is formed, is found free in the oogonium.
All of the known menxbers of this family are endophytic parasites and
the different stages of their development are short-lived.
Lagenidium entophytum lives in the zygospores of species of Spiro-
gyra. L. Rabenhorstii parasitizes the cells of Spirogyra, Mesocarpus,
Mougeotia. L. pygmatim lives in the pollen grains of diverse species
of Pinus.
BIBLIOGRAPHY OF OOMYCETALES
Atkinson, George F.: Damping Off. Bull 94, Cornell University Agricultural
Experiment Station, May, 1895; Notes on the Occurrence of Rhodochytrium
spilanthidis Lagerheim in North America. Science, new ser., xxviii: 691-692,
Nov. 13, 1908; Some Fungus Parasites of Algse. Botanical Gazette, xlviii:
321-338, November, 1894.
Clinton, G. P.: Oospores of Potato Blight, PhytopMhora hifeslans. Report Conn.
Agricultural Experiment Station. Part x. Biennial Report of 1909-1910:
753~774; Oospores of Potato Blight. Science, new ser., xxxiii: 744-747,
May 12, 1911.
Claxjssen, p.: Ueber Eientwicklung und Befructung bei Saprolegnia, monoica.
Ber. d. Deut. Bot. Gesellsch., xxvi: 144, 1908.
CoKER, W. C: Another New Achlya. Botanical^ Gazette, 50: 381-382, November,
1910.
Davis, Bradley M.: Cytological Studies on Saprolegnia and Vaucheria. The
American Naturalist, xlii: 616-620.
OOMYCETALES II9
DE Bary, a. : Compaialixe Morphology and Biology of the Fungi, Mycetozoa and

Bacteria, 1887: 132-145.
DuGGAR, Benjamin M.: Fungous Diseases of Plants, 1909: 135-173.
Engler, Adolf, and Gilg, Ernst: Syllabus der Pflanzenfamilien, 7th Edition,
191 2: 38-42.
Holder, Chas. F. : Methods of Combating Fungous Disease on Fishes. Bull.
of the Bureau of Fisheries, xxviii: 935-936.
Jones, L. R., Giddings, N. J. and Lutman, B. F.: Investigations of the Potato
Fungus, Phytophthora infestans. Bull. 168, Vermont Agricultural Experiment
Station, August, 191 2.
Kerner, Anton: The Natural History of Plants, 1895, ii: 668-672.
Magnus, P.: Kurze Bemerkung iiber Benennung und Verbreitung der Urophlyctis
bohemica. Centralblatt f. Bakteriologie, Parasitunkunde u. infektions-
krankhciten, 895-897, 1902; Ueber eine neue unterirdisch lebende Art
ix:
der Gattung Urophlyctis. Ber. der Deutschen Bot. Gesellschaft., xix: 145-
153, 1901; Ueber die in den KnoUigen Wurzelanswuchsen der Luzerne lebende
Urophlyctis, do., xx: 291-296, 1902; Erkrankung des Rhabarbers durch Perono-
spora Jaapiana, do., xxviii: 250-253, 1910.
Melhus, I. E.: Experiments on Spore Germination and Infection in Certain Species
of Oomycetes. Research Bull. 15, Agricultural Experiment Station, June,
1911.
MiYAKE, K.: The Fertilization of Pythium de Baryanum. Annals of Botany,
xv: 653.
Petersen, Henning E.: An Account of Danish Fresh-water Phycomycetes, with
Biological and Systematical Remarks. Annales Mycologici, viii: 494-560, 1910.
RosENBAUM, J.: Studies of the Genus Phytophthora. Jour. Agric. Res. 8: 233-276,
with pis. 7 and key, 191 7.
Spencer, L. B.: Treatment of Fungus on Fishes in Captivity. Bull. Bureau of
Fisheries, xxviii: 931-932, 1910.
Stevens, F. L.: The Fungi "Which Cause Plant Disease, 1913: 66-101.
Trone, a. H.: On the Fertilization of Saprolegnieas. Annals of Botany, xviii: 541.
Wager, H.: On the Fertilization of Peronospora parasitica. Annals of Botany,
xiv: 263, 1900.
Wettstein, Richard R.v.: Handbuch der systematischen Botanik, 191 1: 158-160.
Wilson, Guy West: Studies in North American Peronosporales V. A Review of the
Genus Phytophthora. Mycologia, vi: 54-83, March, 1914.
ZiRZOW, Paul: A New Method of Combating Fungus on Fishes in Captivity.
Bull, of the Bureau of Fisheries, xxviii: 939-940, 1910.
ZoPF, Wilhelm: Die Pilze, 1890: 282-313.
Wager-, H.: On the Structure and Reproduction of Cystopus candidus. Annals
of Botany, x: 295, 1896.
CHAPTER XIV
HIGHER FUNGI
SUBCLASS MYCOMYCETES
The higher true fungi are characterized by a mycelium in which the
hyphae, as a rule, are permanently multicellular by the formation of trans-
verse septa dividing the hyphal length into short cells. Some mycolo-
gists, among them Brefeld, think it important to call the fungi which
are transitional between the Phycomycetes and the Mycomycetes
proper by the name MESOMYCETES,but the distinction between
these intermediate forms and the higher fungi, being at times difficult
to make, the writer has thought it best not to use the name MESOMY-
CETES, as that of a subclass. The student will see the justice of this
viewpoint as the discussion proceeds.

Of unsatisfactory position in the fungous system are two families
of fungi, which Brefeld includes in the subclass MESOMYCETES,
which will illustrate his point of view as to transitional forms. Under
HEMIASCINEiE, as a suborder, he includes the families Ascoideaceê
and ProtomycetacEtE. Engler considers that these families have a
doubtful systematic position. They show affinity to the PHYCOMY-
CETES, and yet, they have septate hyphae and a sporangium, known
as an ascus, which contains an indefinite number of spores, hence their
closer affinity to the fungi of the order ASCOMYCETALES. The first
family is represented by Ascoidea ruhescens which lives on wounded
beech tree trunks, particularly in the sap which flows from the wounds.
It forms a brown felt-like growth. The richly septate hyphae cut off
laterally and terminally conidiospores and sporangia are formed in
a series, so that as the numerous derby-hat-shaped spores are dis-
charged and the sporangium is emptied of its contents a new sporangium
forms inside of the walls of the old one, so that ultimately a sporangium
may appear to arise out of a receptacle with a wall composed of three
or four layers. In old cultures, the fruit-bearing hyphae may be united
to form Coremia. The genus Dipodascus belongs to this family. The
I20
HIGHER FUNGI 121
family PROXOMYCETACE.f:; is represented by the {genera Prolotnyces,

Monascus and Thclebolus. Protomyccs is a genus of fungi parasitic
in the higher plants; for example, P. macrosporus lives in Umbelli-
FER^, P. pachydermus in Taraxacum. The coprophilous fungus
Thclebolus stercoreus lives on the excrement of rabbits. It has a large
rounded sporangium surrounded by a cushion of hyphge. Numerous
spores suggestive of the moulds are formed within this sporangium.
ORDER III. ASCOMYCETALES.— The fungi of this order are
characterized by a mycelium which lives either saprophytically, or
parasitically, with animals or plants. has with few exceptions a It
rank, or exuberant, development sometimes with apical growth. The
hyphae are septate and the cells are uninucleate, or plurinucleate. The
reproduction of the majority of species is through endogenous spores
known as ascospores, which are formed in definite numbers, usually
eight, sometimes less (four, two, one), and sometimes more (sixteen,
thirty-two, sixty-four, etc.) inside of a sporangium known throughout
the order as an ascus (dcr/cos = wine-skin, water bottle). Frequently,
they are called sac fungi, because of the sac-like ascus. The asci are
found either isolated, or more generally, they are in fruit bodies where
the asci are usually arranged along with the paraphyses between them
in definite layers, which may be termed ascigeral. The paraphyses
may assist in the discharge of the spores, but more usually their func-
tion is that of packing in which they serve also for the protection of
the adjacent asci. The fruit body is an apothecium, when
it is open
with the ascigeral layer wholly exposed. Such apothecia may be

platter-like, saucer-shaped, cup-shaped, or goblet-shaped, and either
sessile, or stalked, the length of the stalks being a variable character.
The perithecium is a closed fruit body sometimes produced under
ground where it remains subterranean. It may be entirely closed with
no opening (cleistocarpous), or it may open by a pore at the top.
This pore may
be borne directly at the top of the rounded perithecium,
or the perithecium may be drawn out into a larger, or a shorter neck,
so that it becomes flask-shaped, or bottle-like. A narrow canal may
lead through the neck, which may be straight, or variously curved.
Sometimes the paraphyses, which extend through the neck and out
of the pore, are designated periphyses.As accessory fruit forms, we
find the conidiospores, which are of various forms, and which are
borne singly, or in chains, at the ends of vertical hyphae (conidio-
122 MYCOLOGY
phores), or they are inclosed fruit l)odies willi terminal pores known
as pycnidia (pycnidium), and in such the conidiospores are termed
pycnidiospores, or pycnospores. The hyphae also break up into a
disconnected series of spores known as chlamydospores, or the whole
of the hypha set aside for reproductive purposes may break up into a
connected series of spores, the oidiospores. Where the conidiophores
are united together into strands, a coremium is formed. Sclerotia,
or condensed masses of resting hyphae, are not unusual in the order.

,
Certain ascomycetous fungi are lichen fungi, as they are parasitic on
green algae and with them form the lichen thallus, which bears a certain
nutritive relation with the organic or inorganic substratum, so that
we may distinguish the crustaceous, foliose and fruticose kinds of
Hchen thalH. Where such hchen fungi and others of the order ASCOMY-
CETALES live on the surface of bark, they are epiphleoidal; where
beneath the surface, hypophleoidal; where they live on rock surfaces,
they are epilithic; in rock holes, hypolithic; and on the surface of the
earth, they are epigeic; below the surface, hypogeic. The growth on
the surface of animals is ectozoic, in animals endozoic. The growth on
the surface of leaves and other plant parts is designated epiphytic or
cpiphyllous; inside the plant, as endophytic, or endophyllous. Zoospores
are never formed in any of the fungi of the order. A few are aquatic.
That sexuaHty exists in forms of the ASCOMYCETALES has been
determined only recently and these discoveries confirm the views of
de Bary, who claimed that the process existed in this order, although
Brefeld and his disciples claimed the contrary. Thanks to the epoch-
making research of R. A. Harper, seconded by that of Claussen, J.
P. Lotsy, Baur, Darbishire, Guillermond and others, the fact that

sexuality exists has been proven indubitably. The first type displayed
by Pyronema, Boudiera and related genera is where a multinucleate
carpogonium with a trichogyne is fertilized by a multinucleate
antheridium. A uninucleate antheridium unites with a uninucleate
oogonium Erysiphace^. The sexual organs are more or less
in the
reduced in .many genera and in some of the ASCOMYCETALES,
they are wanting completely. In the development of the sexual
organs and in the behavior of the egg-cell, there is represented here a
type of sexual reproduction which has its closest parallel in the red
algje (RHODOPHYCEiE). There is a suggestive similarity between
the structure of the sexual organs and the process of development
HIGHER FUNGI 1 23
following fecundation in SphcBrotheca, Pyronema and Collcma, and in

such red algae as Batr actios permum, Nemalion and Dudresnaya. A
sketch of the process will not be amiss. The antheridia and oogonia
arise in Pyronema from theapical cells of thick hyphal branches,
which from the substratum. These organs stand side
arise vertically
by side. Soon a trichogyne is formed on the oogonium, as a papillar
outgrowth, and subsequently it is cut off from the oogonium proper by
a transverse wall. The antheridium and oogonium are multinucleate
from the start and a broad stalk cell is cut off from the base of the
oogonium. The tip of the trichogyne curves over to meet the tips of
the antheridium, and the wall between them is dissolved enough to
form a pore by which the cytoplasm of one organ becomes continuous
with the cytoplasm of the trichogyne in which the nuclei have already
disintegrated. The antheridial nuclei migrate into the trichogyne, and
while this is happening the nuclei of the oogonium move to the center,
where they become collected into a dense, hollow sphere. Now the
basal wall of the trichogyne breaks down and the antheridial nuclei
pass into the oogonium and become mingled with those of the egg cell.
The antheridia and carpogonial nuclei now become paired without
fusing. Out of the oogonium grow ascogenous hyphae and the paired
nuclei pass into them. The young ascus develops from a penultimate
cell of a bent ascogenous hypha with two nuclei which fuse, after the
ascus has been formed and this fusion represents a sexual process. The
end cell of the ascogenous hypha and the stalk cell are uninucleate,
and these two cells may fuse to form a binucleate cell out of which a
penultimate cell may arise. This single nucleus of the ascus then
divides to form the series of eight ascospores usually found in the ascus.
The synapsis stage of this single nucleus is immediately followed by a
reduction division.
Claussen^ has found that the formation of the ascus is not as simple
a process, as described by Harper, and he has added materially to our
knowledge by his reinvestigation of Pyronema confluens (Figs. 38, 39
and 40). He finds that the conjugate nuclei do not fuse in the asco-
gonium (carpogonium), nor in the ascogenous hyphae, nor in the pen-
ultimate cell, nor when the tip cell of the ascogenous hook fuses with
the stalk cell to form a binucleate cell. He finds that the penultimate
1 Claussen, p.: Zur Entwickelungsgeschichte der Ascomyceten, Pyronema con-
fluens. Zeitscrift fiir Botanik, 4, Jahrgang, Heft: 1-64 with 6 plates.
124 MYCOLOGY
cell mav )enultiniatc, Up and stalk cells

and this another, and during
this process of proliferation,
the nuclei derived by descent

from the antheridial nuclei
remain distinct from those
of the ascogonium (carpogo-
nium). Even the two nuclei
derived from the tip and
stalk cells show this dif-
erence, and their descendants

remain distinct with the pro-
liferation of a new hook with
stalk cell. The series of ac-
companying figures taken

from the paper by P.
Claussen will enable the
student to understand the
process better than a lengthy
description.
The antheridia and
oogonia of Sph(Erothe.ca arise
as lateral branches of neigh-
boring myceUai filaments.
The oogonium is cut off from
the rest of the hypha by a
transverse septa, and pos-
sesses a single nucleus. The
antheridial branch appears
quite near its base and grows
upward pressed closely to the
side of theoogonium. The
antheridial cell with one
nucleus is also cut off by a
transverse septum. This
Fig. 38.— Diagrammatic representation of the
observed methods of Ascus formation. {After nucleus now divides and one
Claussen, Ziir Enlivicklungsgeschicte den Ascomy- of the two nuclei passes into
ceten, Pyronema conflucns, Zeilschr. fiir Bolanik
Jahrb., 1912.)
the attenuated end of the
4
HIGHER FUNGI 125
antheridium, which is cut off by a partition wall. The walls

between the two organs are dissolved and the male nucleus passes
through the opening formed wanders toward the egg nucleus with
which it fuses. Immediately after fertihzation, the oogonium begins
steady growth, and some of the outer cells formed become the cover
Fig. 39. — Diagrammatic representation of the development of the ascogenous

hyphal system. {After Claussen.)
cells of the perithecium. But ascogenous hyphae are formed, which

contain two nuclei, then four nuclei by division with karyokinetic
figures. Two of the nuclei wander to the curved side and this is cut
offby two partition walls to form the binucleated penultimate cell,
which becomes the mother cell of the ascus. The two nuclei of the
126 MYCOLOGY
ascus now unite. The fusion or nucleus then divides to form those
of the eight ascospores, and the walls of the perithecium grow to inclose
the asci thus formed, including the paraphyses, which develop between
the asci.
All of the typic ASCOMYCETALES have uninucleate hyphal cells,
while the ascogenous hyphse are binucleate, and in this case the nucleus
has a double chromosome number. Hence is suggested an alternation
of generations.
The life cycle of Pyronema may be displayed in a graphic form
beginning with the ascospore and ending with its production again.
The diploid, or twenty-four chromosome condition, may be repre-
sented by the double lines. This life cycle is contrasted with the
well-known one of the fern where a well-marked alternation of genera-
tion is shown.
Fern (After Claussen) Pyronema

Spore Spore
Prothalluim Mycelium
/ \
Antheridium Archegonium Antheridium Ascogonium
Spermatozoid
I.
Egg
I
cell
I
1
T. i
I I
Antheridium Ascogonium
Sperm nucleus Êgg nucleus (Sperm) Nucleus (Egg) Nucleus
/
\
Sporophyte Ascogenous hyphge
il h
Spore mother cell Uninucleate ascus
4 Spores 4 Nucleate ascus
Brown, in his studies of Leotia, has shown that the asci are formed
at the tips of the ascogenous hyphae in several different ways (Fig.
41). In some cases, to quote him, "a hypha forms a typical hook,
HIGHER FUNGI 127
Fig 40. — Diagrammatic representation of the development of the ascogenous

hyphal system and of the mature ascus. (After Claiissen.)
128 MYCOLOGY
Fig. 41.— 9, Vegetative hyphse giving rise to storage cell; 10, paraphyses grow-
ing out from storage cells; 11-14, fusion of nuclei in storage cell; 15, 16, nucleus with
two nucleoli in storage cell; 17, large storage cell with single very large nucleus; 18,
storage cell with very irregularly shaped nucleus; 19, storage cell containing one
large and two small nuclei; 20, an irregularly shaped storage cell; 21, 22, tip of as-
cogenous hypha with two nuclei; 23, two nuclei in tip of hypha have divided to four;
24, walls have come in, separating sister nuclei; 25, hook in which there is no wall
cutting off uninucleate ultimate cell; 26, hook in which two nviclei have fused to
HIGHER FUNGI 129
consisting of a binucleate penultimate and a uninucleate ultimate and

antepenultimate cell. In this case, the two nuclei of the penultimate
cell may fuse to form the nucleus ofan ascus, or they may divide and
give rise to four nuclei of another hook. The uninucleate ultimate
cell usually grows down and fuses with the antepenultimate cell,
after which the nuclei of the two cells may give rise to the nuclei of
another book or they may fuse to form an ascus. The walls separating
the nuclei may fail to be formed without affecting the fate of the nuclei.
In this process there is a conjugate division comparable to that in the
rusts. Frequently the ascogenous hyphae- do not become markedly

bent,and in this case, when the two nuclei in the tip divide, a wall
may separate two pairs of sisters. Either of these pairs may divide
and give rise to the nuclei of another hook or fuse to form the nucleus
of an ascus. Any of the methods described above by which the number
of asci is increased may be repeated many times. Large storage
cells are formed in rows which give rise to the paraphyses. They are
at multinucleate but the nuclei fuse as growth proceeds. This
first
process continues until often the cells contain a single very large nucleus
many times the size of the largest nucleus in the ascus. The nuclei
are very irregular."
Blackman, V. H. AND Fraser, H. C, Jr.: Fertilization in Sphjerotheca. Annals

of Botany, 19: 567-569, 1905.
Brown, W. H. The Development
: of the Asocarp of Leotia. Botanical Gazette,
50: 443-459-
Claussen, p.: Zur Entwickelung der
Ascomyceten Boudiera. Bot. Zeit., 68
(1905): Zur Entwickelungsgeschichte der Ascomyceten Pyronema confluens.
Zeitschrift fiir Botanik, 4 Jahrgang, Heft i: 1-64; Ueber
neuere Arbeiten zur
Entwickelungsgeschichte der Ascomyceten. Bar. der. deutsch. Bot. Gesellsch.
Jahrg., 1906, Band xxiv: 11-38 with complete bibliography.
Engler, A. AND GiLG, Ernst.: SyUabus der PflanzenfamiUen, 1912:
47.
form nucleus of ascus, and tip has fused with stalk of hook;
27, ultimate cell has
fused with antipenultimate; nucleus of latter has migrated
into former, which is
growing out to give rise to ascus or another hook; 28, two nuclei
of penultimate cell
have fused to form nucleus of ascus; ultimate cell has fused with
antepenultimate
and nucleus of latter has migrated into former, which has grown out
to form another
hook; 29, bmucleate penultimate cell has given rise to hook;
ultimate cell has fused
with penultimate, and the two nuclei have fused; ultimate
cell has not developed
further; 30, binucleate penultimate cell has formed
ascus, which fusion product of
ultimate and antepenultimate has given rise to second ascus;
31, diagram illustrating
multiplication of number of asci by method shown in
26-30; 9-20 Xr400. 21-30 X
2100. (Aftey Brown. William H., The Development of the Ascocarp
of Leotia. Botanical
Gazette, 50: 443-359, Dec, 1910.)
9
I30 MYCOLOGY
Fraser, H. C. J. AND Welsford, E. T.: Further Contributions to the Cytology of

the Ascomycetes. Annals of Botany, xxii (1908).
GuiLLERMOND, A.: La quest, d.l. sex chez. 1. Asc. et les. rec. trav. Rev. gen. de
Bot., XX (1908): Recherches Cytologique Taxonomique sur les Endomycetes.
Rev. gen. de Bot., 21: 353-39; 401-419 (1909) and a number of papers on the
same subject in vol. 20.
Harper, Robert A.: Die Entwicklung der Peritheciums bei Sphasrotheca Cast-
agnei, Ber. d. Deutsch. Bot. Gesellsch., 13: 475-1895; Kerntheilung und
freie Zellbildung in Ascus Jahrb. f. wiss. Bot., 30: 249-284, 1897; Cell Division
in Sporangia and Asci. Annals of Botany, 13: 467-524, 1899; Sexual Repro-
duction in Pyronema confluens and the Morphology of the Ascocarp. Annals
Botany, 14: 321-400, 1900.
of
MoTTiER, David M.: Fecundation in Plants. Publ. 51, Carnegie Institution of
Washington, 1904.
Sands, M. C: Nuclear Structure and Spore Formation in Microsphasra alni.
Trans. Wise. Acad. 733-752; Botanical Gazette, 46:79.

Sci., 15;
Ward, H. Marshall:: P'ungi, Encyclopedia Britannica, nth Edition.

Wettstein, Richard R.v.: Handbuch der systematischen Botanik, 1911: 169-172
CHAPTER XV
SAC FUNGI IN PARTICULAR (YEASTS, ETC.)
Suborder A. Protoasciineae.— The fungi of this suborder are charac-
terized by the absence of definite fruit bodies, that is the asci are not
enclosed, but are free and at the ends of hyphae. Usually they are of
unequal length. Four is the typical number of ascospores in each
ascus. These are one-celled and may increase in number by gemmation.
—
Family i. Endomycetace^. This family is a small one of four
genera of saprophytes and parasites. The two species of the genus
Podocapsa are parasitic on Mucorace^, Eremascus albus, the single
species of that genus grows on spoilt malt extract. The genus En-
domyces with five species is represented by the cosmopolitan Endo-
myces decipiens, which forms a snow-white parasitic growth on the
toadstool Armillaria meltea. Its hyphae are branched richly and the
asci are pear-shaped and borne singly at the ends of the branches,
each producing four helmet-shaped ascospores, 6 to S/jl broad and
sû
high. Conidiospores are more frequently foimed than ascospores.
Oidiospores are also found, as well, as chlamydospores. Oleina nodosa
and O. lateralis are the two species of the fourth genus. The first
grows in ohve oil.
Family 2. ExoASCACEifc.— This family includes parasitic fungi

which cause abnormalities of more or less marked character of the
leaves, fruits and branches of mostly woody plants. The malforma-
tions are in the nature of witches' brooms of the smaller branches,
leaf curls, and deformed fruits, such as the plum pocket. Stone fruits
are especially subject to attack and
some cases the stone formation
in
is suppressed entirely. The myceUum may be deep-seated and
perennial, or may be subcuticular, or sometimes found growing
it
between the epidermal cells, as in Magnusiella flava, while in other

forms, the hyphae may
be below the epidermis and grow throughout
the leaf tissue. The formed on the surface of the
asci are generally
host breaking through from the more deep-seated mycehum beneath.
They are generally stalkless and arranged in close proximity to each
131
—
132 MYCOLOGY
other without paraphyses, so that they form a velvety layer on the

surface of the host plant. Eight ascospores are generally found, as in
the genus Exoascus, but in Taphr'ma (Taphria) the number may be
increased considerably by budding, so that the whole ascus will be
-::#\
^h
y ,
,; ff
^^^ ^rrf^f
Fig. 42. Exoascus and Taphrina. A-F, Exoascus pruni, A. Appearance on

diseased twig; B, cross-section of diseased fruit; C, mycelium in tissues of host; D,
young asci; E, mature ascus with spores; F, germination of spores; G, E, Exoascus
alnitorquus; H, Taphrina aurea, ripe and unripe asci; J, Taphrina Sadebeckii. See
Die naturlichen Pflanzenfamilien I. i, p. 159.
crammed full of them (Fig. 42). The ascospores are generally ellip-
and always one-celled with colorless, yellow, or orange contents.
soidal
The perennial mycelium is responsible for the formation of witches'
brooms in a variety of trees and woody plants. Most of them are the
SAC FUNGI IN PARTICULAR -
1 33
result of the parasitism of species of Exoascus. The "hexenbesen"

are brush-Uke, or tufted masses of branches, which suggest the presence
of other plants (Uke the mistletoe) parasitically or epiphytically
growing. They result mainly by the infection of a bud which de-
velops a branch with increased growth. On this branch, all dormant
buds are stimulated to activity and the whole infected system of
branches consists of negatively geotropic branches. These brush-
like excrescences are called the thunder-bushes, and are -sometimes
nest-like in appearance. An anatomic study shows that the parenchy-
— —
matous tissues pith, hypodermis, etc. are greatly increased; wood
and bark are traversed by abnormally broad medullary rays, the ducts
have short members, the wood fibers wide lumina, which are sometimes
thin-walled and septate. The bast fibers are few, or entirely wanting.
The cork cells are enlarged and retain their protoplasmic contents a
longer time. The form of the witches' brooms are various. Many
of them are pendent, some are nest-like, owing to the death of some of
the branches. In some the branches are elongated, while some have
short twigs. The end of the original branch from which the lateral
branches developed usually dies and its food substances are absorbed
by the hypertrophied branches. The family includes three genera,
distinguished, as follows:
A. Asci found at the end of intercellular mycelial branches.

I. Magnusiella.
B. Asci developed on a more or less subcuticular ascogenous mycel-
ium.
(o) Asci eight- (exceptionally four-) spored. 2. Exoascus.
ib) Asci many-spored by gemmation of the spores. 3. Taphrina.
The genus Magnusiella comprises five species, four of which are found
in Europe and two in America. Magnusiella flava forms small pale
yellow specks on the leaves of the gray birch, Betula populifolia in
North America. The genus Exoascus includes about thirty species
arranged in two subgenera, the first of which includes those species which
deform fruits, which form witches' brooms, and the second those which
cause a spotting of the leaves of various plants. It would lengthen
this book unduly to enumerate all of the species of Exoascus with an
account of the deformities of branches and fruits which they produce.
Only a few of the more important species will be enumerated here,
134 MYCOLOGY
and the diseases which they cause will be described later. Exoascus
pruni (Fig. 42) is the cause of an important disease of plum trees,
producing the so-called plum pockets. It also attacks Pruniis domestica

and P. padus in middle Europe, and P. domestica and P. virginiana in
North America. Exoascus communis attacks the fruits of several
American species of Prunus among them P. maritima. Exoascus
alnitorquus infests the pistillate spikes and cones of species of alder
(Alnus), such as Alnus glutinosa and A. incana in middle Europe, and
A. incana and A. rubra in North America, causing an enlargement of
the fruit scales into twisted, tongue-like, reddish outgrowths. Exoascus
deformans is the cause of peach-leaf curl. Exoascus cerasi is responsible
for the formation of witches' brooms on the cherry. The genus
Taphrina causes witches' brooms and leaf spots. Taphrina purpuras-
cens attacks the leaves of a North American sumac, Rhus copallina,
causing a puckering of the leaves with the formation of a reddish-
purple color.T. aurea (Fig. 42) forms yellow blotches on the leaves of
several European and North American poplars, viz., Populus nigra and
P. italica of Europe, and P. Fremontii, P. grandidentata and P. deltoides
of North America. T. Laurencia causes witches' brooms on a fern in
Ceylon, Pteris quadriaurita.
Suborder B. Saccharomycetiineae. —A true filamentous mycelium
is absent in the fungi of this suborder. The plants are single-celled
and reproduce by budding, or gemmation. Occasionally under ex-
perimental treatment where the culture media are varied, the cells
develop into hyphae and together form a myceHoid growth. Spore

formation consists in a single cell, developing one to eight spores.
It, therefore, may be looked upon as an ascus and the spores are as-
cospores. Many ofthem cause fermentation.
Family i. Saccharomycetace^. Many — species of the genus
Saccharomyces are called generically yeasts, and are of economic
importance, because they induce the alcoholic fermentation of car-
bohydrate substances. The action is accompl shed through a soluble
enzyme formed in the protoplasm of the yeast cell, and first isolated
by Buchner by grinding the yeast cells in sand and extracting the
ferment zymase. The general shape of yeast cells is oval, ellipsoidal,
and pyriform (Figs. 43, 44). The cell wall is well defined and consists
of modified forms of cellulose which may be called fungous cellulose,
because it does not react to the reagents used for true cellulose. This
SAC FUNGI IN PARTICULAR 135
much can be said that the wall consists of a carbohydrate, probably

some isomer of cellulose. Lining the inner surface of the cell wall is a
layer of protoplasm which may be called the ectoplasm, which probably
serves as an osmotic The cytoplasm fills the rest of the cell
membrane.
with the exception of spaces occupied by the vacuoles of glycogen,
nuclear vacuoles, oil globules, the nucleus and nuclear granules. The
glycogen is gradually used up as it probably serves as reserve food,
the same as starch in the higher plants. These glycogen vacuoles
generally coalesce until one large vacuole may almost fill the cell.
J
6 7 8 12
Fig. 43. Fig. 44.
Fig. 43. — Yeast
cell, Saccharomyces cerevisicB. {After Marshall.)
Fig. 44. — Yeast,
Saccharomyces cerevisicB. i-io. Young cells with nucleus,
showing structure; 6-8, division of nucleus; 11-13, cells after twenty-four hours"
its
fermentation with large glycogenic vacuole filled with lightly colored grains. {After
Marshall, Microbiology, Second edition, p. 62.)
the cytoplasm and nuclear bodies being pressed against the cell wall
and forming a thin protoplasmic hning to the inner cell wall surface.
Wager in 1898 demonstrated the nuclear apparatus in a number of
1
yeast species. The nuclear apparatus consists in the earliest stages of

fermentation of a nucleolus in close touch with a vacuole (Fig. 44, No.
4)
which includes a granular chromatin network suggesting a similar struc-
ture in the higher plants. The vacuole may disappear and then the
chromatin granules are scattered through the protoplasm, or are gathered
around the nucleolus, which is present in all of the cells, as a perfectly
homogeneous body. Numerous chromatin vacuoles are often found
1 Wager, Harold: The Nucleus of the Yeast Plant. The Annals of Botany
.xii: 400-539-
136 MYCOLOGY
in young and these ultimately fuse to form a single vacuole which

cells
occurs in the during the earher and the later fermentation. The
cells
process of budding is associated with the stretching of a network of nu-

clear granules and its final constriction in the neck between the mother
and the daughter cell. The nucleolus moves to the constriction where
it becomes dumbbell-
shaped, one half press-

ing into the daughter
cell (Figs. 44 and 45).
There are no stages of

karyokinesis dis-
by the sim- w^
played, but
pie process described ^^ -"^
f 1,«k
^^^ ^W ^^ /<^
above the daughter

cell receives approxi-
mately one-half of the
nuclear substance of
r^ ^
y^^r.
Fig. 45. Fig. 46.

Fig. 45. — Young yeast Saccharomyces
cells, with nuclei and division
ellipsoideus,
of nuclei. (After Marshall, Microbiology, Second edition, p. 64.)
Fig. 46. — Yeast, Saccharomyces the variety known as brewers' bottom
cerevisice,
yeast; a, spore formation; h, elongated cells. {After Schneider, Pharmaceutical Bac-
teriology, p. 144.)
the mother cell. In spore formation, the chromation which is scattered

through the cytoplasm is absorbed more or less completely into the
nucleolus which elongates and divides by a constriction in its middle
part. Subsequent divisions result in the formation of four nucleoli
around which protoplasm collects and thin membranes which become
the walls of the ascospores which remain at first small, but later increase
in size (Fig. 46). The formation of spores can be secured by taking
a Sterile block of plaster of Paris with a saucer-shaped hollow on

top. This block is placed in sterilized water and the top is seeded
with vigorous, young well-nourished yeast plants which develop spores
if kept at 25°C., in from twenty-four to forty-eight hours. The tem-
perature at which spore formation occurs and the time which it takes
for sporulation are points which have been obtained by experimenta-
tion for all the more important species of yeasts. The data which has
been obtained is used in the physiologic diagnosis, or identification of
the various kinds of Saccharomycetace^, which react differently
under experimental treatment. Film formation is also of diagnostic
importance, where economic yeasts form floating films on the nutrient
liquid media in which they are grown. The time
required for the development of the film differs, /^
other conditions being equal, with the species of r^\^(^ ^^
the yeast and is longer the lower the temperature
of the culture. Hansen obtained the following data
for Saccharomyces cerevisicB ;
Film formation takes place at:

Q^^
about ^/^- 47-— Yeast.
S3° to 34°C. in 9 to 18 days. ^
o i "
no,-^ • . ^ i 1
Saccharomyces cerevi.
20 to 28 C. in about 7 to 11 days. ,-^_ growing repro-
13° to I5°C. in about 15 to 30 days. duction by germina-
6° to 7o°C. in about 2 to 3 months. tion, or budding; a,
single cells; b, bud-
No formation of film occurred above 34°C. or below ding cells. {After
5°C. Another point of importance is that species

ÎZiJ^X'^'^J''
of Saccharomyces form films so that this process is
not entirely associated with the fungi belonging to the so-called genus
Mycoderma. In fact some authors recognizing that Saccharomyces
cerevisicB (Fig.47) produced films have named that yeast, Mycoderma
and have thus confused its identity.
cerevisicB,
Hansen in a paper published in 1888 classified the yeasts essentially,

as follows:
1. Species which ferment dextrose, maltose, saccharose: Saccharo-
myces cerevisicB I, S. Pastorianus I, S. Pastoriamis II, S. Pastorianus
III, S. ellipsoideus I, S. ellipsoideus II.
2. Species which ferment dextrose and saccharose, but not maltose:
Saccharomyces Marxianus, S. exiguus, S. Ludwigii S. saturnus.

3. Species which ferment dextrose, but neither saccharose nor
maltose: Saccharomyces mali Duclauxii.

138 MYCOLOGY
4. Species which ferment dextrose and maltose, but not saccharose

Saccharomyces n. sp. obtained from stomach of bee by Klocker.
5. Species which ferment neither maltose, dextrose nor saccharose:
Saccharomyces anomalus var belgicus, S. farinosus, S. hyalosporus, S.
memhranifaciens.
The general chemic phenomena associated with the formation of
alcohol by fermentation out of sugar may be expressed by the formula:
CeHioOe = 2C2H6O + 2CO2

Alcohol Carbon
dioxide
The carbon dioxide passes off in bubbles as a gas, while the alcohol
remains in solution.
The most important yeast is the beer yeast Saccharomyces cerevi-
sicBwhich is a unicellular plant of spheric or elliptic shape 8 to 12/1
long and 8 to 10// broad. Sometimes the cells formed by budding
remain connected to form a chain consisting of the mother, daughter,
granddaughter and great-granddaughter cells. Spore formation is
characteristic and the size of the spores varies from 2.5 to 6/i. There
are usually four spores in each cell. The following gives the tempera-
ture conditions of spore formation in this species:
At 9°C. no spores develop.

At 11° to i2°C. the first indications are seen after 10 days.
At 3o°C. the first indications are seen after 20 hours.
At 36° to 37°C. the first indications are seen after 29 hours.
At 37.S°C. no spores develop.
The temperature limits for film formation are 33° to 34°C. and 6°
to 7°C. There are a number of races of the common beer yeast, which
may be separated into the bottom yeasts and the top yeasts. The bot-
tom yeasts are those which live within the hquid and mostly at the
bottom even from the start. Some of these yeasts form spores with
difficulty. The top fermentation yeasts are those which grow on the
surface of the liquid and cause a brisk'fermentation with a large amount
of froth, or head, as exemplified by the Munich lager-beer yeasts.
Yeasts are among the oldest of cultivated plants, as in biblical times
leavened and unleavened bread were known.
(yeast-raised) The
leaven was a lump of dough kept from one baking to the next. Un-
leavened bread was simply flour mixed with water and baked, and as
a result, a hard tough bread was obtained. The .use of yeast as a
—
Starter began in Roman times, but the art was lost until the seventeenth
century, whenwas regained. One of the earliest methods of obtain-
it
ing yeast was which consisted in adding to a quantity of

salt raising,
milk a little salt sufficient to delay the growth of bacteria, while the
yeast found entrance to the milk through the air and grew rapidly.
This milk was then mixed with dough for the raising process. Bakers
also sometimes used a brew called barms. Scotch barms were prepared
by taking hops and flour with other ingredients which were allowed to
^ '^^ ^ ^^^
C^^^
Fig. 48. Saccharomyces ellipsoideus. A

common yeast in jams, jellies, etc.
Budding process is shown in many of the cells as also the vacuoles. Fig. 66, p. 145,
Schneider, Pharmaceutical Bacteriology, 1912.
ferment spontaneously, and the fermented material was used in

bread baking (see page 667).
Saccharomyces ellipsoideus (Fig. 48) is known as the wine yeast and
may be classed as a wild species, while the beer yeast is found only in
cultivation. The vegetative cells are ellipsoidal 6/x long, single, or
united into a row of loosely connected cells. The cells are two- to four-
spored. The spores are spheric 2 to 4/i broad. It is important in the
fermentation of grape juice, gaining entrance from the skin of the
grape fruit upon which it lives. In the spore form, it overwinters in
the soil, being blown as dust to the developing grape fruits. The
I40 MYCOLOGY
bouquet, or flavor of the wine seems to be clue to the variety of wine

yeast used in the fermentation of the juice, for every wine-producing
region seems to have its especial form of wine yeast and the growth is
different. Some yeasts, such as those of Burgundy and Champagne,

form a compact sediment, which quickly settles leaving the liquid
clear, while others remain for a long time suspended and settle slowly.
Saccharomyces ellipsoideus II is a very dangerous disease yeast, produc-
bottom fermentation breweries.
ing turbidity in the liquid of
Saccharomyces Pastorianus I was first discovered in the dust of a
Copenhagen brewery and also in diseased beer. Its growth in wort
consists of sausage-shaped cells. II produces a feeble
S. Pastorianus
top fermentation. S. Pastorianus III was found in bottom fermenta-
tion beer affected with yeast turbidity.
Saccharomyces ilicis and 5. aquiJolU were found on the fruits of the
holly, Ilex aquifolium.
Saccharom,yces Vordemanni is similar in appearance to wine yeast,
its cells being onion-shaped, or pear-shaped. It is present in Raggi,
which is employed in Java in the manufacture of arrack. It forms 9
to 10 per cent, alcohol.
Saccharomyces pyriformis was discovered by H. Marshall Ward to be
active in the formation of ginger beer in conjunction with Bacterium
vermiforme, for when these organisms are added to a sugar solution
containing ginger, an acid beverage with considerable head is formed
known as ginger beer.
Saccharomyces exiguus occurs in pressed yeast, and it is capable of
developing considerable alcohol from dextrose and saccharose solutions.
Saccharomyces anomalus has been found in impure brewery yeast
in Hungary, also in Belgian beer, on green malt, on bran, in syrup of
Althaea, in soil, and on plum fruits. It ferments wort readily forming
a gray film, a turbidity in the liquid, and an odor like fruit ether.
The spores are helmet-shaped, suggesting those of Endomyces decipiens,
which is parasitic on the caps of A rmillaria mellea, a toadstool. Saccha-
romyces memhranifaciens grows in a gelatinous mass on the injured roots
of elm trees, in polluted water, and in white wines, where'it destroys the
bouquet of the wine. It completely consumes acetic and succinic
acids, and quickly forms gray corrugated films on the surface of wort.
The organisms of Kefir are Saccharomyces cartilaginosus and S. fragilis.
Kefir is a beverage prepared originally in the Caucasus region by fer-
menting milk. Kefir grains, which include the above yeasts, a Torula,
and 3 bacteria {Bacillus caucasicus, etc.) are added to the milk as a
starter. The fermentation of the milk results in the formation of alcohol
lactic acid and carbonic acid. Mazum (Matzoon) an Armenian drink,
is prepared by adding a white, fatty cheese-like mass, to milk.
The
starter includes colored yeasts Oidium laclis, mould fungi, a yellow
Sarcina, Bacillus subtilis, some cocci. Bacterium acidilactici a.nd Saccharo-
niyces anomalus. The only species of yeast, which can be recognized
immediately by microscopic examination, is Saccharomyces Ludwigii,
with its lemon-shaped vegetative cells, on the point of which a wart
makes its appearance, which is cut off by a septum from the rest of the
cell. This species is transitional to those included in the genus Schizo-
saccharomyces. The form of Saccharomyces Ludwigii suggests S.
apiculatus, which is unequally dumbbell-shaped. The genus Tonda
according to Hansen includes yeasts similar to Saccharomyces, but
which do not form endospores, a typical mould growth, and which
produce alcohol in all percentages. They are widely distributed in
nature.
Schroter in Engler's "Die naturlichen Pflanzenfamihen" recognizes
only two genera in the yeast family, namely, Saccharomyces and Mono-
spora. The reproductive cells of the former have two to eight (seldom
one to three) spores and the spores are spheric, or ellipsoidal, while the
needle-shaped spores of Monospora are borne singly in reproductive
cells,or asci. Hansen^ considers Monospora to be a doubtful form of
yeast {Saccharomyces douteux), as also the genus Nematospora. He
recognizes the following genera: Saccharomyces, whose spores have a
single membrane and the cells reproduce hyhxiddrng; Zygosaccharomyces,
where the asci are associated with conjugation; Saccharomycodes, whose
spores have one membrane and sprout into a promycelium; Saccharo-
mycopsis, whose spores have two membranes; Pichia with hemispheric
or angular sporesand Villia with citron-shaped spores. Lafar in his
book on "Technical Mycology" (II, part 2, page
274) gives an analytic
summary of the genera which he believes should be recognized. The
position of such genera as Zygosaccharomyces, Saccharomycopsis,
Schizosaccharomyces with respect to nearly related fungi is presented
and discussed with a diagrammatic scheme of relationship by
^ Hansen, E. Chr.: Grundlinien zur Systematik der Saccharomyceten. Centr.
f. Bak., 1904.
142 MYCOLOGY
Guillermond/ who suggests the probable evolution of such forms from

Eretnascus and Endomyces. Dr. H. Will discusses in Lafar's book
the family Torulaceê, species of which are widely disseminated on
field and garden fruits and on plants of all kinds finding suitable condi-
tions for their growth during the decay of these fruits, and during the
technic processes of fruit preservation, such as the making of pickles
and sauerkraut. A number of them will no doubt prove to be budding
stages of other fungi for our knowledge of them is decidedly imperfect.
The character of the so-called pink yeast, red yeast, and black yeast is
even less well known. As they some have even

are budding fungi,
classed them with the genus Saccharomyces. The genus Mycoderma
was created to include the budding fungi, which form true films and
which are formed rapidly on nutrient liquids, particularly on beer
and wine with air between the cells, which are usually short and sau-
sage-shaped. They are strongly aerobic and form, when exposed to
the air, a wrinkled skin on the surface of the liquid. Like the true wine
yeasts, these various species of Mycoderma have their natural habitat
in the soil and they are carried to their appropriate nutrient substances
by insects, rain or wind. They are probably not true yeast plants, but
may represent growth conditions of other fungi, as related to certain
nutrient materials. Curious chemic activities are possessed by species
of Mycoderma, for example, the formation of acids and their destruc-
tion both at the same time. Citric and succinic acids for example are
consumed by them.
1 GuiLLEEMOND, M. A.: Rcchetches Cytologiques et Taxonomiques sur les
Endomycetees. Revue Generale de Botanique, 21: 401-419, 1909.
CHAPTER XVI
SAC FUNGI CONTINUED
Suborder C. Plectasciinese.^This suborder includes fungi with a
well-developed mycelium on which are developed either on the surface
of the substratum or within it, as in the subterranean forms, closed
perithecia without an opening at the top. The wall of the perithecium
is sometimes called the peridium. The asci are developed on hyphae
of irregular branching, and in considerable numbers, forming irregular
layers of the perithecial interior. Each ascus is rounded and three- to
eight-spored. The spores are one- to many-celled. Condiospores
occur in a few of the forms, such as Aspergillus, Meliola and Penicillium.
Many of the fungi of this suborder are saprophytic, but some are de-
cidedly parasitic, as Thielavia basicola, which destroys the roots of pea
plantsby its parasitic growth and species of the families Terfeziace^
and Elaphomycetace^, the mycelia of which form mycorrhiza with
roots of flowering plants. Economically, the suborder is interesting,
because it includes the common blue and green moulds and species of
Aspergillus used in the fermentation industries. The fruit bodies
of several kinds of Terfezia are used as food by the Arabs of North
Africa, Arabia, Syria and Mesopotamia.
Family i. Gymnoascace^. —The fungi of this family are of interest,
because of the structure of their fruit bodies. In the genus Gymnoascus,
the spheric asci arise on short lateral branches of hyphae which form a
dense rounded mass inclosed by loosely branching hyphae, which form
a basket-like inclosure of the ascus-bearing portion Gymnoascus Reesii
is coprophilous. Some of the shorter branches of this outer envelop-
ment are sharp-pointed and spiny. Ctenomyces serratus, the single
representative of its genus, grows on decaying bird feathers. It
has branches with short hook-like extremities. The fruit body in this
fungus is similarly rounded and covered with hyphae that form an
open basket-Hke peridium.

—
Family 2. Aspergillace^. This family includes fourteen
genera, the most important of which are Aspergillus, Penicillium and
143
144 MYCOLOGY
Thlelavia. The perithecia are never subterranean. They are usually

small, spheric, usually closed, and their walls are made up of pseudo-
parenchymatous hyphge. They rarely open by a pore, more usually
they break up at maturity to allow the escape of the ascospores. The
inclosed asci are spheric to pear-shaped and two- to eight-spored.
The moulds of the genus Aspergillus (Figs. 49 and 50) are usually
saprophytic, and are found upon decaying vegetables, moldy corn and
other cereals. After the conidiospores are formed, the color of the
mould develops and various shades of green, white, blackish-brown,
brownish-yellow, brown and reddish are found in the different species
of the genus. The recognition of this genus is made easy by the shape
of the conidiophores, which are elongated unicellular (unseptate) and
terminate in a globular swelling, the top of which is covered with a large
number of closely set stalks, or sterigmata, of variable length and shape
on which the conidiospores develop. In the related genus Sterigmato-
cystis, the sterigmata are branched (Fig. 51). The conidiospores are
spheric, or ellipsoidal, always smooth or granular
unicellular with
walls, and are formed in long chains (concatenation) from each sterigma
imparting the characteristic color to the whole growth. The perithecia
are fragile spheres with thin walls which may be yellow {A. herbari-
orum) dark red {A. pseudo-clavatus) , or even black (A. fumigatus)
in color. The perithecia and asci are unknown in many of the species,
so that the classification of the species cannot be based on the characters
of that organ and of the ascospores. Only about six to ten species are
known have perithecia out of a possible total number of 120 species
to
included in the genus. This number will probably be considerably
reduced when these moulds are better known. The accompanying
figures show some of the specific differences of the conidiophores and
conidiospore production. The .common green mould, Aspergillus
herbariorum (= Aspergillus glaucus, Eurotium Aspergillus glaucus)
grows on many substances such as dried plants in the herbarium,
(hence its specific name), on old black bread (pumpernickel), on jellies,
on jams, on old leather, on herring pickle and other objects of domestic
use. At first the mycelium is white and as the young conidiospores begin
to form it turns to a pale green, later becoming a dirty grayish green,
while the feeding hyphse change color to a pale yellow and finally a
brown color by the deposit of pigment granules. The globular part of
the conidiophore is 60/x across and crowded with simple sterigmata
—
SAC FUNGI CONTINUED I45
(7M by i4m), bearing prickly, spheric conidiospores 7 to 30^ in diameter

which are larger than any other well-known species. It produces
perithecia also with readiness and in abundance. The at first pale
brown-yellow perithecia, later brown, are about 100 to 200/i in diame-
ter in closing numerous asci which contain five to eight colorless
smooth ellipsoidal spores, exhibiting a furrow directed longitudinally
and 5 to 8/i broad by 7 to IOf^ long. The perithecium develops gradu-
ally from spirally coiled hyphae. The hyphae of the screw are divided
Fig. 49. Aspergillus oryza associated with yeasts in the making of the Japanese
beverage Sake. Vegetative hyphae (a) and spore-forming hyph« {b. c, d) are shown.
Fig. 71, p. 152. {Schneider, Pharmaceutical Baderiology, 1912,
19.)
transversely into as many cells as there are turns of the screw. The
bottom hyphal cells of the screw send up two or three branches of
which grow toward the apex. One of these branches
irregular thickness
looked upon as an antheridium grows more rapidly than the others and
its contents serve to impregnate the inclosed carpogone. These outer
erect hyphae then branch copiously to completely envelope the carpo-
gone and the perithecial wall is thus formed. From the carpogone are
now formed the numerous ascogenous hyph;p, which branch plenti-
146 MYCOLOGY
fully and bear terminally asci of a pyriform shape. These contain

eight grooved ascospores. Aspergillus herbarioriini, as a domestic
and industrial fungus, is selective. It does not thrive on liquid sac-
charine media with mineral salts and inorganic nitrogenous food, while
black bread and wort gelatin are suitable media. Moderate tempera-
tures (8 to io°C.) are best for its growth, and it ceases growth entirely
at blood temperatures. The temperature limits are 7° to 3o°C. with
optimum at 20 to 25°. It grows on tobacco, cigars, hops, cotton-seed
meal, acid pickles, and smoked meats. It causes the blackening and
spoiling of chestnuts and is found on the kernels of various nuts even
before they are removed from the shell (see Appendix VII, pages 702
to 721).
The rice mould, Aspergillus oryzece (Fig. 49), is of practical impor-
tance as a saccharifying fungus, and it has been cultivated for centuries
by the Japanese and used by them in the preparation of the rice mash
for Sake, as well as in the production of Miso and Soja sauce. It grows
luxuriantly and is usually yellow-green in color turning brown with age
with large closely set tough conidiophores about 2 mm. tall. The tops
of its conidiophores are obovate, or spheric. The sterigmata are radially
arranged producing yellowish-green spheric conidiospores (6 to 7/i) in
chains. The sterigmata are larger than in A. herhariorum 4 to 5^1 by 12
to lOjj. . No perithecia
have yet been observed. This mould secretes a
very active diastase andit has been used in the making of pharmaceutic
preparations, such as Taka diastase, which is used in the dose of 2 to 5

grains either in tablet, capsule or solution in cases of indigestion im-
mediately after meals. It converts the starchy food into dextrin and
sugar. The discovery of this diastase in Aspergillus was made by
Takamine, a Japanese zymologist, and his product has been used over
the civiHzed world.
Aspergillus Wentii, which is readily kept in culture on glucose or
beerwort agar, is used in the preparation of Tas Gu in Java. It appears
spontaneously on boiled soy beans that have been covered with leaves
of Hibiscus and it causes a loosening and disintegration of the firm
tissues of the bean. The growth of this species is of a pale coffee color
with conspicuous conidiophores about 2 to 3 mm. in height, their thick
brown heads up to 200/i in diameter are on pale smooth stalks. The end
of the conidiophore globular 75 to 90/x in diameter and is covered
is
with slender simple sterigmata (4^ by 15/x) which bear small globular
to elongated conidiospores, 4 to 5/1 diameter. The mycelium at first
SAC FUNGI CONTINUED 147
is snow-white; later it becomes reddish brown. The discovery of

perithecia is yet to be made.
Aspergillus flavus plays an important part in the cocoon disease of
silkworms. The stipe portion of its conidiophore is roughened by
colorless granules.
Aspergillus luchuensis, according to Inui, is used in the preparation
of a beverage Awamori, which resembles whisky and is used in the
Loochoo islands.
Aspergillus tokelau is found in Tokelau, or Samoan disease, attack-
ing the natives of certain of the Pacific islands. An important patho-
genic species, which causes an epidemic disease of pigeons and lives in
the human ear and the lungs of various birds, is Aspergillus fumigatus,
which was the cause of a false tuberculosis of a calf in Philadelphia.
An autopsy by Ravenel and the writer showed the lung tissue of the
calf penetrated by the myceUal hyphae of the fungus, and its conidio-
phores bearing the conidiospores in a fan-Hke manner were seen project-
ing into the lung cavities almost completely filling them. It, therefore,
grows well at blood temperature, and if its conidiospores are introduced
into the arterial circulation of animals they germinate and produce
serious illness, which may terminate fatally. It also acts injuriously
in certain fermentation processes carried on at high temperatures as
certain lactic acid fermentations. It attacks tobacco, decaying
potatoes, bread, malt and beerwort. It has dwarf conidiophores o.i
to 0.3 mm. long, with club-shaped globules 10 to 20/x thick, upright
15/1 long and with long chains of conidiospores (2 to 3^)-
sterigmata 6 to
Nut-brown globular perithecia are found, 250 to 350M in diameter, in-
closing oval thin-skinned asci (9 to i4ju) with eight red lenticular tough-
walled spores (4 to 4.5^)- As a parasite of the human skin it was called
Lepidophyton. The green mould, which usually grows on malt, is
Aspergillus clavatus causing a moulding of the substratum. The largest
species of the group is Aspergillus giganteus, which looks at first super-
ficially hke a Mucor, but later owing to its grayish-green conidiospores
it is readily separable from the mucor vegetation. Its sterigmata seem
to be hollow, communicating with a pore-like opening with the center
of the conidiophore. No perithecia have been found. Other species
are nidulans (Fig. 50), which can be cultivated readily, A. varians
.4.
and A. ostianus, the latter distinguished by an ochraceous pigment.

The black mould Aspergillus niger more properly Sterigmatocystis niger
148 MYCOLOGY
(Fig. 51) has a copious literature. Lafar cites forty workers of recent
date, who have studied it. The physician finds it as an occupant of
Fig so — Aspergillus nidulans. A, Mycelium with conidiophores; B, branched

conidiophore, C, spore chains at end of conidiophore D, small conidiophores; E,
;
young fruit showing development of covering; jp, hyphae with swollen ends; G,
hypha from interior of fruit-body; H, hyphae with young asci; J, developing perithe-
cium. (See Die jtaliirlichen Pflanzenfamilien I. i, p. 302.)
the human ear in a disease otomycosis. It is associated with the cork

disease which imparts a taste to bottled wine. It grows well in acid
substrata, as gall-nut extract, tannic acid and has a decided capacity
—
for producing oxalic acid. It has stiff slender conidiophores several

millimeters in height. The terminal part can be studied only after
the bleaching or removal of the dark masses of conidiophores.
Fig. 51. Slerigmatocyslis niger {Aspergillus niger) showing conidiophores and coni-
diospore formation with stages in germination of spores. {After Henri Coupin.)
The genus Thielavia is represented by a common pathogenic species,

T. basicola, whose life history and pathogenic character will be de-
150 MYCOLOGY
scribed later. It attacks the roots of a large series of plants including

the tobacco, at least 105 species of plants being attacked according to
the latest account.^ The parasitic mycelium is intercellular, abun-
dantly septate and hyaline. It produces conidiospores, which are
abjointed acrogenously from the conidiophore, and are not as was
supposed formerly endospores formed by free cell division within an
endoconidial cell. The first conidiospore is liberated by the differentia-
tion of its walls into an inner wall and a sheath and by the rupture of
the latter at its apex. The later conidiospores grow out through the
sheath of the and are freed by a spHtting of their basal walls.- This
first
same process is probably that of all " endoconidia " in fungi.

—
Family 3. Elaphomycetace^. The fruit bodies of the fungi of
this family are subterranean with a distinct, mostly thick peridium
whose surface is marked by a more or less strongly developed rind. The
asci borne within the closed fruit body are irregularly arranged and
united into large groups, which are separated by radially arranged vein-
like masses of sterile hyphae. The asci are spheric, or pyriform, and
mostly eight-spored. The whole spore-bearing interior of the fruit
body, when ripe, is transformed into a powdery mass with the sterile
hyphae remaining as a number of capillitia-Hke threads. There is no

spontaneous opening of the fruit body at maturity. The family in-
cludes a single genus, Elaphomyces, which comprises about twenty-two
species, found mostly in northern Italy, in Germany and France, a few
in England, northern Europe and North America. Such species, as
Elaphomyces papillalus, E. atropurpureus from the oak and chestnut
woods of northern Italy, E. mutabilis with a silvery-white mycelium
growing in the oak, beech and birch woods of northern Italy, France
and Germany, E. citrinus with an orange-yellow mycelium, also from
northern Italy, all have delicate thin rinds which become wrinkled
when and belong to the section Malacodermei. The section Sclero-

dry,
dermei includes those species with compact brittle rind, which is not
wrinkled when dry. Here belong E. maculatus with strongly de-
veloped, green myceUum, surface of fruit body blackish brown with
greenish markings, found in the oak forests of northern Italy, French
1 Johnson, James: Host Plants of Thielavia basicola. Journ. Agric. Res.,

vii: 289-300, Nov. 6, 19 16.
^ Brierley, William B.: The Endoconidia of Thielavia basicola; Zopf, W.,
Vnnals of Botany, xxix: 483-491, with i plate, October, 1915.

Jura and the Tyrol. E. cerviims, which is found under oaks, beeches
and pines in Europe and North America, has a fruit body the surface
of which is brownish yellow, or reddish brown, and is covered with
numerous pyramid-shaped projections. The inner layer of the peri-
dium of this species is not veined like E. variegaius, another widely
distributed species throughout Europe. The fruit bodies of the last
two species are frequently parasitized by Cordyceps ophioglossoides

and C. capitatus (see ante, Fig. 21).
—
Family 4. Terfeziace^. The fruit bodies of the fungi of this
family are more or less deeply subterranean, tuber-like, infrequently
galleried {Hydnobolites). The fruit bodies differ from those of the
preceding family in that the interior spore-bearing portion does not
break down into a powdery mass, hence there is no so-called capillitium,
and as in that family the fruit body does not open spontaneously.
The terfas, or kames, of arid Mohammedan countries belonging to the
genera Terfezia and Tirmania were known
Greeks and Romans.
to the
The and associated with the roots
species of Terfezia are found under
of the herbaceous or shrubby forms of Artemisia, Cistus and Helian-
themum. A North African terfa, Terfezia conis, is found in the moun-
tain forests of pine and cedar and in the sands of Sardinia from March
to April. The desert terfas include T. Boudieri, T. Claveryi, T.
Hajizi and Tirmania ovalispora. Duggar,^ an American mycologist,
has gathered these fungi at the base of Artemisia herba-alba found
growing in the sandy soil of small oueds, or stream beds, in southwestern
Algeria. They by the breaking of the soil surface and are
are located
dug out -by the Arabs with a pointed stick. They form a valuable
food, as they are rich in protem.
—
Family 5. Tuberace^. General reference has been made to the
members of this family in a description of the special ecology of the
EUMYCETES. The mycelium of the truffles is well developed and
septate, producing mostly subterranean, tuber-like fruit bodies, which
have more or less numerous chambers lined with the ascigeral tissue
supported by sterile hyphae. The asci, which are arranged irregularly
in the ascigeral tissue, are one to eight-spored. The ascospores are
unicellular, and in the truffles {Tuber) usually spiny. The mycelium
is subterranean and is connected with the roots of coniferous and
broad-leaved trees forming the so-called mycorrhiza. The simplest
' DuGGAR, B. M.: Mushroom Growing, 191 5: 207-217.
—
152 MYCOLOGY
Vj
'm^^^m^-
Fig. 52. A, Tuher ceslivum frtiit-body; B, Tuber magnatiim fruit-body; C Tuber
,
brumale f. melanosporum, section through fruit-body; D, Tuber excavalum, section of

fruit-body; E, Tuber ceslivum f. mese7!tericuni, piece of fruit-body near pcridium en-
larged; G, piece of Tuber excavation enlarged;. H, Tuber rufum, fruit-body magnified
showing asci and ascospores; J, Tiihrr lirumalc, ascia with spores; K, Tuber magnalum,
ascus with spores. {See Die natiirlidicu Fjhnizenfamilien I. i, p. 287.)
SAC PUNGI CONTINUED 1 53
fruit body in the subfamily Eutuberine.^ is found

Genea liispidula in
where forms a hollow sphere with definite opening. Generally, it is
it
provided with a system of tubes, passageways or galleries, which vary

in their arrangement in the different genera. These galleries are hollow
in some, in others filled with hyphje, constituting the vencs externce.
The supporting hyphte between these passageways constitute the
sterile
vena inferncB. In the subfamily Balsaminace^, the fruit body has a

single, hollow chamber, or numerous hollow closed cavities. The
ascigeral layers constitute the walls of these chambers.
The fungi of the genus Tuber (Fig. 52) are of the most interest
economically, as several species, such as T. CBstivum (Spring), T.
brumale, T. melanosporum (^Winter), T. uncinatum (Autumn), T.rufum
are edible, and are known as truffles (Fig. 52). These species occur in
deciduous woods of north Italy, France and Germany and elsewhere
in Europe. They are gathered for food by men (rabassier), who make
a livelihood by selling the truffles for immediate use, or for canning
purposes. As the fruit bodies emit a characteristic odor, they are
located by the aid of specially trained dogs, and pigs, whose keen scent
enables them to find the underground fruit bodies. As they are dug
up, the animal is rewarded by his master with some other attractive
morsel of food, and the newly discovered truffle is placed in a leathern
pouch slung over the shoulder of the rabassier. The tin cans in which
the tfuffles (Tuber melanosporum in Perigord mainly) are preserved
for shipment to all parts of the world are usually labeled with a state-
ment as to the contents of the can, and with a hunting scene, where the
man and his truffle dog prominently figure.
Near here should be placed the family Myriangiaceae repre-
sented by the genus Myriangium with three species of wide distribu-
tion. This family has been monographed by von Honel.^
ivoN Honel: Sitzungsber. Math. Naturw. Klasse k. Akad. Wiss. Wien.,

118, Abt. i: 349-376, 1909-
;
CHAPTER XVII
MILDEWS AND RELATED FUNGI
Suborder D. Perisporiineae. — The mycelium of the fungi which
belong to this suborder is filamentous, superficial, light- or dark-
colored, rarely forming a stroma. The fruit bodies are superficial,
spheric to egg-shaped without a pore and break up irregularly. Peri-
thecia are usually dark-colored and in many cases surrounded by
accessory hyphae, or suffulcra. The asci are spheric, egg-shaped, or
elongated, and range within the closed perithecia from one to many
in number. Paraphyses are usually absent. The following families
are recognized:
A. Perithecium spheric, poreless or breaking irregularly at the top.

(a) Aerial mycelium white, perithecium with appendages or suffulcra
accessory spores belonging to the genus Oidium.
I. Erysiphaceae.
(b) Aerialmycelium absent, or dark-colored, perithecia without

appendages or suffulcra, accessory spores not belonging to
Oidium.
2. Perisporiace^.
B. Perithecium peltate flat, opening at top by a round pore.
3. MlCROTHYRIACE^.
Family i. Erysiphace^. —
The fungi of this family are popularly
called "white" or "powdery mildews." During the summer their
conidial fructifications (Oidium) are found on hops, maples, peas,
rosesand vines imparting to the surface of the host a dusty appearance,
due to the white conidiospores. Later in the summer, the globular
dark brown, or black, perithecia appear and these are provided usually
with appendages, or suffulcra, which are frequently branched in a way
characteristic of the different genera of the family. The white
mycelium upon which the fruit bodies arise is truly parasitic, for short
haustoria are formed which pierce the wall of the epidermal cells, and
swell out into a bladder-like form for absorptive purposes. The haus-
i.'54
MILDEWS AND RELATED FUNGI 1 55
toria are confined to the epidermal cells in all of the genera of the
family except Phyllactinia, which forms special hyphal branches which
enter the stomata, penetrate the intercellular spaces of the leaves and
finally send haustoria into the cells of the loose parenchyma. With
the exception of these haustoria, the mycelium of the "powdery
mildews" is entirely superficial. The conidial forms of the different
fungi of the family were classified formerly under the name of Oidium,
but with a more detailed knowledge of their life history, this name has
been relegated to the synonymy. The conidiospores, which are formed
in greatnumbers, are carried by the wind, or by snails in the case of
Erysiphe polygoni on plants of Aquilegia and are capable of immediate
germination on reaching the epidermis of a suitable host plant, the
germ-tube penetrating the outer wall of some epidermal cell. True
sexual reproduction has been discovered in some of the mildews by
R. A. Harper, thus verifying the earlier observations of de Bary.
Sphcerotheca Castagnei serves to illustrate the process. The oogonium
and antheridium, which are formed where two neighboring hyphse
approach, each contains a single nucleus. The cell wall between these
organs is dissolved at the time of fertilization and the male and female
nuclei unite and a fresh wall is laid down between the two organs.
Now the wall of the future perithecium begins to form by the develop-
ment of a number of upright hyphal branches around the oogonium,
forming a pseudo-parenchymatous tissue, while other branches later
absorbed grow into the interior of the developing perithecium, while
the outer wall cells become flattened and darker in color. The fol-
lowing growth takes place in Sphcerotheca, which develops only a
single ascus. The carpogonium elongates, divides and a curved row
of five or six cells is formed. The penultimate cell of this row contains
two large nuclei, while the other cells of the row have one nucleus each.
The young ascus develops from this penultimate cell in which the two
nuclei fuse followed by a rapid increase in size of the ascus, which presses
against the inner wall cells of the perithecium and absorbs them.
The nucleus of the ascus finally divides three times, producing the
nuclei of the eight ascospores, which subsequently are formed by free
cell formation. From the half-grown perithecium there arise apical,
equatorial or basal hyphae which grow out as the appendages, or
suffulcra, which in Phyllactinia are acicular and bulbous at the base
(îg- 53)) in Uncinula hooked at the apex and in PodosphcBra and Micro-
156 MYCOLOGY
Fig. 53. — Mildew of chestnut leaves due to Phyllaclinia corylei with ascus and
perithecium to left. (Martic Forge, Pa., Nov. 2, igiS-)
MILDEWS AND RELATED FUNGI 157
sphara (Fig. 54) dichotomously branched. These appendages prob-

ably assist in the distribution of the perithecium, serving to attach
the perithecia to plants, if wind-borne, or to the bodies of insects by
which they are carried to other plants. The number of asci found in a
perithecium and the number and character of the spores in the asci
vary generically (see Appendix VIII, pages 721-726).
As the fungi of this family are especially suitable for systematic
study, a key is given not only of the principal genera, but also of the
anth^
Fig. 54.— Lilac mildew, Microsphara alni. A, Perithecium with appendages;

B, perithecia showing asci (a); C, ascus with ascospores; D, conidiophore (cph),
bearing conidiospores {c.s.); E, beginning of fertilization; anth, antheridium; car,
carpogonium; F, later stage of fertilization showing the fusion of two nuclei (/).
(From Gager with E and F after R. A. Harper.)
principal species of the different genera. These keys (p. 721) have been
taken from a monograph of the Erysiphace^ by Ernest S. Salmon, pub-
lished in 1900, as vol. ix of the Memoirs of the Torrey Botanical Club, to
which the mycologic student is referred for detailed descriptions of the
various species. The material for the systematic study is easily kept
in the dry condition and the perithecium can be studied in situ on the
dried leaf or other plant parts, and later treated with weak alcohol
158 MYCOLOGY
to remove the air, washed and mounted permanently stained, or

unstained in acetic acid with a ring of asphalt, or in glycerine jelly
for a study of the asci and ascospores.
For a study of the distribution
of the haustoria and for a detailed examination of the sexual organs,^
small pieces (2 by 4 qcmm.) of hop leaves on which myceha of the
mildew (Sphcerotheca) are found in various stages of development
should be fixed in weaker Flemming's solution, as described by Zim-
mermann on page 178 of his "Botanical Microtechnique," and then
hardened in alcohol and carried through to paraffin. The sections
should be cut 5 to 7.5^ thick stained with safranin (one to one and
one-half hours), gentian-violet (one-half to one hour), and orange
G. (quickly), then treated with absolute alcohol, cleared in oil of cloves
and mounted in balsam.
The members of
material for systematic study should be handed to
the class in mycology, mounted and then studied as unknowns by the use
of the generic and specific keys given in Appendix VIll, pages 721-726.
—
Family 2. Perisporiace^. The aerial mycelium of these fungi
is superficial black, filamentous, or wanting, or rarely as a firm stroma.
The perithecia are situated on the aerial myceUum, or on the stroma.
They are black, + spheric, rarely elongated, poreless, or weathering ir-
regularly at the apex and without appendages. The wall is mostly
membranous, or brittle. The asci are clustered and mostly elongated.
The shapes of the spores are various. Paraphyses are usually wanting,
and are present in only a few cases.
The genus Scorias has been described incidentally in a foregoing page
(72). It is represented in America by a single species, spongiosa, ^h.\c\\
liveson beech twigs and leaves associated with some species of wooly
aphis, or on the ground where the droppings of the aphis in the form
of honey-dew have collected. Its mycelium is greenish-black, much-
branched, rigid, septate and the hyphae are glued together by an
abundant mucilaginous substance forming a loose spongy mass, bearing
an abundance of pyriform, coriaceous perithecia, which enclose narrow,
thick-walled, eight-spored asci. Elongate pycnidia and perithecia are
also frequently seen.
Family 3. —
Microthyriace^. The mycelium of the fungi of this
family is superficial and dark in color. The perithecia are superficial
1 Harper, R. A.: Die Entwickelung des Peritheciums bei Sphcerotheca Castagnei,
Bericht. der Deutsch. Bot. Gesellsch., xiii, Heft. 10: 475-481, 1895.
— ,
shield-shaped, unappendaged, black, membranous to carbonous

formed of radiating chains of cells. The asci are four- to eight-spored
short and associated with paraphyses. Two fungi which attack the
cofifee plant are the most important pathogenic species of the family:
Fig. 55. A-D, Nectria cinnabarina. A, Stroma of conidia and fruit-bodies of

fungus; B, stroma in section; C, ascus; D, mycelium with conidiospores; E, F, Neclria
dilissima; F, conidia layer; G, H, Nectria sinaplica; G, ascus; H, pycnidia-like layer.
J, Nectria inaiirita; K, Neclria oropensoides coremium. {See Die natiirlichen Pflanz-
enfamilien I. i, p. 35 7-)
and Microthyrium cofcB. There are twenty-

Scolecopeltis aeruginea
one genera, and more than 300 species not well understood.
Suborder E. Pyrenomycetiinaee.— The mycelium is always present
in these fungi. The perithecia are either located upon the substratum,
i6o MYCOLOGY
or in the substratum, and are mostly spheric. A wall (peridium) is
present inclosing the clustered eight-spored asci which arise from the
interior basal part of the perithecium. The perithecium opens by an
apical mouth or pore and is either isolated or imbedded in a stroma
which takes manifold forms. The formation of conidiophores and
conidiospores varies in the different families and genera. Sometimes
a distinct conidial layer is formed; at
other times the conidiospores are
formed in pycnidia. The suborder
includes many saprophytic and para-
sitic fungi found upon plants and
animals.
Family i. Hypocreace^. —The
perithecium of these fungi is spheric
and opens terminally by a definite
pore. In color, it may be pale,
sprightly colored, or colorless, never
black. Hypomyces with sprightly
colored perithecia arises from a thick
crust-hke stroma. It lives parasitic-
ally on a number of different fleshy
fungi. For example, Hypomyces
lactifluorum transforms a species of
Lactarius into a cinnabarred growth
roughly resembling a toadstool and
without gills, while the original color
of the host is completely lost in the
higher color produced by the parasite.
Nectria without stroma has its peri-
Fig. 56.— Ergot {Claviccps pur-
thecia developed on the surface of the
purea) on rye head. {After Clinton,
G. P., Rep. Conn. Agric. Exper. Stat., N. cinnabarina is a par-
substratum.
1903-)
on various deciduous trees (Fig.
asite
55). Its conidial form known as Tubercularia vulgaris produces flesh-

colored eruptions through the bark of various host plants. Nectria
ditissima grows on the beech. Polystigma has a crust-like stroma on
the leaves of trees of the genus Prunus, while Epichloe typhina con-
fines its parasitic attack to grasses upon which it develops orange-
yellow stroma. The genus Cordyceps consists of species which live
—
MILDEWS AND RELATED FUNGI [6l
Fig. 57. A, Balansia claviceps on ear of Paspalum; B-L, Claviceps purpurea;

B, sclerotium; C, sclerotium with Sphacelia; D, cross-section of sphacelial layer; E,
sprouting sclerotium; F, head of stroma from sclerotium; G, section of same; H,
section of perithecium; J, ascus; K, germinating ascosnore: ^. conidiosnores pro-
duced on mycelium. (See Die nalurlichen Pflanzenfamilien I. i, p. 371.)
l62 •
MYCOLOGY
parasitically on insects and their larva and some in subterranean fungi.

The myceHum kills the insect or larva and mummifies it. Out of the
host grow conidiophores (Isaria) in early stages of development, and
later stalked stroma, in which on enlarged terminal portions the per-
ithecia with asciand ascospores are found. C. militaris and C. cinerea
occur on insects, or insect larvae. C. sinensis is found on caterpillars
in eastern Asia, while C. ophioglossoides grows on the fruit bodies of
species of Elaphomyces (see ante, page 70) (Fig. 21). Claviceps is a
genus of fungous parasites found in the developing caryopses of various
grasses. Its conidial stage was formerly known as Sphacelia. Claviceps
purpurea and C. microcarpa are important species and their life his-
tories will be described in the third part of this book. As ergot, the
purpurea are used in medicine (Figs. 56 and 57).
sclerotia of Claviceps
Fifty-seven genera and three doubtful ones are recognized and described
in Engler's Die natiirlichen Pflanzenfamilien.
Family —
Dothideace^. This family comprises twenty-four
2.
genera among most important of which is Plowrightia (Fig. 22)

the
and Phyllachora. The fruit bodies of these fungi is spheric with definite
mouth and without distinct peridium, as they are found imbedded in a
black stroma. Plowrightia includes twenty species of fungi, which
form stroma in the interior of host plants, and which break through to
the surface, and form pimples in the center of which the opening to
the perithecium is found. The spores are egg-shaped, two-celled,
hyaline, or bright-greenish. Plowrightia ribesia is found on dried
twigs of species of currants Ribes in Europe and North America.
P. virgultorum occurs on brick in northern and middle Europe, P.
Mezerei grows on dead branches of Daphne in middle Europe and Italy.
P. insculpta is found on dried branches of Clematis vitalba in Bel-
gium, France, Germany and Italy and P. morbosa is the cause of black-
knot of the cherry and plum (Prunus) and will be described subsequently.
Phyllachora is a large genus of some 200 species found mostly on the
leaves of various plants; P. graminis is the commonest species of cos-
mopolitan distribution on grasses and sedges. The warty spot of clover
is Phyllachora trifolii.
Family 3. Sordariace^. —The perithecia in this family are

superficial, or deeply sunken in the substratum and often break through
at maturity. The stroma is usually absent, but when it occurs the
perithecia are sunken with projecting papilliform beaks. The perithecia
are thin and membranaceous to coriaceous, slightly transparent to

black and opaque. The asci are usually very delicate, surrounded
by long paraphyses, The dark-colored
or intermingled with them.
spores are one- to many-celled, surrounded by a hyaline gelatinous
envelope, or ornamented with hyaline gelatinous spicula. The
SoRDARiACE^ are entirely saprophytic and grow on manure, hence,
they are coprophilous fungi. Special mechanical devices are shown by
the asci for eruptive spore discharge and the distance to which the
spores are shot may be between 5 and 9 cm.^
—
Family 4. Chêtomiace^. This is a small family of two genera,
ChcBtomium and Bommerella, which are found on waste paper, manure
and on small living fungi, which resemble the fungi of the family
PerisporiacecB, if the mouth to the perithecium is wanting. Bom-
merella has three-cornered ascospores. The perithecia of such forms
as ChcBtomium spirale and C. crispatum are provided apically with
masses of spirally wound ha^'rs.
Family 5. Sph^riace^. —This important family includes parasitic,

or saprophytic fungi showing exceptional diversity on dead parts.
They have rounded perithecia with definite opening. The peridium is
evident, mostly dark-colored, membranous to leathery never fleshy,
usually free from the substratum, or more or less depressed. A
stroma may or may not be present. Some authors include a number
of families which perhaps may be subordinated here and ranked as
subfamilies. Rosellinia quercina is a disease of oak seedlings. Myco-
sphcerella fr agarics is the cause of leaf spot of strawberry; M. strati-
formans produces leaf-splitting blight of sugar cane. Gulgnardia

Bidwellii is a most important parasite, being responsible for the black
rot of the grape and G. vaccinii causes cranberry scald. Apple
scab and pear scab are due to the attack of Venturia pomi and Venturia
pyrina. A serious disease of sycamore leaves in the spring known as
anthracnose is caused by Gnomonia veneta.
Family 6. Valsace^. —The stroma of these fungi is black and is
formed in the substratum which is more or less altered. The peri-

thecia have a regular border and take various forms in the different
genera. The asci and long-stalked, alternating with
are cylindric
paraphyses. Pycnidiospores are formed in pycnidia and conidiospores
1 Griffiths, David: The North American Sordariace^. Memoirs of the
Torrey Botanical Club, xi, No. i, May 7, 1901.
1 64 MYCOLOGY
on definite conidiospores. Of the ten genera of the family, the

genera Valsa and Diaporihe are the most important. Both genera
include about 400 species, which are most saprophytic in wood and the
bark of woody plants. Valsa oxystoma is the cause of the disease and
death of the branches of Alnus viridis in alpine regions; Diaporthe
farinosa grows on the branches of the linden, Tilia americana in North
America and D. eucalypti on Eucalyptus globulus in California.
—
Family 7. Melogrammatace.e. The stroma are mostly like those
of the genus Valsa and rarely like those in Diatrype. They are hemis-
pheric and are formed beneath the bark and later break through to the
surface, where they are more or less isolated. The perithecia are
imbedded in the stroma. Conidial fructifications are formed on the
surface of young stroma, or pycnidiospores are produced in pycnidia.
The most important genus of this family is Endothia, which is repre-
sented by the Chestnut-blight fungus E. parasitica, which lives in the
cambium and inner bark of chestnut trees causing a final girdling of
the branch and the death of the part beyond the girdled area. It has
caused untold injury to the forest groves of America, where the chest-
nut tf-ee abounds, and its morphology and its ravages will be described
subsequently.
—
Family 8. Xylariaceê. The stroma of these fungi is developed
strongly and is frequently upright and branched. The perithecia
are borne in the branched club-shaped portions of the fruit bodies.
Early in their growth the surface is covered with conidiospores. The
ascospores are unicellular and blackish-brown. The genus Num-
mularia, which includes forty species, is represented typically by
N. Bullardi, which causes black charcoal-like eruptions on thick
branches of the beech, Fagus. Ustulina, with nine species, includes
U. vulgaris found on old stems of broad-leaved trees and Hypoxylon
with about 200 species is confined mostly to damp wood and old
tree stumps. Xylaria digitata, one of the 200 species of that genus,
grows on old wood, and X. polymorpha on old tree stumps. This
family completes the list of pyrenocarpous fungi.
—
Suborder F. Discomycetiineae.^ The discomycetous fungi have a
filamentous mycelium. Reproduction is by the union of two hyphal
branches either of similar size, or differentiated into oogonia and anthe-
ridia. The fertilized egg cell either develops directly into an ascus,
or it develops ascogenous hyphas from which the asci are formed.
The apogamous formation of fruit also occurs in this suborder. The

asci are united into definite, usually fiat layers, which are
open in
fruit bodies known as apothecia. Conidiospores are also found in
some of the forms and the conidiophores are of diverse character.
The asci are usually eight-spored. The fungi of this suborder are
either parasitic, or saprophytic in habit, and a few of the fleshy members
of the family Pezizace^ are edible.
Family i. Hysteriaceê.^ —The apothecium is elongated and the
opening is a long wide cleft between the approaching walls of the
apothecium, so that the ascigeral layer is exposed at the time of the
spore discharge.
Some species of the genera Lophodermium and Hypoderma are
dangerous parasites of leaves; for example, L. pinastri attacks pine
leaves; L. nervisequum attacks the spruce tree; while Hypoderma
hrachysporum found on the white pine, Pinus strobus. Such genera
is
as Lophium, Hysterium, and Glonium include species which are sapro-

phytic on bark and wood.
Family 2. PHACioiACEiE. —The apothecium is rounded, seldom
elongated and its walls are separated through a star-shaped opening,
rarely a cleft-like opening, so that the ascigeral layer is fully open at
maturity. The family includes such parasites as Nemacyclus niveus
on coniferous needles; Rhytisma acerinum, which produces black tar-
like blotches on maple leaves; and R. salicimim, which causes similar
black areas on willow leaves. Several species of Trochila are found
on the leaves of different plants.
Family 3. Pyronemace^. —^The fruit body is placed on fine
hyphge or on a felt-like cushion of hyphae. At first it is spheric; later,
it is flatly expanded. The hypothecium is occasionally feebly de-
veloped, at other times strongly so. The peridium is poorly formed,
it is
or entirely absent. The most interesting genus is Pyronema. P.

confliiens has a fruit body i mm. across, and of a yellow or reddish
color. It is often found in spots where
have been kindled in the fires
woods. The and the method of its forma-

structure of the apothecium
tion following the sexual union of an antheridium and oogonium have
been described by Harper^ and the essential details have been given
on a former page of this book {ante^ pages 123 and 126).
^ Harper, R. A. : Sexual Reproduction in Pyronema confluens and the Mor-
phology of the Ascocarp. Annals of Botany, 14: 231-400, 1900
—
i66 MYCOLOGY
Family 4. Ascobolace^. —The apothecia of the fungi of this

family are unstalked. They and grow up on manure.
are superficial
The peridium is mostly thin, or wanting, and the hypothecium, which
is well developed, consists of rounded parenchyma-Uke cells. In
Ascoholus, the ascospores are discharged from the asci by a squirting
Fig. 58. A, B, Lachnea ^culellala. A, Habit, B, ascus with paraphysis; C, D,

Lachnea hemispharica; C, habit; D, ascus with paraphysis; E, Sarcosphara arenosa
habit; F, G, Sarcosphara coronaria; F, ascus; G, habit; H, Sarcosphcera arenicola
ascus with paraphysis. {See Die nalurlichen Pjlanzenfamilien I. i, p. i8i.)
action,and this is accomplished probably by the pressure of the cell

wall upon the cell sap. The end of the ascus breaks open suddenly, the
ascus collapses, and the eight spores are discharged simultaneously
along with the cell sap. In Ascoholus, which is related to Pyronema,
the ascogonium is at first multicellular, but all the cells empty their
contenls into a single large one, from which the ascogenous hyphiB
then arise.
Family 5. Pezizace^. —^The apothecia of this family are saucer-

or cup-shaped, sessile from a mycelium which is
or stalked, arising
found in the substratum. The peridium and hypothecium consists
of rounded cells and they are of fleshy, or leathery consistency. The
asci, which are usually eight-spored, are separated by distinct para-
physes. The spores are Lachnea and Peziza are

usually hyaline.
the most important Lachnea scutellata (Fig. 58) has a
genera.
scarlet to vermilion-red cup, whose margin is beset with a fringe of
Fig. 59. — Saucer-shaped fruit-bodies oi Peziza re panda. (Photo by W. H. Walmsley).
large brown grows on wet sticks and logs in damp, or wet

bristles. It
L. hemisphcerica has a cup i to 4

places, especially at the water's edge.
cm. wide with a bluish-white to gray disk and with brownish outside
bristleswhich fringe the margin of the apothecium. It grows on much-
decayed wood. Peziza aurantia, which is found in the fall in woods,
and is edible, has a bright orange cup i to 5 cm. wide, powdery outside.
At first, it is cup-shaped, then saucer-shaped and irregular. It is
stemless, or nearly so. The and strongly

spores are clear, elliptic
netted. A woodland form, P. coccinea, is scarlet in color and suggests
a wine glass in its stalked apothecium. P. badia grows on the
ground in grassland and woodland, and is also edible. It has a
i68 MYCOLOGY
dark brown to paler brown apothecium, i to 4 cm. across and almost

stemless. P. ceruginosa is a stalked, green form whose mycelium pene-
trates the wood of beeches and oaks and imparts to them a copper-
green color, which makes it valuable for the manufacture of the famous
"Tunbridge ware." The attempt has been made to extract the pig-
ment, or to manufacture it synthetically for use as a shingle stain, but
without much success. P. Willkommii produces on larch trees a disease
known as larch canker. Other species of Peziza grow on bark (Fig. 59),
horse and cow manure, and are, therefore, typically coprophilous.
—
Family 6. HelotiacE/E. The apothecia in these fungi are super-
ficial from the beginning and rarely arise by break-
ing through the substratum. Sometimes they de-

velop from a sclerotium {Sclerotinia). In texture,
they are waxy, leathery and thick, and stalked, or
unstalked, smooth or hairy. The asci are eight-
spored. The spores are round, elongated, or fila-
mentous, and one to eight-celled, hyaline. The

paraphyses are filamentous. The fringe cup,
Sarcoscypha floccosa, has a slender, white, hairy
stem, I to 3 cm. long by 2 to 3 mm. wide, and
bearing an apothecium 4 to 10 mm. wide with a
scarlet disk, so that the whole fruit body is goblet-
shaped. The outside of the cup is covered densely
with long white hairs forming a fringe at the margin.
The spores are clear and elliptic 20 by 11//. The
-Scleroiinia fringe-cup fungus grows on decaying twigs from
(After
spring to autumn. Sclerotinia is the most impor-
tant genus economically. It includes about forty
species. The apothecium arises from a sclerotium. Sclerotinia haccarum
forms sclerotia in the fruits of Vaccinium myrtillus; S. urnula (Fig. 71)

in those of Vaccinium vitis-idcea. Sclerotinia Fuckeliana forms sclerotia
on the grape-vine. form was long known as Botrytis cinerea.

Its conidial
Sclerotinia sclerotioriim (Fig. 60) is parasitic and pathogenic on a number
of cultivated plants, such as beets, and bears its sclerotia forming on the,
subterranean parts of these host plants. The black disease of hyacinth
bulbs connected with the growth of Sclerotinia hulhosum. Apples,
is
pears and stone fruits are attacked by S. fructigena. S. libertiana.

causes lettuce drop. S. trifoUorum is responsible for the stem rot of
clover. Other fungi without sclerotia are parasitic and destructive.

Such are Dasyscypha Willkommn, the cause of larch canker. D.
Warburgiana is parasitic on cinchona in the tropics. Such genera as
Coryne, Helotium, Lachnum and Rutstroemia are saprophytic on wood.
—
Family 7. Mollisiace^. ^The fungi of this family differ from
those of the preceding largely in texture, the former being tougher with
hyphal cells modified in a fibrous manner. The spores are hyaline.
Pseudopeziza is the only important germs with apothecium formed
its
beneath the epidermis, which is subsequently ruptured with the pro-

trusion of a shallow fruit body. The asci show eight unicellular spores.
Pseudopeziza medicaginis is the cause of alfalfa leaf spot. Ps.
ribis causes anthracnose of currants.
The remaining famiUes of the suborder are Family 8, Celidiace^,
Family 9, Patellariace^., Family 10, Cenangiace^.
—
Suborder G. Helvelliineae. This suborder includes fungi with
a well-developed mycelium which is filamentous and largely functional
for nutritive purposes. From this mycelium, which penetrates the
substratum, arises a fleshy, waxy or gelatinous fruit body, which usually
possesses a stalk upon which is raised an expanded portion; sometimes
club-like, in other forms constituting a distinct pileus. The expanded
part, which may be smooth and gelatinous, wrinkled or with variously
contorted folds, or of deep pits separated from each other by anastomos-
ing ribs, is covered with the ascigeral layer, which consist of asci and
paraphyses standing on end-like pahsade tissue. The asci are typically
eight-spored, rarely, two-spored, and open at the apex through the
removal of a lid, or through a tube-like mouth. The ascospores are
unicellular, or multicellular.
FAivnLY I. Geoglossace^. —The fruit body is fleshy, waxy, or
gristly, and is separable into a stalk, or stipe, and an enlarged fertile
portion, the pileus, which is club-shaped or knobbed, and its color is
some shade of yellow, green, or black. The asci are club-shaped,
opening by a pore at the apex. This family includes twelve genera,
and it has been carefully monographed by Massee.^
Geoglossum hirsutum is an American ground form with pileus flat
and black, 2 to 3 cm. long and i to 2 cm. wide. It is wrinkled and
hairy (Fig. 61). The stem is 6 to 8 cm. tall, black soUd and hairy.
'Massee, George: A Monograph of the Geoglosseae. Annals of Botany, ii;
225-306 with 2 plates, 1897.

— ,
lyo MYCOLOGY
The and many-celled, loo to 120 by 4 to

spores are brown, very long
7/1. American species, grows on the ground
G. glutinosum, another
among the grass. It is black and smooth with the ascigerous portion
one-third the entire length of the fruit body and in shape oblong-
lanceolate, slightly viscid. The upper portion passes imperceptibly
into the stalk. The spores are eight in number, arranged parallel to
each other with obtuse ends and three-septate, 65 to 75 by 5 to 6m,
and brown in color.
Leotia chlorocephala is a fungus found in West Virginia, New Jersey
and Pennsylvania. It is cespitose
in habit and grows in mixed woods
on moist ground, from July until
late frosts. It is green and has a
gelatinous appearance. The pileus
is depressed globose, more or less
wavy and with an incurved border,
in color a dark verdigris-green. It
is Another species, L. lubrica,

ecUble.
is found on the ground in woods from
North CaroUna and Minnesota to

Massachusetts. It is yellowish, olive-
green with an irregular hemispheric,
inflated, wavy cap.
Family 2. HELVELLACEiE. The —
fruit body in these edible .fungi is
Fig. 61. Geoglossum liirsiUum. fleshy and divided into a hollow stalk
A, Appearance of fungus; B, asci with
paraphyses; C, spore. A, natural and ascigerous expanded portion.
size; B, 300/1; C, 400/1. {Die naliir- The upper part is cap-Hke and
lichen Pflanzenfamilien I. i, p. 165.)
covered externally by the ascigeral
layer. The asci are club-shaped and open by the lifting off of a distinct
hd. The spores are ellipsoid, colorless, or bright yellow and smooth.
Five genera are included in the family: Morchella, Gyromitra, Verpa,
Cidaris and Helvella. This family includes the largest of the sac fungi.
Some species of Gyromitra weigh over a pound and forms of Morchella
may grow a foot tall. The cap of Morchella is more or less deeply ridged
crosswise and lengthwise and has a delightful odor. The broad stem
Morel, Morchella crassipes, has a cap 4 to 10 cm. tall and 3 to 6 cm.
wide at the base, in color tan to tan-brown, with deep pits and wavy to
irregular ridges, the whole cap being more or less conic. The stem is
3 to 12 cm. by 2 to 6 cm., white and hollow. The spores are elliptic,

clear, smooth, 20 to 22 by 10 to 12/x. M. esculenta, the common
Morel, has a cap 3 to 7 cm. tall and 2 to 4 cm. wide, of a yellowish-
brown to brown color, covered with very regular ribs with a blunt
edge. The spores are smooth, elliptic, clear, 14 to 22^ by 8 to 14/i.
It grows on the ground in woods and forest openings, and is a delicious
morsel.
Gyromitra has a more irregular cap more or less inflated and folded,
the edge united in places with the stem. G. esculenta has a rounded
lobed pileus, irregular, gyrose-convolute, smooth and bay-red. Its
stem is stout, stuffed, or hollow. The ascospores are elliptic, yellow-
ish, 20 to 22M long. It grows in wet ravines, or springy places in the
vicinity of pine groves, or pine trees. G. brunnea is brown in color and
is figured by Clements in his "Minnesota Mushrooms," page 143.
Verpa digitalijormis grows on ground in woods. It has a brown, or
dark brown, smooth, bell-shaped cap with a long finger-like stem,
beneath, hence the specific name. Verpa bohemica is the "ribbed
verpa" and is delicious eating.
The cap in the genus Helvetia hangs loosely over the stem and it is
saddle-shaped more or less lobed. The stem is ribbed. The ascigeral
layer is confined to the upper side of the cap. All of the species are
edible. Helvella crispa is a common species and has been collected in
West Virginia, Pennsylvania and New Jersey. It is white or whitish
in color, whileH. lacunosa is gray to almost black.
Family —
Cyttariace^. This family is represented by the
3.
single genus Cyttaria with a tuber-like stroma in which the apothecia

are sunken. The stroma, which arises on the antarctic beech, Notho-
fagus, in South America and Tasmania, is stalked. The asci are cyhn-
dric and eight-spored. The spores are ellipsoidal and hyaline. The
paraphyses are filamentous, breaking down into mucilage. The cylin-
dric asci bear elliptic hyaline spores. Six species have been described
from Patagonia, Tasmania and Terra del Fuego.
—
Family 4. Rhizinace^. The fruit bodies of the fungi of this
small family are stalkless and they are fleshy and waxy in consistency.
Four genera are included.
Suborder H. Laboulbeniineae. —We owe our knowledge of these
eccentric or singular fungi to four botanists: J. Peyritsch, G. Lindau,
172 MYCOLOGY
Roland Thaxter and J. Faull. They are parasitic on insects, mostly

which live in moist situations and are long-lived and hiber-
beetles,
nating. They are often highly specialized, as to the parts of the
insect on which they grow, occurring only on certain joints of the legs
and on certain legs of the host. The vegetative mycelium is very much
reduced, consisting of one to a few cells, which are attached to the body
of the insect and their usually minute size renders them difficult of
study. The host is not destroyed nor even inconvenienced by these
fungi which appear as minute, usually dark-colored, yellowish bristles
or bushy hairs projecting from the chitinous integument of the insect.
Stigmatomyces BcbH lives parasitically on house flies. The bicellu-
lar spore with its mucilaginous coat becomes attached at its lower end.
The upper cell develops an appendage which bears a number of unicel-
lular flask-shaped antheridia from which the naked spermatia are shed.
The lower cell divides into four cells which represent the female repro-
ductive organ, where the carpogonium, or egg cell develops a trichogyne
to which the spermatia become attached. The three fundamental parts
of which these plants are composed are a main body, the receptacle;
one or more spore-producing portions, the perithecia; and lastly, one or
more appendages which, in the majority of cases, are associated with
the formation of the male sexual organs. The receptacle is that por-
tion of the fungus on which the appendages together with the perithe-
cia, or their stalk cells, are inserted. The sterile appendages, which
form dense tufts and sometimes are more conspicuous than the main
plant itself, serve to protect the delicate trichogyne which is subse-
quently developed. Sometimes, the primary appendage develops a
spine-Uke process. The male organs and male elements in the Laboul-
BENiACE^ may be designated as antheridia and antherozoids, the former
consisting of a single antheridial cell or a group of such cells, the latter
of a single naked, or thin- walled cell, so that the antherozoids are pro-
duced either endogenously or exogenously. Among the antheridia
which produce endogenous antherozoids we may distinguish the
simple and the compound. A simple antheridium discharges its
antherozoids through its special pore or opening, the compound an-
theridium consists of several antheridial cells each of which dis-
charges its contents into a common cavity from which they es-
cape. The female organs are formed from a segment of the lower
cell of the receptacle rarely from the terminal cell. The perithe-
cium, as in many other Ascomycetales, originates from a cell of the

receptacle situated below the female organ. The procarp consists of
three distinct parts: the trichogyne, the trichophoric cell and the part
lowest the carpogenic cell, which is fertilized and undergoes further
development. FaulP has shown in two species of Laboulhenia that after
the procarp mature the carpogonium and trichophoric cell become
is
continuous. Meanwhile, the nucleus of the carpogonium is succeeded

by two which are apparently daughters of the carpogonial nucleus, and
almost simultaneously the trichophoric nucleus undergoes division.
Later, a uninucleate trichophoric cell and a uninucleate inferior sup-
porting cell are septated off from the now four-nucleated fusion cell.
After further nuclear divisions a binucleate superior supporting cell and

sometimes a binucleate inferior supporting cell are cut off. The binu-
cleate ascogonium now begins to bud off asci, or divides into two asco-
genic cells, each of which contains a pair of nuclei. Up to this stage no
nuclear fusions have been observed. The nuclei of an ascogenic cell
divide conjointly, a daughter of each passing into a young ascus. This
process is repeated at the birth of every ascus. The pair entering the
ascus soon fuse. The fusion nucleus divides by a reduction mitosis
after a period of growth and the number of chromosomes is the same
as in other mitoses. There are two other mitoses prior to spore forma-
tion, and both are homotypic. The spores are delimited by the method
characteristic of the ordinary sac fungi. Each ascus in Stigmatomyces
BcBfi produces four spindle-shaped bicellular spores. In other genera
eight two-celled spores are formed. It is to be noted in closing that the
sexual organs of these curious fungi are similar to those of the red
seaweeds, Floride^. Thaxter^ has done more than any other botanist
to make this order known systematically.
Phylogeny oj Ascomycetales. —Atkinson in a philosophic discussion of
the phylogeny of the Ascomycetales suggests six series or lines of
development and his suggestions are incorporated in the accompanying
chart.
I. Apocarp line from Dipodascus-\\ke forms and by reduction.
1 Faull, J. H. : The Cytology of the Laboulbeniales. Annals of Botany,

XXV 649-654, July, 191 1. The Cytology of Laboulhenia chsetophora and L.
:
gyrinidarum. Annals of Botany, xxvi: 355-358, with 4 plates, April, 191 2.

- Thaxter, Roland: Contributions toward
a Monograph of the Laboulbeniaceae
part I, 1896; part II, 1908, Mem. Amer. Acad, of Arts and Sci.
174 MYCOLOCxY
2. Plectocarp line from Dipodascus-like forms, perhaps similar to

Monascus.
3. Perispore line arising from Monascus-Ukc prototype, before split-
ting of archicarp, or from Aspergillace^.
4. Pyrenocarp near Monascus-like prototype, Laboul-
line arising
BENiALES side near base, and some of the Mycothyriales as reduced
from Sph^riales.
Those who adhere to the behef that the AscomycAtales have
descended from the red algae interpret their belief in three ways: first,
sac fungi with highly developed trichogyne of the Collema type with cer-
tain red algae of existing forms; second, sac fungi with highly developed
trichogyne of the Polystigma type with hypothetic algae with trichogyne
representing the common original stock of both groups; and third, sac
fungi with simple generalized copulating gametes of the Gymnoascus

type with hypothetic algae having a simple procarp representing the
stock from which both groups started. It will be noted that Atkinson
Ascomycetales have been derived from
believes that the fungi of the
the simple Phycomycetes, and that thePROXOASCOMYCETES are der'ved
by descent and degeneration from such a primitive form as Dipodascus,
Endomyces Magnusii being the nearest known form to the generalized
condition seen in Dipodascus. The Euascomycetes are derived from
fungi similar to Monascus and Gymnoascus with generalized archicarp.
Six distinct lines as previously noted arise from these primitive forms.
Atkinson gives a chart which is purely provisional, and which suggests
the probable relationship of the principal groups to each other and to a
probable common ancestor.
GENERAL BIBLIOGRAPHY OF THE ASCOMYCETALES

Atkinson, Geo. F. : Phylogeny and Relationships in the Ascomycetes. Annals
of the Missouri Botanical Garden, ii: 315-376, February-April, 1915.
Barker, B. T. P. : The Morphology and Development of the Ascocarp in Monascus,
with 2 plates. Annals of Botany, xvii: 167, 1903.
Blackman, H. H. and Welsford, E. J. : The Development of the Perithecium of

Polystigma rubrum. Annals
Botany, xxvi: 761, 191 2, with 2 plates.
of
Brown, Horace T.: Some Studies in Yeast. Annals of Botany, xxviii: 197, 1914-
Carruthers; D.: Contributions to the Cytology of Helvella crispa, with 2 plates.
Annals of Botany, xxv: 243, 191 1.
Clements, F. E.: Minnesota Plant Studies: iv, Minnesota Mushrooms, 1910:
138-151-
Conn, H. W. Bacteria, Yeasts and Moulds in the Home, 1903, with 293 pages.
:
DuGGAR, B. M.: Mushroom Growing, 1915, pages 188-224, dealing with European
Truffles, Terfas and Morels.
Ellis, J. B. and Everhart, J}. M.: The North American Pyrcnomycetes, 1892,
pages 793, with 41 plates.
Engler, a.: Die Natiirlichen Pllanzenfamilien, I. Teil, i Abt. : 142-505 with
separate parts by Ed. Fischer, G. Lindau and J. Schroeter.
Faull, J. H.: The Cytology of the Laboulbeniales. Annals of Botany, xxv:
649-654, July, 191 1.
Faull, J. H.: The Cytology of Laboulbenia chajtophora and L. Gyrinidarum.
Annals Botany, xxvi: 325-355, with 4 plates, April, 191 2.
of
Eraser, H. C. I. and Ullsford, E. J.: Further Contributions to the Cytology of
the Ascomycetes, with 2 plates. Annals of Botany, xxii: 331, 1908.
Eraser, H. C. I. and Brooks, W. E. St. T.: Further Studies on the Cytology of
the Ascus. Annals of Botany, xxiii: 537, 1909.
Eraser, H. C. I. and Gwynne-Vaughan Mrs. D. T.: The Development of the
Ascocarp in Lachnea Cretea, with 2 plates. Annals of Botany, xxvii: 553, 1913.
Grant, James: The Chemistry of Bread Making, 191 2: 125-152.
Griffiths, David: The North American Sordariacese. Memoirs Torrey Botanical
Club, xi, 1901.
Jorgensen, Alfred: Microorganisms and Fermentation, transl. 3d Edition
by Alex. K. Miller and A. E. Lennholm, 1900, with 318 pages.
Kohl, Dr. F. G.: Die Hefepilze ihre Organisation, Physiologic, Biologic and Sys-
tematik ihre Bedeutung als Giirungsorganismen, 1908.
Klocker, Alb.: Fermentation Organisms: A Laboratory Handbook, transl. by
G. E. Allan and J. H. Millar, 1903, with 391 pages.
Lafar, Dr. Franz: Technical Mycology, transl. by Charles T. C.Salter. II, Part
I: 99-189: Part II: 191-481.

Massee, George: A Revision of the Genus Cordyceps. Annals of Botany, ix: i,
with 2 plates.
Massee, George: A Monograph of the Geoglosseae. Annals of Botany, 11:

225-301, 1897.
Massee, George: The Structure and Affinities of the British Tuberacese, with i
plate. Annals of Botany, xxiii: 243, 1909.

Massee, George: Text-book of Fungi, 1906: 261-313.
Salmon, E. S. On Endophytic Adaptation Shown by Erysiphe graminis.
: Annals
of Botany, xix: 444.
Salmon, E. S. : On Oidiopsis taurica, an Endophytic Member of the Erysiphaceae.
Annals of Botany, xx: 187, 1906.
Salmon, Ernest S. : A Monograph of the Erysiphaceae. Memoirs Torrey Botan-
ical Club, ix, 1900, pages 287, with 9 plates.
Stevtens, F. L.: The Fungi Which Cause Plant Disease, 1913: 113-297, with
bibliography.
Thaxter, Roland: Contributions toward a Monograph of the Laboulbeniaceie,
part I, Mem. Amer. Acad. Arts & Sci., 1896; part II, do., 1908.
1^5 MYCOLOGY
Bull. ii8, U. S.
Thon, Charles: Cultural Studies of Species of Penicillium.
Bureau Animal Industry, 1910.
Wager, Harold: The Nucleus of the Yeast Plant. Annals of Botany,
xii: 499-
540, with 2 plates, 1898.

Systematischen Botanik (2d
Wf.ttstein, Dr. Richard R. von: Handbuch der
Edition), iqii: 168-192.
.
CHAriER XVIII
BASIDIA-BEARING FUNGI (SMUTS)

ORDER BASIDIOMYCETALES
The fungi of this order have mostly a strongly developed mycelium,
multicellular and at times with apical growth. Sexual reproduction is
entirely absent, yet in the rusts, we find certain nuclear fusions which
are looked upon by some mycologists as of a sexual nature. The
characteristicmethod of reproduction is non-sexual by means of conidia,
which in the most primitive forms are of indefinite number,
while in the most highly differentiated forms the conidiospores are
definite in number two to eight, and are borne on special conidio-
phores known as basidia (basidium-ia). In many forms, the basidia
are arranged in definite parts of fleshy fruit bodies and in special layers
known as hymenia (hymenium-ia). Besides the conidiospores other
kinds of spores, known as chlamydospores, are formed. Zoospores are
entirely absent. The fungi of the order are either saprophytes, or
parasites, and occasionally, they are facultative saprophytes, or faculta-
tive parasites. None of them live in the water (nicht wasserbewohnend)
The Basidiomycetales do not follow the Ascomycetales in the direct
line of evolution of the fungi. They may be considered to parallel the
sac fungi. The group is supposed, in this regard, to represent the results
of extreme simplification; the sexual organs, if ever present, have in
the phylogenetic history of these fungi long since disappeared and
simple nuclear fusions function in all probability in lieu of the sexual
act.
Key to Suborders of the Basidiomycetales (After Stevens)
Chlamydospores at maturity free in a sorus, produced intercalary,

from the mycelium; basidiospores borne on a promycelium and resem-
bling conidiospores. i. Hemibasidii.
Chlamydospores absent, or when present, borne on definite stalks.
Basidia septate, arising from a resting spore, or borne directly on a
hymenium. 2. Protobasidii.
Basidia non-septate, borne on a hymenium. 3. Eubasidii.
177
178 MYCOLOGY
Suborder Hemibasidii. —The conidiophore, or more correctly the

basidium, arises from the chlamydospore and bears an indetinite
and usually large number of basidiospores. All cells of the mycelium
and the spores, as far as known, are unicellular. The position of this
suborder in the family tree of the fungi is uncertain. The majority
of the funguses are strictly parasitic on the higher plants, and their
mycelia live in the tissues of the same, mostly as intercellular parasites,
certain hyphae known as haustoria penetrating the interior of the host
cells. Infection of the host takes place, as a rule, very early and in
some cases at the time of seed formation, so that the parasitic mycelium
keeps pace with the growth of the host plants and at definite times and
places, such as anthers, ovaries and the like, which are mostly de-
formed, the spore-bearing portion of the fungous parasite appears.
The which are formed in such places, are known as chlamydo-
spores,
spores, and the mass of spores and diseased host parts are mostly
black and soot-like. The chlamydospores give rise to a promycelium,
which cuts off basidiospores. The basidiospores give rise either to
conidiospores, or they infect some host plant, if deposited upon it at
the susceptible time. Brefeld first suggested the name Hemibasidii for
the UsTiLAGiNACE^ and Tilletiace^ which he considered as repre-
senting the link connecting the lower fungi and the true BASIDIO-
MYCETALES. Two famiUes are recognized by mycologists, viz.,
USTILAGINACE^ and TlLLETIACE^.
—
Family i. Ustilaginaceê. ^The fungi of this family are all para-
sitic. They can be recognized readily by the outbreaks of dusty
material that they produce on certain parts of their hosts, when they
reach their reproductive stage. An important genus, Ustilago, the
type genus of the family, derives its name from ustio, a burning. The
smut wheat is called locally in England "bunt ear," "black ball,"
of
" dust brand" and " chimney sweeper." All of these names are indica-
tive of the sooty-black character of the spores. There are two chief
phases in the development of a smut fungus, the mycehal phase and
the spore phase. The hyphae of the mycelium mostly push between
the cells through the intercellular spaces and form short special branches,
or haustoria, which enter the host cells and absorb from them nutritive
material. The mycelium may be locahzed, or it may be spread gen-
erally throughout the host. Where the mycelium gains entrance to
the host through the germinating seeds, it remains in the vegetative
BASIDIA-BEARING FUNGI (SMUTS) 1 79
condition and without external manifestation of infection until in its
fruiting stage, when it breaks through the tissues of the host, appear-
ing at the surface. In perennial plants, the mycelium may live in the
perennial parts, each year extending into the new growth. Eventually,
the mycelium becomes conspicuous in certain organs of the plant. It
may develop abnormal growths, or cause swellings in the stem leaves,

Here the hyphae break
flowers (anthers, ovaries), or fruits of the host.
up into chains of spores, which develop thicker walls than the hyphal
cells from which they arose and are known as chlamydospores (xXayuus,
xXctfxvdos = a cloak + (rwopa = a seed). The hyphal cells between the

spores undergo almost complete gelatinization, which gelatinized cells
are used probably to nourish the developing spores, as at maturity the
spores lie loosely surrounded in part by the diseased cells of the host
ready to be discharged as the adjoining hyphal and host cells dry up
and completely disappear. The chlamydospores, which make up the
smutty, or sooty masses, are usually thick-walled and, being small,
4 to 35^t, they are easily disseminated. They are usually spherical, or
spheroidal, but may be ovoid, eUipsoidal or even oblong. They are
simple, i.e., consisting of single cells, but they may be united into spore
balls,which may have an external coating of sterile cells. The galls
of the chlamydospores may be smooth, or echinulate, or reticulate with
a network of ridges, or wings. Their color may be yellowish, reddish
or olive-brown, violet, or purplish, and the dark-colored spores in mass
may appear to be black or dark amber-brown. Sori are masses of the
spores that break out singly, or in clusters, on the various organs of
the hosts. These clusters are protected by their coverings of the tissue
of the host. The sori may be dusty and easily broken up, while in
other species, they may be hard and the spore mass is gradually
disintegrated.
The wind is undoubtedly one of the principal agents in the dissemi-
nation of the smut spores, but it was found that no smut spores could
be demonstrated in spore traps set up at the University of Manitoba
by BuUer farther distant from the infected fields than 250 yards. Man
distributes the spores through unclean agricultural methods, such as
using old grain bags over and over again, and in sowing seed to which
the smut spores are attached. The threshing machine is an active
agent in the spread of smut spores, and the farmer should see that his
machine is carefully cleaned from one operation to another.
i8o MYCOLOGY
—
Fig. 62. Germination of smut spores, a, Chlamydospores; b, basidium; 5,
basidiospores; d, infection threads; e^ detached pieces of mycelia; /, knee-joints, i.
Germination of Ustilago avenae in 1/ 50 per cent, acetic acid 24 to 48 hours after being
placed in liquid. 2. Same as in i but in distilled water. 3. Germination of Ustil-
ago levis in Cohn's modified solution at end of 24 hours. 4. Same as 3 but at end of
2 or 3 days. 5. Germination of Ustilago Iritici in Cohn's modified solution. 6. Ger-
mination of Ustilago striafortnis from red top in 1/ .50 per cent, acetic acid at end of
2 days. 7. S'ame as 6 except in Cohn's modified solution. {After Bull. 57, Univ.
III. Agric. Exper. Stat., March, igoo.)
BASIDIA-BEARING FUNGI (SMUTS) l8l
Experiments to determine the vitality of smut spores have shown

that those of the stinking smut of wheat, covered smut of barley and
oat smut are long-Hved under favorable conditions for seven, or eight
years, and in a dry condition are resistant to frost. Where vegetative
reproduction occurs, as in the loose smuts, the spores lose their vitality
after five to six months. It has also been determined that stinking
smut spores passing through the bodies of animals lose their power of
germination in a great majority of cases. Only those passing through
pigs retain their vitaHty a longer time. The presence of occasional
viable spores in the manurial offal of animals suggests a danger of
the spreading of smut diseases through manure applied to fields as
fertihzers.
Germination (Fig. 62). —The spores, when placed in a drop of
water, send out a single hyaline thread several times the length of
the spore, and this thread, or promycelium, becomes divided into four
cells by cross-partitions, or septas. Usually the apex of these four cells
produce one or more elongated thin-walled spores, the basidiospores,
or sporidea. These basidiospores are pinched ofif at the base, and
others are formed to take their place. When the basidiospores reach
the proper host, whether in the seed, seedling, partly grown or mature
condition, it forms on germination an infection hypha, which bores
through the surface and enters the interior of the host. Once inside a
mycelium is formed.
Modes of Infection. —
(i) Certain smut spores, as those of the
stinking smut of wheat, covered smut of barley, naked and loose smuts
of oats and others, adhere to the outside of the grains and are sown
along with the grain. In the soil germination takes place and the spore
produces a short stout mycelium, which develops secondary, or even
tertiary spores, which by means of infection threads attack the young
grain seedlings as they grow upward through the soil. This mode of
infection is called seedling infection. (2) In the so-called loose smuts
of wheat and barley, the chlamydospores, which are mature at the time
of flowering of these commercial grasses, fall upon the female organs
of the wheat, or barley, and germinating the infection hypha pushes
its way into the developing grain where it remains dormant as a deli-
cate mycelium. The normal development of the grain is not inhibited,

so that when it is planted as seed, the mycelium begins to grow with
the seedling and keeps pace with the future growth of its host until
I 82 MYCOLOGY
the maturity of the spores at the time the wheat, or barley, come into
bloom. This mode of infection is known as flower infection. A third
method is shown by the corn smut which may infect its host at any
time by entering the young and tender parts of the plant. A knowledge
of these facts is important, for the treatment of seeds will be efficacious
with smuts, which infect seeds, while it would be useless with infection
accomphshed by the second and third methods.
Grain smuts cause a considerable loss to the farmer every year.
Oat smut, it has been estimated, causes a loss of $10,000,000 per annum
in the United States. Smut explosions have been recorded recently. ^
In the wheat-growing regions of the Pacific Northwest in the summer of

1 914, 300 threshing machines were blown up or burned by smut ex-
plosions. Passing into the cylinder of the threshing machine, the smut
balls were broken up and the highly combustible smut dust oily and
dry filled the interior of the separator. It is when this condition ob-
tains, that the explosions and flames occur. The smut dust was prob-
ably ignited by static electricity in the cyHnder of the threshing machine.
The drier the conditions, the more static electricity is formed, and the
easier it is to ignite the smut.
The family Usttlaginace^ includes eleven American genera. Only
three genera out of the seven will be considered in this book. They are
Ustilago, Sorosporium and Tolyposporiiim. The genus UsHlago, of
which there are about seventy-two American species, is distinguished
from the other two less important genera by its single spores which
form dusty masses at maturity without any kind of inclosing membrane.
Sorosporium has its spores agglutinated into balls which form more or
less dusty masses. The spore balls are usually evanescent and the
spores are very dark. The spores are agglutinated into balls in Toly-
posporium, forming more or less dusty spore masses. The spore balls
are rather permanent, the spores adhering by folds, or thickenings of
the outer coat.

Family 2. Tilletiace^. —The name Tilletia which is that of an
important genus (Fig. 63) of the family is derived from Matthieu Tillet,
who published a book in Bordeaux, France, in 1755. The sori form
dusty spore masses, which break out to the surface, or are imbedded
permanently in the plant tissues, often without causing any malforma-
1 AsHLOCK, J. L.: Smut Explosions. The Country Gentleman, April 10, 1915,
P- 703-
BASIDIA-BEARING I'UNGI (SMUTS) 183
—
Fig. 63. Bunt or stinking smut of wheat (Tilletia Irilici). a. Whole head af-
fected with smut; h, smutted grains; c, normal grains; d, smutted grain broken to
show spores; e, normal grain divided in the middle; /, chlamydospores enlarged; g,
germination of a spore. {Draivings by Pool, Venus A., from Bull. 135, Set. Ser. 141,
Univ. of Tex., Nov. 15, 1909.)
184 MYCOLOGY
tion of these parts. In germination, a promycelium is formed, which

usually gives rise to a terminal cluster of elongated basidiospores, or
sporidia,which sometimes bear whorls of secondary basidiospores.
Sometimes the primary sporidia fuse in pairs, and these with or without
fusing may give rise to infection hypha?; or in nutrient media to a
mycehum bearing dissimilar secondary sporidia (aerial conidia). As
in the preceding family the hyphae break up into chlamydospores which
break through the host tissue, as a sooty mass of dust. When these
chlamydospores germinate, they give^ rise to an undivided basidium
with basidiospores borne at the apex not on the side, as in the preced-
ing family. This is the principal morphologic difference, as the two
groups of smut fungi approach each other so closely tîat in external
appearance they resemble each other. Brefeld described the structure
and life history of Tilletia tritici {T. caries), the bunt of wheat very
carefully. In England, this disease of the wheat plant is called in
various districts pepper brand, smut balls, bladder brand, stinking
smut, stinking rust (Fig. 63) In the fields, it is difficult to distinguish
diseased from sound wheat, as there is little to indicate the presence of
the hidden parasite, but it excites an abnormal development of chloro-
phyll, so that the spikes of the affected plants are usually greener than
the healthy ones. The brand spores are found in all the grains of a single
ear. The burst grains are shorter and wider than healthy ones and
pointed toward the base. When cracked, a black dust is discharged,
which under the microscope is seen to consist of reticulate-walled spores
of an olive-brown. They germinate readily and even after eight and a
half years, they have been known to grow. On rubbing the black
powdery mass between the fingers, the smell of herring brine is given
off,and this decayed fish odor has originated one of the common
names, that of stinking smut. A curved unicellular basidium arises
from the chlamydospore on its germination. This produces a bundle
of elongated condiospores, or basidiospores, according to one's bias.
Sickle-shaped secondary conidiospores arise from the primary kind.
The primary conidiospores may unite by bridge-like connections so
that two united spores look like the letter H. Wheat becomes infected
in the seedling state, the spores being sown with the grain, and the
infection hypha which enters the host forms a mycelium which grows
along with the host until the spores break out again.
Tilletia is the most important genus. In it the sori may occur in
BASIDIA-BEARING FUNGI (sMUTS) 1 85
various parts of the host, usually in the ovaries, where are formed a
dusty dark spore mass. The spores are simple, separate and originate
singly at the ends of special hyphse, which almost disappear through
gelatinization. The spores varies in size from i6/x to 35/1. Fifteen out
of the fifty-three species recorded by Saccardo have been found in
North America. The important species are Tilletia fcetens bunt or
stinking smut of wheat; Tilletia tritici on wheat; Tilletia horrida
in the ovaries of cultivated rice; Tilletia anthoxanthi in the ova-
ries of the sweet vernal grass, Anthoxanthum odoratum; and Tilletia
Maclagani on a wild grass, Panicum vigatum. Urocystis cepulcB is the
onion smut; Urocystis occulta on the stems and sheaths of rye; Urocystis
violcB on the stems, rootstocks, petioles and leaves of violets, Entyloma
crastophilum levis on such grasses as Agrostis, Poa, E. Ellisii forms pale
white spots on spinach leaves in New Jersey. Entyloma lineatum grows

on wild rice, Zizania aquatica; Entyloma thalictri on the meadow rice,
Thalictrum polygamum; Entyloma lobelice or Lobelia inflata; Entyloma
nymphcece on the leaves of Nuphar advena and Nymphcea odorata.
The species of Doassansia mostly grow on plants, such as: S a git-
tar ia, Potamogeton, etc., growing in moist situations. Ten species
occur in North America.
BIBLIOGRAPHY OF THE SMUTS
Arthur, J. B.: Rapid Method for Removing Smut from Seed Oats. Bull. 103,
vol. xii, Agric. Exper. Stat. Purdue University, March, 1905.
Clinton, G. P.: The Smuts of Illinois Agricultural Plants. Bull. 57, Agric. Exper.
Stat. Urbana, March, 1900.
Clinton, G. P.: North American Ustilagineas. Journal of Mycology, 8: 128-156,
October, 1902
Clinton, George P.: North American Ustilagines, Proceedings Boston Society
of Natural History, 31: 504, 1904.
Clinton, George P.: The Ustilaginea?, or Smuts, of Connecticut. Bull. .5,
State Geological and Natural History Survey, 1905.
Clinton, George P.: Ustilaginales (Ustilaginaceae, Tilletiaceae). North American
Flora, 7, part I: 1-82, Oct. 4, 1906.
DiETEL, P.: Hemibasidii. Die naturhchen Pflanzenfamilien, I. Teil, Abt. i,
1900: 2-24.
DuGGAR, B. M.: Fungous Diseases of Plants, 1909: 370-383.
Eriksson, Jakob: Fungoid Diseases of Agricultural Plants, 191 2: 44-62.
Garrett, A. O.: The Smuts and Rusts of Utah. Mycologia, II: 265-304, No-
vember, 1910.
1 86 MYCOLOGY
Gusspw, H. T. Smut Diseases of Cultivated Plants. Their Cause and Control,

Bull. 73, Division of Botany, Central Experimental Farm, Ottawa, Canada,
March, 1913.
Henderson, L. F. : Smuts and Rusts of Grains in Idaho. Bull. 11, Agric. Exper.
Stat., Idaho, 1898.
Hitchcock, A. S. and Norton, J. B. S.: Corn Smut. Bull. 62, Exper. Stat.,
Kansas State Agricultural College, December, 1896.
Massee, George: Text-book of Fungi; 1906: 313-325.
Massee, George and Ivy: Mildews, Rusts and Smuts: a Synopsis of the Families,
Peronosporaceae Erysiphaceae, Uredinacefe and Ustilaginacea?, 1913: 182-205.
Smith, Worthington G.: Diseases of Field and Garden Crops, 1884: 245-262.
Stevens, F. L.: The Fungi Which Cause Plant Disease, 1913: 298-323.
Swingle, Walter T.: The Grain Smuts: How They Are Caused and How to
Prevent Them. U. S. Farmers' Bull. 75, 1898.
.
Underwood, L. M.: Moulds, Mildews and Mushrooms, 1899: 81-85.

von Tubeuf, K.: Pflanzenkrankheiten, 1895: 289-340.
VON Wettstein, Richard R. Handbuch der Systematischen Botanik,
: 1911: 193-
195-
CHAPTER XIX
RUST FUNGI
Suborder Uredine^. —-Usually in systematic works placed as
ORDER UREDINALES. The fungi belonging to this suborder are
characterized by basidia which are divided either by transverse or
longitudinal septae. In this character, they are contrasted with the
EUBASIDII, which have unseptate basidia. Including the rusts this
suborder embraces some of the most important disease-producing
fungi, the study of which concerns the mycologist.
The uredineous fungi are those which are strictly parasitic and
which in some cases are so specialized, that their growth is confined to
the species of a single host. Those fungi in which the different stages
of the life cycle are passed on the same host are known as autoecious,
while those which grow on two or more hosts are known as heteroecious.
The plant on which the final stage is passed is called the final host,
while the other plant on which some of the stages occur is designated
the alternate host. So speciahzed is the nutrition of the rust fungi,
that they never have been grown on culture media off the host
plants on which they live. Hence, they are obligate parasites. The
myceUum is septate, much-branched, usually ramifying between or in
the walls and sending haustoria into the cell cavities.
of the cells
The reproductive spores are borne in more or less definite clusters, or

sori, below the surface of the host, or rarely singly, and the spores are
set free by the breaking open of the overlying tissues of the hosts.
Five different kinds of spores may be found in the uredineous fungi,
but they are not all present in every genus (Fig. 64). The final spore
form is known as the teliospore, or teleutospore, which determines the
name which is to be appHed to the parasite. Such spores are borne
known as a teUum.
in a sorus When these teliospores germinate, they
produce a four-celled promycelium known as a basidium, and this
abstricts sporidia, ormore properly basidiospores, which are minute,
thin-walled spores without surface sculpturings. These are succeeded
by spermogonia (spermogonium), which are now called by most
187
I 88 MYCOLOGY
American mycologists, pycnia (pyciiium), in which spermatia, or

pycniospores, are formed. Pycnia indicate the nature of the life cycle
and furnish positive characters for identification. Arthur has shown
that if pycnia and urediniospores are found arising from the same
mycelium, aecidia do not occur in the cycle; and if pycnia and telio-
spores are found there are neither uredinia nor secia in the life cycles.
These pycnospores are accompanied or succeeded by aeciospores
(aecidiospores), which appear in the cluster cups, or aecia in long chains.
The peridia of the different kinds of aecia are variable, and hence
Fig. 64.— Spore forms of wheat rust, Pucainia graminis. A, Section through
barberry leaf showing pycnia on upper surface and secia on lower; B, two uredinio-
spores; C, germinating urediniospore D, teliosorus showing several teliospores; E,
;
single two-celledjteliospore F, germinating teliospore with four-celled basidium and

;
two basidiospores; G, basidiospore growing on barberry leaf. {Adapted from deBary.)
mycologists have described four different kinds of form genera: Cceoma

= peridium absent; Mcidiiim = cup-shaped and peridium toothed;
Roestelia = peridium elongate fimbriate; Peridermium = peri-
and
dium irregularly and broken. Urediniospores (uredospores)
split
succeed the aeciospores and they appear in sori known as uredinia

Curedinium). Amphispores are special forms of urediniospores formed
in arid, or semi-arid climates and usually have a thick cell wall and a
persistent pedicel. They are in the nature of a resting spore. Meso-
spores are exactly of the same nature as the two-celled teliospores, but
they arise merely by the omission of the last nuclear division, and hence.
RUST FUNGI 189
have only one cell. These different kinds of spores, representing stages
in the life histories of the different genera and species of rusts are
designated, as follows: O = pycnium; I = aecium; II = uredinium;
III = telium. The determination of the presence or absence of these
life histories has been made for a large number of
spores in the various
rusts,and we are now in a position to tabulate the results of this study
and to give names to the different forms of rust life cycles which have
been found. We call a fungus possessing:
Auteu-form, if all four kinds are found on one plant
(Ex. Puccina Asparagi on Asparagus officinalis).
O I II III an Eu-form Hetereu-form,if O, I occur on one species and II, III
}
on another (Ex. Puccinia gramiitis is on wheat and
I barberry).
O I III an opsis-form (Ex. Gymnos porangimn Jutiiperi-virginiance, O, I on
apple, and III on red cedar).
O II III a Brachy-form (Ex. Puccinia suaveolens on Canada thistle).
[O] III a Micro-form pycnia (spermogones) sometimes absent (Ex. Puc-
cinia ribis on currant).
A Lepto-form is one, of whatever kind, in which the teliospores

grow as soon as mature without any period of rest, as Puccinia malva-
cearuni on hollyhock. W. B. Grove in his "British Rust Fungi,"
page 40, gives a diagram which represents all of the possible life cycles
of the different forms of rust fungi. It is reproduced here (Fig. 65).
As a fungus which shows a complete life history passed on two dis-
tinct host plants, we will take the black rust of cereals, Puccinia
graminis (Fig. 64), first carefully studied by the German botanist,
Anton de Bary, in 1864-65. It infests all the common cereals, wheat,
rye, barley and oats, also many grasses. It appears on the wheat
plant, when the host is about ready to produce its spikes of flowers. It
appears on the leaves and culms of the wheat plant, as orange-red

lines, which represent cracks in the epidermis of the host exposing the
sori, or uredinia filled with rust-red spores, urediniospores. These
summer spores are yellowish and their surface spinulose with four equa-
These urediniospores may follow each other on
torial germ pores.
summer. This summer stage is succeeded

several crops during the early
by the autumn stage in which the sori become filled with stalked,
'
two-celled, dark-colored spores with thick walls. The common name
of this stage is "black rust." Wintering in the open these two-celled
teliospores germinate. Each of the two cells may sprout out a pro-
mycelium, or only one may do so. This basidium (promycelium) is
I go MYCOLOGY
upright and divided transversely into four cells, each of which cuts off
a basidiospore. These basidiospores are blown to the leaves, twigs, or

fruits of the barberry where a mycelium is formed. Later pycnia
(spermogonia) appear on the upper side of its leaf. These are accom-
panied by round, fringed depressions, the cluster cups or ascia, which
appear in the spring on the lower side of the leaves. The agciospores
are arranged in chains. These spring spores, aeciospores, are carried to
the wheat plant where they induce the characteristic rusted appearance
basidium
teleutospore
basidiospore
uredospore
mycelium
secidiospore
fusion-cell
Fig. 65. — Relations of various spore forms of rusts to each other. {After Grove, W.
B., The British Rust Fungi, 19 13, 40.)
of the cereal. The wheat plant is not killed by the attack of the fungus
which, however, prevents the reserve foods from being properly stored
in the grains; hence, they are mushy and unfit for storage, or for bread-
making purposes. been recently shown that in Australia and the
It has
plains of India, where the barberry is unknown, the black rust of wheat
does serious damage. Three methods are open to the wheat rust to
winter over: (i) The fungus may winter by its urediniospores, (2) by a
perennial mycelium, (3) by Eriksson's mycoplasm. Arthur, in Amer-
ica, and others have shown that it winters by its urediniospores, or
RUST FUNGI 191
amphispores, as they have been termed by some, but in conversation

with Arthur he insisted that the perennating spores are typical uredinio-
spores, so that the postulation of a perennial mycelium, or a hibernating
fungous protoplasm in the cells of the grain (mycoplasm) is unneces-
sary. Eriksson has proved that in Sweden six forms of Puccinia
graminis may be distinguished; which he enumerates as follows:
A. Not distinctly fixed (occasionally going over to other forms of
grass): (i) f. sp. tritici on wheat (seldom on rye, barley and oats).
B. Distinctly fixed (firmly confined to the indicated species): (2)
f. on rye, barley and on couchgrass, Agropyron repens, Ely-
sp. secalis
mus Bromus secaUnus and others; (3) f. sp. avenae on oats

arenarius,
arid on Avena elatior, Dactylis glomerata^ Alopecurus pratensis, Milium
efusuni and others; (4) f. sp. poae on Poa compressa and P. pratensis;
(5) f. sp. airae on Aira ccespitosa and ^4. hottnica; (6) f. sp. agrostis on
Agrostis canena and A. stolomfera. An oat plant infected with this rust
can in its turn infect wheat, rye, barley and so forth. The black rust
of cereals is the classic example of an heteroecious rust.
The asparagus rust, Puccinia asparagi, may be used to illustrate the
life history of an autoecious species. All the spore forms are pro-
duced on stems and twigs. The aecia appear in long, light green cush-
ion-like areas, which are short cylindric with a white peridium. The
aeciospores are orange-colored and the wall is hyaline. The pycnia
appear in yellow clusters followed by the aeciospores in early sum-
mer. The uredinium is filled with yellowish-brown, thick-walled uredi-
niospores w'ith three or four germ pores. The black rust stage (telium)
appears later in the season, when the two-celled stalked teliospores push
out from beneath. The whole life cycle is passed on the asparagus
plant.
—
Cytology of the Rusts. According to the earlier researches of
V. H. Blackman (1904), A. H. Christman (1905), O. H. Blackman and
Miss H. C. Fraser (1906), Edgar W. Olive (i9o8),Kurssanow (1910) and
Dittschlag (191 6), supplemented by the research of other botanists, a
flood of light has been thrown on the nuclear behavior in the rusts, and
accordingly on their sexuality, or non-sexuality. Blackman discovered
in Phragniidium violaceum (Fig. 66), that in the formation of the
aecidium, there was a fusion of two cells by which the nucleus of one
passed over into the adjoining cell. In the formation of spores the
paired nuclei of the fusion cell divide side by side and simultaneously
(conjugate division) so that we find that the basal cell, the aecio-
—
192 MYCOLOGY
pores and intercalary cells all have two nuclei, which are not sister
nuclei. The upper cell, cut off from the fusion cell, is the secio-
spore mother cell; the lower grows a little longerand then divides again
in the same way, and thus a vertical series of aeciospore mother cells is
formed, the oldest at the top. Each of the aeciospore mother cells.
Fig. 66. A, Chain of young jeciospores of Puccinia caricis; a, fusion tissue;

b, basal (fusion) cell with conjugate nuclei; c, asciospore mother-cell; d, intercalary
cell; e, young aeciospore; B, germinating teciospore of P. caricis; C, teliospore of P.
caricis; D, formation of teliospores of P. falcaria {after Dittschlag); E, development
of aecium {after Blackman) of Phragmidium violaceum; e, epidermal cell; s, sterile
cell; below these cells a nucleus is seen migrating into the adjacent cell /; F and G,
conjugation of two female cells to form basal cell of aeciospore chain {after Ditlschlog).
In G the first conjugate division is just completed. {Adapted frotn Grove, British
Rust Fungi.)
as soon as it is formed, cuts off by conjugate division a small cell below,

called the intercalary cell; this sooA disorganizes and disappears, while
the other portion remains as the aeciospore. The succeeding uredinio-
spores have two nuclei in the conjugate condition and this is continued
over into the cells of the young teliospores (Figs. 67 and 68). Before
RUST FUNGI 193
the teliospore reaches maturity, the nuclei fuse, and the uninucleate
condition then continues again until the formation of the
gecia. In the
micro- and which have no aecium or uredinium, we find that
lepto-iorms,
the association takes place at points in the ordinary mycelium, but
Fig. 67. —Portion of a section of cedar apple about 5 mm. below a teliosorus.
Note (i) Binucleate intercellular mycelium; (2) the haustoria in various stages of
development; (3) the doubling of nucleoli in the nuclei of some of the parenchyma
cells of the host. Material collected on March 31. {After Reed, H. S., and Crabill,
C. H., Techn. Bull. 9, Va. Agric. Exper. Stat., May, 1915.)
always before the formation of the teliospores. Whether the

association of nuclei in the ordinary mycelium takes place by the
migration of a nucleus from one cell to another, or whether two daughter
nuclei become conjugate one cell has not been settled definitely.
in
The pycnospores are probably abortive male cells. They have never
13
194 MYCOLOGY
Fig. 68. —
Portion of a teliosorus of cedar apple in February showing mycelia
stroma and the binucleate condition of the cells of young teliospores. (After Reed,
H. S., and Crahill, C. H., Techn. Bull, g, Va. Agric. Exper. Stat., May, IQ15.)
basidiospores
teleutospore
uredospoTe.
SPOEOPHYTE GAMETOPHYTE
(2?i generation) (n generation)
spermatium
uredospore $ gamete
aecidiospore gametes
intercalary cell
fusion-cell
Fig. 69. — Diagram of the alternation of generations of a typical rust. {After Grove,
W. B., The British Rust Fungi, 1913, 27.)
RUST FUNGI 195
been known to germinate, and the large size of their nuclei suggests that
we arc dealing with male cells.
The mature tcliospore, which may be looked upon as a spore
mother cell, has a single fusion nucleus. "The fusion nucleus is large,
round and (when unstained) perfectly clear and homogeneous, but for
its nucleolus, so that it looks like a vacuole; it occupies almost invari-
ably the middle of a cell. The dense chromatin mass is loosened out
into a kind of spireme which becomes shorter and thicker; the nuclear
membrane then disappears, and the spireme thread splits longitudi-
nally, though the splitting is often indistinct. It then divides trans-
versely into segments which become arranged, or strung out, on a
spindle (sometimes, but more rarely, in an equatorial plate) then the ;
daughter nuclei are formed at the poles, and the next division, which
is homotypic, follows immediately" (Harper and Holden, 1903;
Blackman, 1904). These nuclei are found in each of the four cells
which form the basidium, and ultimately, they pass into each of the
four basidiospores which are uninucleate and haploid.
The alternation of generations which has thus been determined by
the various cytologic studies of recent years may be displayed in a
diagram adapted from Grove (Fig. 69).
The same life cycle may be represented in another way.
Basidiospore
Mycelium
Gametophyte
(w generation)
Pycnium
Female cells Pycnospores
Fusion cell
II
.^ciospore mother cell Jicium

. ^
iEciospore
\
Intercalary eel
Sporophyte
(2« generation) Urediniospore (repeated)
Teliospore
0000
4 Basidiospores
196 MYCOLOGY
EndophyUum sempervivi which attacks the house leek, Semper-

vivum, and causes its rosette of normally spreading leaves to stand
erect, shows a somewhat different condition, which has led to the sup-
position that it represents the primitive life cycle of the higher ure-
dineous fungi. Its life history has been investigatecl by Hoffman
(191 1). The spores mature on the house-leek leaves in April and
May. They germinate at once in the secidioid telium and a four-
celled basidium is formed; hence, the spore looks like an seciospore
and partakes of the nature of a teliospore and may be called an secio-
teliospore. Each basidium produces four basidiospores on long sterig-
mata, and they are blown to the leaf of a house leek, where they
begin growth at once by boring through the cuticle, and the mycelium
then grows through the intercellular spaces of the host sending haus-
growing down to the base of the leaf and into the
toria into the cells,
axis up to the growing point, where it perennates until the following
spring, when it enters the freshly formed leaves, which become yellow,
longer and more erect.
Pycnia are formed in March and April followed by aecio-telia,
which repeat the cycle. Hoffman has established the most interest-
ing point about this rust, that the aecio-teliospore chain arises from a
cell produced by the fusion of two adjacent cells of the spore bed
after the manner described by Christman except the conjugating cells

were not in any definite plane. The binucleate secio-teliospores then
become uninucleate by the fusion of the conjugate nuclei. The for-
mation of the basidiospores from these oecio-teliospores probably
follows a reduction division.
Kunkel (191 4) has shown that a study of the binucleate seciospores
of CcBoma nitens during germination shows that they become uninu-
cleate previous to the production of the promycelia. The normal ger-
mination of the aecio-teliospore consists in the pushing out of a germ tube
into which the protoplasmic contents of the spore passes. The nucleus
which travels out into the tube divides producing two nuclei which may
divide again immediately and cell division may follow at once, but in
other cases the four nuclei of the promycelium (basidium) may be
present before cross walls are formed. Ultimately, four cells are found
filled with protoplasm and uninucleate. The basidiospore arises as an
enlargement of the sterigma and the nucleus enters when it is one-half
developed. Cceoma nitens although like EndophyUum sempervivi in
some respects is more primitive, since it possesses a simpler aecium.
RUST FUNGI 197
Phylogeny of the Uredine^ (Uredinales)
In looking for the primitive types of rust fungi, it has been assumed
by some mycologists, that, as the rusts are a specialized group of para-
sites, the most primitive forms will be found on hosts which are lowest
in the phylogenetic scale of the higher plants. This consideration

would place Uredinopsis, which grows upon ferns, as one of the primi-
tive rusts, while many of the more advanced types of Puccinia are found
upon the Composite. The absence of a germ pore is considered primi-
tive, as instance its absence in the aecio-teliospore of EndophyUum.
When these first appeared, they were numerous and indefinitely scat-
tered, while in the higher rusts, they are reduced in number and
restricted to a definite part of the cell wall. The formation and ger-
mination of teliospores approaches that of the smuts a more primitive
group, hence the formation of a basidium and basidiospores must
have been inherited by both from their ancestors. Now among the
red algge, such as Grifjithsia, the sporophyte bears tetraspores, these
develop into a thallus which bears the gametes. Hence one would look
for the ancestors of the UREDINE^ among red algge. Again, it has
been suggested that the female cells of the ascium have a trichogyne,
such as the red seaweeds (Florideae) possess. In the rusts, it has become
abortive.
The Endophyllace^ are considered by Grove to constitute the
starting point from which the varied forms of the Pucciniace^ have
been derived. In EndophyUum, we have seen that the seciospore, which
is the product of the fusion cell, is also the teliospore from which the
basidium and basidiospores arise. The aecium is accompanied by the

pycnium here. The first stage of evolution was the separation of this
spore form into two: one the geciospores, germinating like conidio-
spores; the other, the teliospore, germinating with the formation of a
basidium and basidiospores. Pucciniopsis suggests these stages. The
summer spores are probably modified geciospores formed as a device
for repeating the spore generations without the intervention of another
fusion cell. The fusion of the two nuclei in the teliospore is from a
cytologic standpoint paralleled by a similar fusion in the BASIDIO-
MYCETALES, for a division into four basidiospores follows in both
cases, although the mechanism is different. The paired condition
of the nuclei found in the ascogenous hyphae of the ASCOMYCETALES,
such SLsPyronema confluens investigated by Claussen (1912), and in the
•
198 MYCOLOGY
formation of the ascus, the two nOn-sister nuclei fuse after which the
fusion nucleus divides, the first division being heterotypic (meiotic,
reducing, possessing synapsis and diakinesis stages), and the two fol-
lowing ones, which result in the formation of eight ascospores, are

homotypic. From this point of^view, the ascus is a spore mother cell
comparable to the teliospore of the rust fungi, but forming an octad,
not a tetrad of spores. The probable phylogeny and relationship of
the Uredine^ to the other groups has been set forth in a family tree
by Grove.
Arthur, who has studied the rusts carefully for many years, pro-
posed at the International Congress of Botanists held in Vienna in
1905 an arrangement of the famihes, genera and species of the rusts,
which differs materially from the older classifications.
As this classification of Arthurs has not been elaborated in detail,
it has been considered best to follow the arrangement of families, sub-
families and genera given in Engl er and Gilg's "Syllabus der Pflanzen-
familien" (7th Edition, 191 2) as following the conservative and
older treatment.
—
Family Endophyllace^. The teliospores are abstricted suc-
cessively in long rows and are surrounded by a peridium which is
formed like that of a typic aecidium of Puccinia from the peripheral
cell rows, but is sometimes less strongly developed. These teliospores
are perhaps more correctly called aecio-teliospores, as they are separated
from each other by intercalary cells like true seciospores and arise
from a fusion cell, but they germinate by the formation of a basidium
and basidiospores like true teliospores. The germ pores are impercep-
tible and the spore wall is colored. Pycnia are present and both kinds
of sori are subepidermal.
Endophyllum sempervivi lives parasitically on the house leek, Sem-
pervivum tedoruni, and several other species of Sempervivum in Europe
from April to August. It has been proved by de Bary, Hoffmann and
others, that the basidiospores produced by the secio-teliospores infect
the leaves of the house leek and from them arises a mycehum which
livesover the winter in the stem. The following spring, it forms
pycnia and secio-teliospores and the affected leaves are more erect
than normal ones, twice as long, narrower and yellowish at the base.
Family MelampsoracetE.—The tehospores are unstalked, one- to
but placed singly on dilated hyphie in the tissues of the
four-celled,
RUST FUNGI 199
host, or arranged side by side in flat crusts. Germination of the teHo-

spore results in the formation of a four-celled basidium, each cell of
which forms a single basidiospore. The secium is typically without a
peridium, hence, a cseoma and the urediniospores appear in long chains
without a peridium, or arising singly, and then mostly surrounded
by the peridium, or mixed with paraphyses.
The genus Melampsoropsis includes fungi whose teliospores are in
cushion-like layers, which break through the epidermis of the host.
M. ledi has its teliospores on Ledum and its aecia on the spruce, Picea
excelsa, in Europe, and on P. rubra in this country. The secia of Cronar-
tium have a broad, inflated irregularly torn peridium. The uredinium
is enclosed in a hemispheric peridium, which opens at the summit by a
narrow pore. Its teliospores are abstricted in long chains and remain
united into cylindric columns, which are horny when dry. The
European C. asclepiadeum has its gecia on the branches of Plnus sihestris
in May and June, and its urediniospores and teliospores on PcBonia
officinalis in gardens, as also on Vincetoxicum, Cynanchum and Verbena.
C. quercmim has its aecia on Pin us and its urediniospores and teliospores
on at least twenty species of oak in North America. C. ribicola is a
dangerous parasite called the white pine blister rust and against it the
United States Government has an active quarantine. Its aecium is con-
fined to the five-leaved pines, one of which is Pinus strobus, our eastern
white pine. These are found in the months from March to June. The
urediniospores and teliospores grow on the currants, Ribes nigrum and
R. rubrum. The fungi of the genus Melampsora are mostly heteroecious.
There are seven species recorded for North America. Of these Melam-
psora meduscB causes the poplar rust. The aecium occurs on the larch,
Larix, and its urediniospores and teliospores on Populus deltoides, P.
tremuloides and P. balsamifera. Calypfospora is a genus of rusts, the
life history of which has been investigated by Hartig, Kuhn and
Bubak. In July to September, the teliospores appear on the stems of

Vaccinium vitis-idcBa, where the stem becomes swollen and elongated
and at first of a pink color passing to brown. It occurs on other species
of Vaccinium, including V. pennsylvanicum in the United States. The
aecia are found in Europe on leaves of Abies pectinata and in America
on A balsamea.
.
—
Family Coleosporiace^. The aecium in this family has a perid-
ium. The flattish, linear pycnia are subepidermal dehiscing by a
—
MYCOLOGY
slit.The teliospores consist of four superimposed cells. There is a

North American species of this family, Gallowaya pini (formerly Coleo-
sporium pini), which has teUospores only and these on the leaves of
Pinus inops, i.e., on trees of the same order on which Coles porimn has
A
Fig. 70. A-D, Uromyces pisi. A, Ascidia on deformed leaves of Euphorbia
cyparissias; B, ascidia enlarged; C, teliosori on leaves of Pisum sativum; teliosori
enlarged; E and F, Uromyces Irifolii on Trifolium hybridum. {After Dietel, Die
natiirlichen Pflanzenfamilien I. lA**, p. 55.)
its aecia. In Coleosporium, the teUospores are adherent closely with a

rounded, thickened, gelatinizingpore. The long sterigmata bear
large, ovate, flattened sporidia. The orange rust of asters and golden
rods,^'C. solidaginis is reported to cause a sickness of horses, some-
RUST FUNGI 20I
times resulting in the death of the animals. Its urediniospores and

teliospores are on compositous plants and its aecial stage on the pitch
pine, Pinus rigida, this stage being known in the older books as
Peridermium acicolum. The species of the genus are all heteroecious,
and ascial stages, whenever found, occur on species of Pinus and
are referable to the form genus Peridermium. Arthur and Kern
enumerate twenty-seven species of Peridermium, ranging from Mexico
to Alaska, and from the Atlantic to the Pacific coasts. The species
are all secia of species belonging to tehal genera, but they cannot
be always satisfactorily assigned because of incomplete knowledge
regarding them. The genus Peridermium embraces all aecial forms
possessing peridia, inhabiting the Pinace^ and Gnetace^. Only
three of the twenty-seven American species have been associated with
telial forms as follows:
Peridermium pini connected with Coleosporium campanulcc on
Campanula.
Peridermium cerebrum connected with Cronartium on oak.
Peridermium elatinum connected with Melampsorella cerastii.
—
Family Pucciniace^. In this family, the teliospores usually con-
sist of a single cell, or a vertical row of superimposed cells sometimes
united into a small bead-like cluster. The teliospores are borne on a

simple, or a compound pedicel. The urediniospores are single, on
hyaline, deciduous stalks. The secia usually have a peridium. The
most important genera of the family are: Uromyces, Puccinia, Gymno-
sporangium, Gymnoconia (Fig. 71) and Phragmidium.
The rusts belonging to the genus Uromyces have one-celled winter,
or teliospores, which are egg-shaped, individually separated and massed
in small, open spore groups. The important pathologic species are the
clover rust, Uromyces trifolii; the rust of beans, U . appendiculata; beet
rust, U. betcB; carnation rust, U. caryophyllinus (Fig. 70). The largest
genus of the rusts, Puccinia, has usually two-celled teliospores, although
unicellular ones may occur in some species. The principal cereal or
grain rusts may be enumerated first, known,
as they are fairly well
owing to the researches of Eriksson and others:
Black Rust of Cereals, Puccinia graminis (Fig. 64) with its aecium
on the barberry, Berberis vulgaris. Six forms of this species may be
distinguished: (i) f. sp. h-itici on wheat (seldom on rye, barley
and oats); (2) f. sp. secalis on rye, barley and couch grass, Agropyron
—
202 MYCOLO&Y
repens, Elymns arenarius, Bromus secalimis and others; (3) f. sp.

avencB on oats and Avena elqtior, Dactylis glomerata, Alopecurus praten-
sis, Milium efusum, etc.; (4) f. sp. po(B on Foa compressa and P. praten-
sis; (5) f. sp. airce on Aira ccBspitosa and A. hottnica; (6) f. sp. agrostis on
Agrostis canina and A. stolonijera.

Brown Rust of Rye, Puccinia dispersa, with its cluster cups on
Anchusa arvensis and A. officinalis.
Crown Rust of Oats, Puccinia coronifera, with its secium on the
buckthorn, Rhamnus cathartica. Of this species there are eight
Fig. 7 1.
A-C, Gymnoconia interstitialis. A ^cidia on leaf of Rubus canadensis;
,
B, piece of leaf enlarged; C, teliospore; D, teliospore oi Sphenospora pallida, 500/ i.

{After Dieiel: Die naturlichen Pflanzenfamilien I. lA**, p. 70.)
speciahzed forms, as follows: (i) f. sp. avencB on oats; (2) f. sp. alope-
curi on Alopecurus pratensis; (3) f. sp. festuccd on Festucas; (4) f. sp.

lolii on rye grass, Lolium perenne; (5) f. sp. glycericB on Glyceria aqua-
tica; (6) f. sp. agropyri on Agropyron repens; (7) f. sp. epigm on Cala-
magrostis epigeios; (8) f. sp. hold on Holcus lanatus.

Crown Rust of Grasses, Puccinia coronata, with its aecium on Rham-
nus frangula. Three special forms of this rust are known: (i) f. sp.
calamagrostis on Calamagrostis ariindinacea; (2) f. sp. phalaridis on
1 Arthur, J. C. and Kern, F. D.: North American Species of Peridermium,

Bull. Torr. Bot. Club, 33: 403-43''î iyo6.
RUST FUNGI 203
Phalaris armidinacca; (3) f. sp. agrostis on Agrostis vulgaris and A,

sfolonifera.
Yellow Rust of Wheat, Puccinia glumarum, without any known
aecial stage. It has according to Eriksson the following specialized
forms: (i) f. sp. tritici on wheat; (2) f. sp. secalis on rye; (3) f. sp.
hordei on barley; (4) f. sp. Elymi on elymus arenarius; (5) f. sp.
agropyri on couch grass, Agropyron repens.
Fig. 72. — Hollyhock rust, Puccinia tnalvacearum. {Nantucket, August 19, igis.)'
Brown Rust of Wheat, Puccinia triticina, with aecia unknown.

Dwarf Rust of Barley, Puccinia simplex.
Timothy Rust, Puccinia phlei-pratensis. Experiments to get this
form to infect barberry leaves have met with indifferent success.
Chrysanthemum Rust, Puccinia chrysanthemi, on leaves of Chry-
santhemum sinensc in greenhouses all the year round.
—
204 MYCOLOGY
Dandelion Rust, Puccinia iaraxaci, on the dandelion Taraxacum

rather common in Europe, North America, Japan and the
officinale,
East Indies.
Reed Grass Rust, Puccinia phragmitis, with aecia on Rumex crispus,
R. ohtusifalius and urediniospores and teliospores on reed grass Phrag-
mites communis.
Fig. 73. Roeslelia aurantiaca on fruit oi Amelanchier intermedia corresponding

to Gymtiosporangium clavipes on red cedar. (Shelter Island, New York, July 16,
•1915-)
Ash Rust, Puccinia fraxinata, on leaves and petioles of ash and

uredinospores and teliospores on salt grass, Spartina Michauxiana.
Asparagus Rust, Puccinia asparagi, develops all of its spore forms
on the cultivated asparagus.
Violet Rust, Puccinia violce, is parasitic on about forty-six different
RUST FUNGI 205
species of violetsjn\\sia/ Europej'^Northând South America. It

autoecious.
2o6 MYCOLOGY
Rust of Stone Fruits, Puccinia pruni-spinQsa:, occurs on various

species of the genus Prunus in the southern and central United States.
Fig. 76.- — Fully expanded cedar apple on red cedar. Long yellow teliosori as
finger-like projections are seen. (After Jones and Barlholomew, Bull. 257, Agric.
Exper. Stat., Univ. Wise, July, 1915.)
The secial stage occurs on Anemone and Hepatica, and is known as

jEcidium punctatum.
Hollyhock Rust, Puccinia malvacearum (Fig. 72), is found over the
world, where the hollyhock, Althcea rosea, is grown.
RUST FUNGI 207
—
Fig. 77. Longitudinal section of a partly'gelatinous teliosorus'after the exten-
sion of the tentacles. (After Reed, H. S., and Crahill, C. H., Techn. Bull. 9, Va.
Agric. Exper. Slat., May, 1915.)
208 MYCOLOGY
Belonging to the genus Gymnoconia (Fig. 92) is the orange rust

of raspberry and blackberry which is found throughout the United
States and Canada. It is also widely distributed in Europe and Asia.
The genus Phragmidium, which is confined entirely to plants of the
rose family, is autoecious. Warts are formed on the teliospores by the
contraction of an outer gelatinous layer which with a rigid middle
lamina and the arrangement of the germ pores distinguishes Phrag-
—
FiG. 78. Teliospores of cedar apple showing germination with formation of
basidia (promycelia) and basidiospores (sporidia). {After Reed, H. S., and Crabill,
midium from neighboring genera. The teliospores are two- to several-

celled by transverse septa. An important species is the Rust of Roses,
Phragmidium subcorticium, which has a spindle-shaped teliospore with
six to eight cells.
Gymnosporangium is a genus of heteroecious rusts the secia of which

occur on Rosacea (except one on Hydrangeac^ and one on Myri-
RUST FUNGI log
cace.e) while the three-, four or five-celled teliospores are found

on CupRESSiNEiE {Chamcecyparis Cupressus, Juniperus, Libocedrus).
,
One autoecious species is G. bermudianum which produces both its

ascia and telia on junipers (/. bermudianum) Kern gives thirty-two
.
species as the number for North America and in vol. 7, North American
Flora, part 3, pages 188-190, gives a useful key for the identification
of the species.
Gymnosporangium botryapites causes fusiform swellings on the white
cedar, Chamcecy parts thyoides, on which swellings the two- to four-
FlG. 79- — Cedar rust on apple, roestelia stage with pustules.^ {After Jones and
Bartholomew, Bull. 257, Agric. Exper. Stat., Univ. Wise, July, 1915.)
celled teliospores are formed. The aecia occur on two species of shad
bush: Amelanchier canadensis and A. intermedia (Fig. 73).
In Gymnosporangium nidus-avis, the telia arise from a perennial
mycelium which often dwarfs the young shoots and causes bird's-nest
distortions inwhich usually there is a reversion of the leaves to the
juvenile form, sometimes causing gradual enlargements in isolated
areas on the larger branches of Juniperus virginiana with gecia on
several species of Amelanchier (Fig. 73).
Juniperus communis is the host of the telial stage of G. clavaricBforme,
which appears on long fusiform swellings of various-sized branches,
14
,
MYCOLOGY
scattered, or aggregated and its aecia on seven species of Amelanchier

one each of Aronia and Cydhnia.
Gymnosporangium Ellisii (Figs. 74 and 75) in its telial form distorts
the younger branches of the white cedar, ChamcBcyparis thyoides, pro-
FiG. 80. — Roestelia,or aecia on apple leaf. {After Giddings and Berg, Bull. 257,
Agric. Exper. Stat. Univ. Wise, July, 1915.)
ducing numerous fasciations. The aecia and pycnia of this fungus

are on Myrica. Gymnosporangium remarkable in forming
globosiim is
aecia on eighty-five different species of hawthorn, Cratcegus, while its

J
RUST FUNGI
teliospores appear on irregular spheric swellings or excrescence on

Jitiiiperus virginiana.
The mycelium of G. jimiperi-virginiance is annual, or biennial,
producing globose swellings known as cedar apples on the leaves of the
— ;:
3= 5
j* 'it
212 MYCOLOGY
a copious bibliography is one by HowardL. Reed and C. W. Crabill issued

as Technical Bulletin 9 (May, 1915) by the Virginia Agricultural Experi-
ment Station. The 106 pages of text are devoted to a careful considera-
tion of all aspects of the disease, which is prevalent throughout the
geographic range of the red cedar. The aecia are found on the apple
and were originally described as RoesteUa pyrata (Schw.) Thaxter,
and frequently the apple stage is known as the RoesteUa stage (Fig.
81). Infection of the leaves (Fig. 80) and fruit is only possible
during their undeveloped condition and not all varieties of apple are
susceptible. Some are rust free. Such are Early Harvest, Golden
Pippin, Winesap, while the badly affected varieties are Grimes Golden,
Smokehouse and York Imperial. The aeciospores are dark brown,
—
Fig. 82. Diagram (left) of aecium (roestelia) of apple rust; right, three Kcio-
spores from the cup highly magnified. {AfUr Jones, L. R., and Bartholomew, E. T..
Bull. 257, Agric. Exp. Stat., Univ. Wise, July, 1915.)
minutely pitted and almost spheric with thick walls and granular con-
tents. The first aecia (Figs. 81 and 82) become mature during the
month of July and viable spores are produced in large numbers during
this and the following two months (Fig. 83). This is the period of
infection of the red cedar, and the mycelium formed from these spores
remains dormant in the cedar leaves until the following spring, when
the cedar apple (Fig. 76), or gall, is formed out of the parenchyma
of the red cedar leaf (Fig. 161). Into the gall a vascular strand extends.
The surface of the galls becomes papillate and in May these papillae
enlarge into gelatinous horns, or teliosori (Fig. 77), made up of the
agglutinated stalks of numerous teliospores (Fig. 77), which are two-

celled and measure 46 to 63/x by 15 to 20/x (Fig. 78). These telio-
RUST FUNGI 213
spores on germination produce a four-celled basidium (Fig. 78),

or promycelium, from which are cut off basidiospores, which infect the
partially developed apple leaves, or apple fruits (Fig. 79). The dis-
ease apparently does little damage to the red cedar trees, but the
214 MYCOLOGY
aecial stage devastates the apple orchards found in proximity to red

cedar trees infected with the rust. Destroying the red cedar trees
seems to be the only feasible plan of combating the disease.
BIBLIOGRAPHY OF THE RUSTS
Arthur, J. C: Cultures of Uredineje. I (1899), Botanical Gazette, 29: 268-276;

II (1900 and Mycology, 8: 51-56; IV (1903), Journal of My-
1901), Journal of
cology, 10: 8-21; V (1904), Journal of Mycology, 11: 50-67; VI (1905), Journal
of Mycology, 12: 11-27; VII (1906), Journal of Mycology, 13: 189-205; VIII
(1907), Journal of Mycology, 14: 7-26; IX (1908), Mycologia, i: 225-256; X

(1909), Mycologia, 2: 213-240; XI (1910), Mycologia, 4: 7-33; XII (191 1),
Mycologia, 4: 49-65; XIII (1912, 1913 and 1914), Mycologia, 7: 61-89;

XIV (1915), Mycologia, 8: 125-141.
Arthur, J. C. and Holway, E. W. D.: Description of American Urcdinea;. Bull.
Lab. Nat. Hist, of State Univ. of Iowa, I, 3: 44-57; Hj 4:"377-402; III, 5: 171-
193; IV, 5:311-334-
Arthur, J. C.: Clue to Relationship among Hetercecious Plant Rusts. Botanical
Gazette, 33: 62-66, January, 1902.
Arthur, J. C: The Uredineae Occurring upon Phragmites, Spartina and Arundinaria
in America. Botanical Gazette, 34: 1-20, July, 1902.
Arthur, J. C: Problems in the Study of Plant Rusts. Publ. 22, Botanical Society
of America, Dec. 31, 1902, 1-182.
Arthur, J. C: Taxonomic Importance of the Spermogonium. Bui. Torr. Bot.
Club, 31: 125-159, March, 1904.
Arthur, J. C: Terminology of the Spore Structures in the Uredinales. Botanical
Gazette, 39: 219-222, March, 1905.
Arthur, J. C: Amphispores of Grass and Sedge Rusts. Bull. Torr. Bot. Club, ^,2:
35-41, 1905.
Arthur, J. C. and Kern, F. D.: North American Species of Peridermium. Bull.
Torr. Bot. Club, 33: 403-438, 1906.

Arthur, J. C: Eine auf die Struktur und Entwicklungsgeschichte begriindete
Klassifikation der Uredineen. Rusultats Scientifique du Congres international
de Botanique Wien, 1905: 331-348, 1906.
Arthur, J. C: New Species of Uredinese. Bull. Torr. Bot. Club, I, 23: 661-666,
December, 1901; II, 24: 227-231, April, 1902; III, 31: 1-8, January, 1904;
IV, 33: 27-34, 1906.
Arthur, Joseph C: Uredinales. Coleosporiaceas, Uredinaces, ^cidiace^ (pars).
North American Flora, 7, part 2, 1907; /Ecidiacese (continuatio), 7, part 3, 191 2.
Arthur, J. C: On the Nomenclature of Fungi Having Many Fruit Forms. The
Plant World, 8: 71-76; 99-103.
Blackman, H. v.: On the Fertilization, Alternation of Generations, and General
Cytology of the-Uredinese. Annals of Botany, xviii: 323-373) i904-
Blackman, H. V. and Eraser, Miss H. C: Further Studies on the Sexuality of
the Uredinea;, with 2 plates. Annals of Botany, xx: 35-47, 1906.

RUST FUNGI 215
Carleton,. Mark A.: Investigations of Rusts. U. S. Dept. Agr., Bureau of Plant

Industry Bull. 63, 1904.
Carleton, Mark A.: Lessons from the Grain Rust Epidemic of 1904. U. S. Ucpt.
Agr., Farmers' Bull. 219, 1905.
Cheistman, a. H.: Sexual Reproduction in the Rusts. Botanical Gazette, xx.xix:
267-274, 1905.
Christman, a. H.: Observations on the Wintering of Grain Rusts. Trans. Wise.
Acad. 98-107, 1905.
Sci., 15:
Coons, G. H.: Some Investigations on the Cedar Rust Fungus, Gymnosporangium

Ann. Rep. Neb. Exp. Sta., 25: 217-245, pis. 1-5, 191 2.
juniperi-virginianas.
DE Bary, Anton: Comparative Morphology of the Fungi Mycetozoa and Bacteria,
1887: 274-286.
DiETEL, P.: Uredinales. Die natiirlichen Pflanzenfamihen I, Teil i, Abteilung**:
24-81, 1900.
DuGGAR, Benjamin M.: Fungous Diseases of Plants, 1909: 384-438.
Eriksson, Jakob: Fungoid Diseases of Agricultural Plants, 191 2: 63-89.
Eriksson, Jakob: On the Vegetative Life of Some Uredineae. Annals of Botany,
xix: 55.
Freeman, E. M.: A Preliminary List of Minnesota Uredineae. Minn. Bot. Studies,
2, part 4: 407, 1901.
Grove, W. B.: The British Rust Fungi (Uredinales): Their Biology and Classifica-
tion, 1913, pp. i-x + 1-412.
Heald, F. D.: The Life History of the Cedar Rust Fungus Gymnosporangium
juniperi-virginianas. Ann. Rep. Neb. Agric. Exp. Sta., 22: 105-113, pis. 1-13,
1909.
Hoffmann, A. W. H.: Zur Entwicklungsgeschichte von Endophyllum sempervivi.
Centralbl. fur Bakteriologie, 2, abt. 32: 137-158, 191 2.
Hume, H. Harold: Some Peculiarities in Puccinia Teleutospores. Botanical

Gazette, 1899: 418-423.
Kern, Frank D. : A Biologic and Taxonomic Study of the Genus Gymnosporangium.
Bull. N. Y. Bot. Gard., 7: 391-494, pis. 151-161, 1909-1911.
Kern, Frank D.: Gymnosporangium. North American Flora, 7, part 3: 188-21 1.
Klebahn, H. Die wirtswechselnden Rostpilze, 1904.

:
Kunkel, Louis Otto: Nuclear Behavior in the Promycelia of Caeoma nitens

Burrill and Puccinia Peckiana. Howe Amer. Jour. Bot., i: 37-47, January,
1914.
KURSSANOW, L.: Zur Sexualitat der Rostpilze. Zeits. Bot., 2: 81-93, iQio-
Magnus, P.: Weitere Mittheilung iiber die auf Farnkrantem auftretenden Uredi-
neen. Berichten der Deutschen Botanischen Gesellschaft, xix. Heft 10: 578-
584, 1901.
Magnus, P.. Ueber eine Function der Paraphysen von Uredolagern, nebst einen
Beitrage zur Kenntniss der Gattung Coleosporium. Berichten der Deutschen
Botanischen Gesellschaft, xx. Heft 6; 334-339, 1902.
Massee, George: Diseases of Cultivated Plants, 1910: 289-338.
Massee, George and Ivy: Mildews, Rusts and Smuts, 1913: 52-182.
McAlpine, D.: The Rusts of Australia. Dept. Agric. Victoria, 1906.
2l6 MYCOLOGY
Olive, Edgar W.: Sexual Cell Fusions and Vegetative Nuclear Divisions in the
Rusts. Annals of Botany, xxii: 331-360, 1908.
Olive, Edgar W. : Origin of Heteroecism in the Rusts. Phytopathology, i: 139-
149, October, 191 1.
Olive, Edgar W.: Intermingling of Perennial Sporophytic and Gametophytic
Generations in Puccinia Podophylli, P. obtegens and Uromyces Glycyrrhizas.
Annales Mycologici, ii: 297-311, August, 1913.
Pritchard, F. J.: A Preliminary Report on the Yearly Origin and Dissemination
of Puccinia graminis. Botanical Gazette, 52: 169-192, 1911.
Reed, Howard S. and Crabill, G. E.: The Cedar Rust Disease of Apples Caused
by Gymnosporangium juniperi-virginianse. Tech. Bull. 9, Virginia Agric. Exper.
Stat., 1915.
Sappin-Trouffy, P.: Recherches histologiques sur la famille des Uredinees. Le
Botaniste, 5: 59-244, 1896.
Stewart, Alban: An Anatomical Study of Gymnosporangium Galls. Amer.
Journ. Bot., 2: 402-417 with i plate, October, 1915.
Sydow, Paul H.: Monographia Uredinearum seu-specierum omnium ad hunc usque

diem descriptio et adumbratio systematica auctoribus, 1904.
Tulasne, L. R.: Second Memoire sur les Uredinees et les Ustilaginee. Ann. Sci.
Nat., iv. 2: 77, 1854.
von Tavel, Dr. F.: Vergleichende Morphologic der Pilze, 1892: 123-133.
VON TuBEUF, Dr. Karl F.: Pfianzenkrankheiten, 1895: 340-434.
Ward, H. Marshall: Illustrations of the Structure and Life History of Puccinia
graminis. Annals of Botany, ii: 217 with 2 plates.
Ward, H. Marshall: On the Relation between Host and Parasite in the Bromes
and their Brown Rust, Puccinia dispersa. Annals oT Botany, xvi: 233, 1902.
VON Wettstein, Dr. Richard R., Handbuch der Systematischen Botanik, 1911:
196-202.
Suborder Auricularine^. —Family Auriculariace^. —^The

fungi of this family are saprophytes, or v^ôod-inhabiting parasites. The
basidia are borne directly on the mycelium, or in variously formed fruit
bodies in which the basidia form a layer. The basidia are transversely
divided into four cells. Auricularia includes about forty species of
which the best known is, Auricularia (Hirneola) Auricula Judce, the Jew's
ear fungus, which develops its fruit body on rotten wood. When wet, it
is gelatinous; when dry, it appears as a dry crust. It is a rather gelatin-
ous, flabby-looking, thin expanded cup or saucer-shaped fungus of

a brownish color when expanded smooth inside, veined and plaited so
as to have the resemblance to a human ear. It grows on a variety of
trees: elm, maple, hickory, balsam, spruce and alder and up to 1900,
it had been collected in Ohio, Maryland, Indiana, New Jersey, Pennsyl-
vania and West Virginia. Outside it is velvety and grayish-olive.

GELATINOUS FUNGI 217
Auricularia (Hirneola) polytricha is the "Mu-esh" of the Chinese, who

gather it as an article of food, in fact oak boughs are cut and allowed to
decay to raise the fungus.
—
Family Pilacrace^. ^This is a small family of two genera, Pila-
crclla and Pilacre, with spheric stalked fruit bodies. The basidia are
in capitate clusters and surrounded at first by a peridium-Uke wall,
which breaks at maturity.
Suborder Tremelline^. —Family Tremellace^. — This family
includes twelve genera, of which Trcmella most important. The
is the
majority are widely distributed and live saprophytically on wood, where
they appear as soft, trembhng, gelatinous masses, when moist, becoming
rigid and horny when dry. The basidia are longitudinally divided by
two septa. The four portions thus formed each bear a terminal basidio-
spore. Some species of Tremella produce conidiospores. Tremella
frondosa has been used as food, but as such is unsatisfactory. Tremella
foliacea is of a smoky-brown color, cold, clammy and trembles in the
hands. When stewed, it becomes a shmy mess relished only by the
Chinese. Tremella mesenterica is brain-like in its convolutions, ge-
latinous in texture and usually the size of a walnut, and of an orange-red
color.
CHAPTER XX
FLESHY AND WOODY FUNGI
Suborder Eubasidii. —The fungi of this suborder are characterized
by the undivided (unseptate) basidia, more or less club-shaped with
generally four, rarely two apical sterigmata each of which
six, eight, or
bears a basidiospoi-e (Fig. 92). These fungi are usually fleshy and the
spores are borne openly on wrinkles, ridges, gills, in pores, on spines,
or in closed fruits, which open regularly, or irregularly, by splitting.
Many of the forms are edible, some are inedible, because of toughness,
or woodiness, while others are poisonous.
—
Cytology. Recent studies by Juel (1897), Maire (1900), Ruhland
(1901), Harper (1902), Levine (1913) have shown that as a general
thing the hyphal cells of the mycelium in the HYMENOMYCETES
and GASTEROMYCETES are binucleate, and sometimes, as in Cop-
rinus radiatus, uninucleate. The cells of the young carpophore are
binucleate, but as the fruit body matures, the majority of the cells in
the stipe and pileus are multinucleate, but this condition arises from
the amitotic fragmentation of the two nuclei originally present in each
cell. The subhymenial cells from which the basidia spring and the
paraphyses are always binucleate. All the cells, which are concerned
directly with the production of basidiospores, are binucleated through-
out their development. The multinucleated condition above noted
development and are found in the organs
arises in cells of strictly limited
of nutrition, support, transportation, etc. Maire found that the pairs
of nuclei divide simultaneously, as conjugate nuclei, so that in the suc-
cessive cell generations which arise in the development of the carpo-
phore the two nuclei in each cell are of widely separated nuclear ances-
try, duplicating exactly the condition found in the rusts previously
described. The young basidium contains only two nuclei just as in
the teliospore of the rust. These two nuclei fuse to form the primary
nucleus of the basidium which then divides twice to furnish the nuclei
for each of the typically four basidiospores. Levine (191 3) who has
studied this nuclear division in a number of species of Boletus, finds the
218
FLESHY AND WOODY FUNGI 219
long axes of the spindles in both divisions are commonly transverse to

the long axis of the basidium. The spores in all of the forms studied
by him are uninucleate at first. Just when the mycelial cells become
regularly binucleate has not been certainly ascertained except in a few-
forms. Presumably in Coprinus radiatus the uninucleate spores give
rise to uninucleate hyphal cells, but Levine finds in his Boletus studies
that the primary spore nucleus divides at once to form two nuclei.
Presumably, the nuclear division in other forms may be delayed, until
the primary mycelium has arisen. An alternation of generations com-
parable to that of the rusts is also present in the Hymenomycetes and
Gasteromycetes. The sporophyte begins at some indefinite point in

the mycelium and extends through the development of the carpophore.
—
A. Hymenomycetes. The undivided basidia of these fungi bear
four basidiospores perched on corresponding points, or sterigmata.
These basidia spring directly from the mycelium in the primitive forms,
but in the more highly evolved types, the basidia are borne on definite
layers (hymenial layers) together with the paraphyses and cystidia
characteristic of some of the forms. The hymenia are carried by special
fruit bodies which differ structurally in the different f amiUes. These
fruit bodies arise from a profusely branched mycelium, which radiates
through the organic substratum, which may consist of leaf mold, rotten
wood, dying tree trunks, and manurial waste. The hyphal cells are
frequently united by clamp connections which probably give greater
strength to them. Such are the saprophytes. Some of the hymeno-
mycetous fungi are parasites and five in the bark and wood of trees,
and some few are parasitic on the woody parts, leaves, flowers and
developing fruits of certain shrubs. Sometimes, as in Armillaria mellea,
the hyphae become united in strands with apical growth. These strands
are known as rhizomorphs and serve in part as the resting organs. True
sclerotia are also formed. The fruit bodies take various forms. The
most highly developed types with stalk, cap and gills are known as
toadstools. Some of the simple forms are club-shaped. Others have
spines and pores instead of gills over which the hymenia are spread.
—
Family i. Dacryomycetace^. The fruit body is gelatinous, or
cartilaginous, and of different shapes. The whole surface of the fructi-
ficationis covered with a paHsade-like layer of long club-shaped basidia
which bear two-forked basidia, each fork with a basidiospore. Conidio-

spore formation occurs in a number of forms. The important genera
220 MYCOLOGY
are Dacryomyces, Guepinia, Calocera. Dacryomyces deliquescens forms

gelatinous, or gristly, lumps on tree stumps. Guepinia peziza is sapro-
phytic on oak stumps. Calocera viscosa is a branched upright form
suggesting the true coral fungi.
Family 2. Exobasidiace^. —The mycelium of the fungi of this
family Hves parasitically in the chlorenchyma of many shrubs. The

fruit body is a thin basidial layer, which breaks out of the tissues of the
host. Each basidium develops four basidio-
spores; rarely 5 to 6 are formed. Some of the
species form galls on the stems, leaves and
flowers of ericaceous shrubs, such as species of
Vacciniiim, Rhododendron, Azalea, Andromeda,
etc. There are two genera: Exobasidium (Fig.
84), with eighteen species; and Microstroma,

with two species. Exohasidium caccinii (Fig.
84) develops swellings on the leaves of species

of Vacciniiim of a whitish-red color. Its
basidia are club-shaped with four sterigma
and four basidiospores. The basidiospores are
spindle-shaped, 14 to i6yu long by 2 to 3^
broad, colorless and smooth. Exohasidium

rhododendri forms enlargements of the leaves
of species of Rhododendron of greater or less
size; colored white, or flesh-colored. Ex-
Fig. 84.— Floral gall
ohasidium ledi occurs on Ledum in Finland.
produced on flowers of
huckleberry, Exohasidium andromedcB grows on leaves and
Gaylussacia
by Exohasidium ^^j^g ^f gpecies of Andromeda in Europe and
resinosa,
vaccina. Note enlarged
and swollen calyx.
°
(Pine America.
,.,.
,
'-
Exooasidium AzalecB
...
found
IS on
mTT^' f î6T"^' ^' ^" ^P^"^^ ^^ ^"^^^^^^ ^^ ^^^^^ America. E.v-

obasidium lauri forms widely spread, yellow
then brownish, horny, or club-like on the stems of the laurel in
galls
Italy, Portugal and the Canary Islands. Exohasidium Warmingii

attacks the living leaves of Saxifraga aizoon in Greenland, Tyrol and
north Italy.
Family 3. Hypochnace^. —The hymenium is cobwebby. The
basidia have two, four or six sterigmata. Cystidia are sometimes present.
Hypochnus occurs on old stumps, on leaves and on mosses. Tomen-
tella is another genus.
—
Family 4.. Thelephorace.^. — Fruit bodies of a simple type are

found in this They form on three trunks, either flat
family.
leathery crusts with the hymenium on the smooth upper surfaces, or the
flat fructifications are raised above the substratum and have bracket-
like outgrowths, which show an overlapping arrangement with the

hymenial layer on the under side. The important genera are Corticium,
Stereum and Thelephora. In Corticium, the fructification is leathery,
membranous, fleshy, rarely wholly gelatinous, crust-like, growing resu-
FiG. 85. a piece of old oak timber rotted by Slereum fruslulosum showing scat-
tered frvtiting bodies. {After von Schrenk, Hermann, Bull. 149, U. S. Bureau of
Plant Industry, 1909.)
pinate. The hymenium is smooth, or pimply, and consists of club-

shaped basidia with four basidiospores. The species are mostly
found on wood. C. vagiim-solani in its sterile form is known as Rkiz-
which apparently has been found on sugar beet, bean, carrot,
octonia,
cabbage, potato, egg plant and a number of other hosts. The hymeno-
phore of this species is white with short basidia and elliptic spores.
It frequently entirely surrounds the green stems of its host near the
ground. The persistent hymenophore of Stereum is leathery, or
22 2 MYCOLOGY
woody, attached laterally, or centrally, sometimes as a bracket with a

smooth hymenium. Stereum hirsutum attacks oak trees in which the
wood becomes brownish at first and in longitudinal section, white or
yellow streaks are found, hence the common name white-piped, or
yellow-piped oak.In the cross-section, these streaks are white specks,
and another name, that of "fly wood," is apropos. Further decom-
position follows. The rot of woods, known as partridge wood, where
the timber becomes speckled with white, is due to Stereum frustulosum
Fig. 86. — Coral-like fruit-bodies of Clavaria flava. {Photo hy W. H. Walmsley.)
(Fig. 85). The fruiting bodies are hard and crust-like, light brown to
grayish in color.
, The smothering fungus of seedlings is Thelephora
terrestris and T.laciniatum. Soft leathery masses are found at the base
young trees of the hard maple. These are numerous, shelf-like fruit of
bodies, hemispheric in shape and in -mass may completely surround
and smother the small tree. Hymenochcete noxia attacks tropic plants,
such as cocoa, tea, bread fruit, camphor and the like.
Family 5. Clavariace^. —The fairy clubs, or coral funguses belong

here. The simple, or branched, club-shaped or antler-like hymeno-
.
phores are fleshy, leathery, cartilaginous, or waxy. The basidia are

clavate, interspersed with cystidia and bear one to four sterigmata.
Pistillaria, Typhula, Clavaria and Sparassis are important genera.
Many of the species of Clavaria are edible (Fig. 86), but some of them
are tough and leathery. The color varies, as noted in the enumeration
of common American species given below:
Clavaria flava (paleyellow) (Fig. 86).
Clavaria aurea (golden).
Clavaria botrytes (red-tipped).
Clavaria cristata (crested).
Clavaria cinerea (ashen).
Clavaria aurantio-cinnabarino (orange-red)
common species, has its hymenial ridges pro-
Sparassis crispa, a
jecting and much convolute, suggesting a mammalian brain. It is too
tough to be edible.
Family 6. Hydnace^. — The highest forms of this family possess
the form of a mushroom, while others are sessile and are resupinate,
others without a distinct cap are effused. The hymenium is spread
over with persistent bristles, teeth, tubercles or spines. The most
important genera are Phlebia, Radidum, Grandinia Irpex and Hydnum
(Fig. 87). The edible forms are included in the last two genera.
The forms of Hydnum are extremely variable. The highest forms,
such as Hydnum repandum, have a cap with a central stipe, while
in other forms it is lateral, or absent. In some of the lower forms,
the pileus is resupinate. Projecting spines are covered with the
hymenial surfaces. A rot of hardwoods is due to Hyd-
in America
num H. diversidens with its yellowish-white sporophore
coralloides.
takes the form of an incrustation,ôr bracket with downward-projecting
spines of unequal length. The hymenium renews itself by a new
hymenium growing through the old one. It causes a decay of timber
known as white rot. Hartig gives a careful description of it, as it occurs
in Europe. H. caput-ursi is a bracket form growing as excrescences on
living oak pendulous spines at first white, then becoming
trees with its
yellowish and brownish. H. caput-medusce. has pendulous tufts of
white to gray spines and is found on elms and oak trees. The spiny
character of H. erinaceum (Fig. 87) suggests a hedgehog, hence its
specific name. The last three are fleshy and edible. Irpex differs
from Hydnum in having the spines connected at the base, and in
224 MYCOLOGY
in their being less awl-shaped and pointed. /. obliquus on stumps, /.

carneus on tulip poplar, /. fusco-violaceus on pine trunks are American
species.'
Family 7. Polyporace^. —The fruit body of the fungi of this
family are of various substance and shape. The hymenium lines the
inner surface of pores, or grooves, or is spread over the under surface of
the fruit body. The depressions are either united vein-like grooves,
tubes, or honeycombed cells, or twisted passages. Concentrically
PLESHY AND WOODY FUNGI 225
three species of which M. lacrymans, the dry-rot fungus, is most impor-

tant. This fungus is of world-wide distribution, where it attacks
structural wood work and timbers. It has been so long associated as
a destructive agent with the structural wood work of men, that it was
supposed to be an entirely domesticated form and not known to exist
in the wild form. Recent investigations have shown that it occurs on
living trees, which when used for structural purposes furnish wood
which is liable to destruction later on. The mycelium of Merulius
lacrymans (Fig. 88), usually gains access to dressed boards, joists, or
Fig. 88. — Immature fruiting stage of dry-rot fungus {Merulius lacrymans) de-
veloping on the front of a board. {After Clinton, G. P., Rep. Conn. Agric. Exper,
Slat., pi. xxviii, 1906.)
rafters by the germination of one of its spores at a point where the beam
may be in contact with a damp wall. Its mycehum penetrates the wood
and usually grows lengthwise at first, the water for its extension being
supplied by larger more tube-like hyphae known as the conductive hyphte,
which carry water to the extreme end of the mycelial growth. The pres-
ence of the fungus results in a decay of the wood, which is reduced to a
brown punky mass, that crumbles between the fingers. When the myce-
lium comes to the surface of the wood, it forms a white felt-like coyering
studded with water drops, hence the specific name lacrymans referring
IS
226 MYCOLOGY
to the tear-like drops of water pressed out of the Hving hyphal cells.
The mature sporophore an amber-brown color covered with anasto-
is
mosing wrinkles (Fig. 89) over the surface of which the basidia bearing
basidiospores are borne. Two basidiospores are borne on pointed
sterigma by each basidium. As the fungi by means of its conducting
hyphag is independent of local water supplies, it can grow in wood, even
if protected by an external coat of paint, or varnish, and the builder is
chagrined to find such wood work crumble away beneath the coats of
paint. Mycodendron is a curious fungus with a fruit body which
suggests a muffin stand, or a pagoda with superimposed, rounded.
-Fruiting stage of dry-rot fungus (Merulius lacrymans). {After Clinton,

G. P., Rep. Conn. Agric. Exper. Stat., pi. xxviii, 1906.)
spore-bearing shelves through which the central stalk runs from one-half
to the next above. Mycodendron paradoxum has been collected on
wood in Madagascar.
—
PoLYPOROiDE^. This Subfamily includes tough or woody fungi
found generally on wood as bracket-hke fruit bodies of different
forms and sizes. The spore-bearing surface, a hymenium, consists of
furrows, or tubes. In the perennial-fruited forms, the tubes are often
found in layers. Mycologists have made a natural division of the dif-
ferent forms of fruit bodies into those which are resupinate, the annual
peroi^ species, the perennial peroid forms and those species which are
like the agarics. The various forms are of interest to the scientific my-
cologist, but to the mycophagist they are of use as food. Only one
poisonous form known, and that is the medicinal one, Fames laricis,
is
but it is so bitter and unattractive, as not to be tempting. Some of

them are destructive to living trees, to timber used for mine props, and
structural purposes, and to wood exposed to the weather, or in contact
with the soil.
The ease with which the polypores are collected and preserved
makes them especially suitable for systematic study in the classroom.
Besides, they retain their characters when dried, so that the keys used
for their identification can be readily followed. Fortunately also we
have several manuals which cover the different sections of our country.
They are reasonable enough in price to be furnished for use in the class-
room. It is suggested that boxes of the different kinds used for this
purpose be filled with enough specimens to furnish each member of
the class in mycology with one specimen of each kind. There should
be a sufiicient number of manuals of the region, where the botanic
institute is situated, to supply every two members of the class with
one, so that the students may use them in groups of two. The
advertisement of the books is here reproduced for the use of teachers
of mycology.
MANUALS OF POLYPORES AND BOLETES

By William A. Murrill, A. M., Ph. D., Assistant Director of the New York
Botanical Garden, Editor of "Mycologia," and Associate Editor of "North American
Flora."
Northern Poljrpores, November, 19 14. Including species found in Canada and
the United States south to Virginia and west to the Rockies.
Southern Polj^jores, January, 1915. Including species found in the United
States from North Carolina to Florida and west to Texas.
Western Pol5T)ores, February, 1915. Including species found in the states on
the Pacific coast from Cahfornia to Alaska.
Tropical Polypores, March, 1915. Including species found in Mexico, Central
America, southern Florida, the West Indies, and other islands between North
America and South America.
American Boletes, November, 19 14. Including all the species found in temperate
and tropical North America, both on the mainland and on the islands, south to
South America.
As satisfactory keys of the different genera and species of the poly-

pores and boletes are given in these manuals, and as it is presupposed
that their use will be adopted, keys of the more common genera and
228 MYCOLOGY
species are not given space in this book. It should be stated, however,
that Murrill classifies his genera and species differently from the authors
that have preceded him where many of the new genera were classified
under the genera Polyporus and Boletus (Fig. 90). The arrangement
of Murrill seems to be a more satisfactory presentation of these groups
than those systems which have gone before and is founded on more
natural characters. The nomenclature which this author adopts in

the several recommended manuals was f()rcshad(nved in \o\. i)art i (>,
Fig. 90.~7)<j/«7;(s /,//, ,i\ in three stages of devel. iinnent. {After Patterson, Flora W. ,
and Charles, Vera K., Bull. 175, U. S. Dept. Agric, pi. x.xxi, Apr. 29, 1915.)
(1907), and part 2 (1908) of the "North American Flora," where keys
will also be found with the synonymy which has been omitted from the
manuals. To connect satisfactorily, the old and the new generic and
names, the treatment of the Polyporace^ in the "North
specific
American Flora" should be consulted.
Trametes roblniophila is found on decayed spots of living trunks of
Rohinia pseudacacia from Pennsylvania to Virginia and Missouri, and
it doubtless causes decay of the wood. T. suaveoleus is found on willow
trees, where it causes serious decay. It has an agreeable odor. T.
subiiivosa is occasional on dead deciduous wood in Florida, Louisiana
and Mississippi. At New Orleans, it was collected on living water

oak and at Eustis, Fla. on cypress. The species become more abundant
in tropic America where nine have been found. T. jalapensis was col-
lected on a railway tie near Jalapa, Mexico. The species of Coriolus
are annual. It includes Coriolus {Polyporns) versicolor found on all
—
Fig. 91. Piece of dead wood with'sporoiiii-iK ,- ,.,.. iomenlaius. {After von
Schrenk, Hermann, Bull. 149, C/.?5. Bureau of Flant Industry, pi. viii, 1909.)
kinds of dead wood. It causes root rot in many trees and becomes a
wood parasite of Catalpa. It has a leathery, thin and rigid hymeno-
phore depressed at the point of attachment. The surface is velvety
and variegated with two-colored zones. The pores are minute rounded
with ragged edges, white then yellowish. Polyporus arcularius is com-
mon in the eastern United States on dead branches and trunks of vari-
MYCOLOGY
ous trees. P. caudicimis is one of the most dangerous enemies of shade

trees in Europe but, fortunately, it is rare in America.
The genus Fames includes the fungi with corky, woody, or rarely
punky hymenophore, which is sessile, hoof-shaped, or applanate (Fig.
91). The substance of the fruit body is white, flesh-colored, or wood-
colored. The tubes are cylindric and usually thick walled. Fomes
annosus will live on trunk and roots of coniferous trees, Fomes
(Pyropolyporus) igniarius causes serious heart rots of trees. It was
formerly the source of tinder. Dcedalea quercina (Fig. 202) is a corky,
or woody, species common on oak and chestnut trees. It is at first
porous, but these pores coalesce to form slits with blunt partitions.
It is very common about Philadelphia. Lenzites betulinus is common
on dead deciduous wood.
—
FiSTULiNOiDEiE, The most important genus of this subfamily is
Fistulina, which comprises about six species. F. hepatica is the com-
monest form, and is known by its English name beefsteak fungus or,
in French, langue de hoeiif. The tongue-shaped fruit body projects
from the tree and is six to ten inches across with a liver-colored and
sticky gelatinous surface. The mouths of the tubes are closely packed.
It is edible, when fully mature, its flavor resembhng beefsteak.
BoLETOiDE^. —The
members of this subfamily are tube-bearing
fungi differing from the Polyporoide^ in their fleshy substance and
terrestrial habit. They have a cap and stipe like a mushroom, but
porous tubes instead of gills on the under cap surface. They occur
usually in forested tracts during summer and autumn. The annual
hymenophore is usually centrally stipitate. Many of the best edible
fungi (few of them poisonous) are found in this subfamily, which in-
cludes, according to Murrill in North America, Central America and
the West Indies, as far as Trinidad, eleven genera.
Boletus (Tylopilus) felleus (Fig. 90) is common iji woodlands. It
is discarded as an edible form, because of its bitter taste. Forty-eight
species of Ceriomyces are listed by Murrill for America. The genus
Boletus proper is made to contain only five species, while Strohilomyces
strobilaceus still retains its old name. This rough shaggy form is
regarded as edible.
CHAPTER XXI
MUSHROOMS AND TOADSTOOLS
Family 8. Agaricaceê. —The mycelium of the fungi of this fam-
ily lives in the substratum, which may be the soil, leaf mould, rotten
wood, old stumps, dead tree trunks, or living trees, as far as the natural
environment is concerned, and in manures, in the decay of agricultural
plants in the fields, offal, spent tan bark and rubbish heaps, as far as
man has influenced the environment. The hyphse may be delicate and
cobwebby, thread-hke, cord-like, or in strands (rhizomorphs). They
are always septate, sometimes with clamp connections and their color
may vary from white to yellow, or brown {Ar miliaria mellea). The
fruit bodies are mostly fleshy, rarely of membranous, or leathery, con-
sistency. Usually of an umbelloid form, they may have a sessile cap,
or pileus, or the stalk, if present, may be attached laterally, although
it is placed centrally as a general rule. The hymenophore consists of
radiately arranged veins, folds, or gills (lamellae), which are generally
free from each other, seldom anastomosing, or dichotomously branched.

As the popular name toadstool is suggestive of the commonest form
of these fleshy fungi, a few words of explanation with regard to the
general structure will be apropos. Attached to the spreading myce-
lium we find arising vertically the stalk, or stipe. The height of this
varies in the different genera and species. Sometimes it is enlarged
at the base, at other times, the stalk is perfectly cylindric. The sur-
face of the stipe may be smooth, rough, reticulate, or stringy, and its
center may be solid, stuffed, or hollow, as may be. An annu-

the case
lus, such as is present in the common mushroom, may in other forms
be absent, or well developed. Placed on the stem, or
stipe, above we
find the cap, or pileus, which
expanded horizontally. It has a
is
domed, convex upper surface sometimes with a projecting boss, or

umbo; in other forms it is depressed (crateriform, umbilicate, etc.).
The gills, or lamellae, are attached to the lower surface of the pileus.
They may run from the stipe to the margin, or they may run only
part way, so that frequently there are secondary gills alternating with
231
232 MYCOLOGY
the primary ones. The gills may be free from the stipe, adnexed, or
even decurrent.
A section of a mature gill shows the following disposition of the
hyphal layers. The central part of the down-

gill consists of parallel,
ward directed hyphffi, that form the trama. Running out obliquely
from the trama are shorter cells which constitute the subhymenium.
The basidia, together with their accompanying paraphyses and cysti-
dia, form a palisade-like layer (the hymenium) whose cells stand at
right angles to the tramal hyphae.
The basidia are furnished with
sterigma, which bear the basidio-
spores (Fig. 92). In such forms
as the common mushroom, the
gill chamber is at first closed by
a veil known as the partial veil,
or velum partiale, which ruptures
when the pileus expands. The
part of this membrane attached
to the stipe becomes the annulus.
while the other part remains at-
tached in a shreddy condition
to the edge of the cap. The
species of A manita have a univer-
sal veil which covers the whole
fruit body, and as this enlarges
the velum universale is torn trans-
FiG. 92.-Coprinus siercorarius with versely, the lower part forming

young and mature sporophores with gills, the death CUp, Or volva, and the
basidia and basidiospores and cystidia. ,• . .
"PP^r P^rt sometimes remammg

,
{After Brefeid.)
in the form of flaky pieces, which
are distributed irregularly over the upper surface of the cap (Fig. 93).
A
frill-like annulus is also found at the top of the stipe in the Amani-
tas. It does not represent a portion of the partial veil in the Amanitas,
but is a membrane which is formed from a thick, loosely felted
layer, which separates as elongation proceeds from the surface of the
stipe, retaining its connection with the stipe where the stalk joins the
cap. It is pulled away from the stipe by retaining its connection with
the edges of the pendant gills as a continuous membrane, which covers
MUSHROOMS AND TOADSTOOLS 233
the gills. As the pileusexpands the membrane becomes detached first

at the margin of the cap,and it falls down around the stipe, as a frill,
plaited in delicate folds, corresponding to the former lines of contact
with the lamellae and is now known as the annulus superus, frill, or
armilla. Special milk tubes are found in such forms as species of Lac-
when these toadstools are wounded a milky fluid oozes out in
tarius for
drops. Each basidium usually bears four basidiospores, sometimes
thereare two. The color of these spores is distinctive, and is used in
Fig. 93. — Deadly amanita (Amanita mitscaria) showing volva at base of stem
and frill, like stem ring. {After Chestnut, V. K., Bull. 175, U. S. Dept. Agric, pi. i,
Apr. 29, 1915-) J
the classification of the genera of the family. We distinguish the

white-spored, rosy-spored, ochre-spored (yellow or brown), brown-
spored, black-spored agarics.
Fungi" (1909) has carried on detailed
Buller in his "Researches on
studies with numerous gill fungi and has studied the physi-
species of
ology and mechanics of spore discharge and fall. The disposal of the
hymenium beneath a pileus on gills, the rigidity of the fruit body, the
growth movements of the fruit body, all faciUtate the distribution of the
discharged basidiospores. The spores liberated from a pileus in per-
234 MYCOLOGY
fectly still air placed above a horizontal sheet of paper fall vertically
downward and produce a spore print of radiating lines of spores cor-

responding to the interlamellar spaces. The number of spores liberated
in Agaricus (Psalliota) campestris (Fig. 94), 8 cm. in diameter, was
1,800,000,000 spores. Coprinus comatus formed 5,000,000,000 spores.
Such discharge under normal conditions is continuous, but by exposing
the gills to ether, or chloroform vapor, it ceases. BuUer determined
that the four spores on each basidium are discharged successively leav-
ing the sterigmata a few seconds or minutes of one another, so that an
entire mushroom will discharge in total about a million spores a minute
Fig. 94. — Meadow mushroom, Agaricus campestris. A, View of under surface;

a, annulus; g, gills; B, side view; s, stipe; p, pileus or cap. {From Gager, after W. A.
Murrill.)
fortwo or more days. The rate of fall of hymenomycetous spores ranges

from 0.3 to 6.0 mm. per second; those of the mushroom shortly after
they have left the gills fall at a speed approximately i mm. per second.
The path described by a spore in its fall has been called a sporabola.
Buller has divided the fruit bodies of the Agaricace^ into two types,
the Coprinus comatus type and the Agaricus campestris type. The
deliquescence in the first type is an autodigestion, which renders impor-
tant mechanic assistance in the process of spore discharge, where the
process proceeds in succession from below upward, so that autodiges-
tion refnoves those parts of the gills from which the spores have been
—
discharged, and permits the spores to fall more easily past the neighbor-
ing gill surfaces.
Development of the Fruit Bodies.—Kikmson^ has studied the develop-
ment of the mushroom {Agaricus {PsaUiota) campestris) (Fig. 94).
The youngest stage is the homogeneous primordium 01 the carpophore
composed of slender, uniform, dense hyphae, intricately interwoven, and
surrounded by a thin layer of hyphse of a looser arrangement. This
layer is the universal veil which grows until the form of the fruit
appears when it is torn into white floccose patches on the pileus. In
the very young primordium then there is no evidence of a differentiation
into stem and pileus and at this stage stained longitudinal sections show
two small deeply stained internal areas near the upper end of the young
fruit body and some distance from the surface. The hyphae here are
richer in protoplasm and form an annular area within the fruit body.
This area now increases in extent and many hyphae grow from its
upper portion downward to form the primordial layer of the hymenium.
These downward growing hyphge are slender and terete and taper
pointed, which enables them to push between the surrounding hyphse.
Soon after these hymenial hyphse grow downward there is a cessation of
growth. Just below this area which results in the rupture and separa-
tion of the hyphse at this point in a corresponding internal annular area,
forming the well-known "gill cavity" which at first is very minute.
With the formation of this annular primordium of the hymenium
the primordia of the stem, veil and pileus are differentiated. The
period of elongation of the parts after they have been organized follows
in succession. The marginal veil completes its period of elongation
first, then the stem, followed by the pileus, and finally, the hymenium
where in examples studied Atkinson secured two-spored basidia.
A somewhat similar development takes place in Agaricus Rodmani,
a form which grows in grassy ground and paved gutters in cities from
May to July. The sequence of events in the growth of the fruit body is
given by Atkinson. ^ He finds that the primordium of the fruit body
is oval in form and homogeneous in structure, consisting of intricately
woven hyphse. The hymenophore primordium arises as an internal
Âtkinson, George F.: The Development of Agaricus campestris. Botanical

Gazette, 42: 215-221, September, 1906.
2 Atkinson, George F.: Morphology and Development of Agaricus Rodmani.
Proceedings American Philosophical Society, 191 5: 309-343, with 7 plates.

236 MYCOLOGY
annular zone of new growth toward the upper part of the young fruit
body (basidiocarp) and with its origin the four primary parts of the
basidiocarp, pileus, stem, marginal veil and hymenophore are differen-
tiated. By the continued growth and multiplication of hyphae rich in
protoplasm, which are parallel and directed downward, the hymeno-
phore primordium becomes more compact to form a level palisade
zone, and as the ground tissue beneath lags behind in growth, the more
rapid growth of hymenophore causes a rupture of the ground tissue
beneath and an annular gill cavity arises. The lamellae project into this
cavity, as downward-growing radial sahents of the level palisade zone,
beginning next to the stem and proceeding in a centrifugal direction.
Cultivation of the Mushroom. —
The commercial growing of mush-
rooms has been placed upon a sure financial basis within recent years
and around Philadelphia, notably in Chester County, there are large
concerns which make the culture of mushrooms a specialty. Mush-
room cultivation is an important business in Europe, especially in
France where certain of the grades are canned and bottled for export
trade. Mushrooms are grown in America in long mushroom houses,
or sheds especially constructed and heated for the purposes of the trade.
Cellars are also devoted to the industry. Sometimes they are grown
under the benches of greenhouses devoted to the raising of other plants.
The beds are so constructed of boards that they rise in tiers of four, or
five with a central aisle, or in the larger houses there are tiers of beds
along the walls and in the center of the house with two aisles running
lengthwise with a cross aisle at the far end or in the middle of the house.
Stable manure is used as the compost for commercial mushroom
culture. Bedding straw should also be included with the manure in
the compost. The manure should be the best that can be obtained.
It should be thrown into about four feet high and forked over
piles
occasionally to assist the fermentation process, which is assisted further
by wetting the fermenting mass occasionally until the fermentation is
completed, which is usually at the end of three weeks. During this
time all objectionable odor should be lost and the temperature should
decline to 120° or i30°F. Out of this compost the beds are constructed
by compressing the mass with blows of a spade, or by a compressing
board. Growers cover the manure bed with a thin layer of garden soil
one to one and a half inches deep. This operation is known as casing,
and is performed after the spawning operation has been completed.
Spawning consists in breaking up the bricks of spawn into about ten

pieces and one piece of spawn, which consists of hard manure pene-
trated by the mushroom hyphae, is used for each square foot of bed
space. The piece of spawn should be covered by about one inch of
compost which should have a temperature of 70° to 75°F. The casing
soil should be well moistened by repeated sprinkhng, and not by a sud-
den drenching. Under favorable conditions, such a bed should come

into bearing in from six to eight weeks after spawning, and during the
period of production constant care in the matter of watering is neces-
sary to keep the beds up to the maximum conditions of production.
The making of spawn is an art in itself and the process is fully described
in a recent book by B. M. Duggar on "Mushroom Growing," published
in 191 5 by Orange Judd Company, New York. Duggar also ascer-
tained in his studies of the mushroom that fragments of growing mush-
rooms obtained under aseptic conditions could be made the starting
point for pure cultures of spawn. This is based on the fact, that a
small piece of the inner stipe tissue of a fresh mushroom will, when
placed on any suitable sterile nutrient medium, promptly develop a
mycelium. The method of making pure cultures is described in Bulle-
tin 85, Bureau of Plant Industry, United States Department of Agri-
culture and in Duggar's "Mushroom Growing" and need not be re-
peated here.
Chemistry and Toxicology of Mushrooms. With the increase —
hving and in our population, which is beginning to feel the
in the cost of
shortage of food supplies, earnest attention has been directed to foods,
such as the edible wild fungi, which are frequently abundant during the
summer months. One phase of this study has been the investigation
of the food value ofmushrooms and toadstools. Chemical analyses
have been made to ascertain what they contain. It has been found,
that such a fungus as Polyporus sulphureus, has over 70 per cent, of
water, while species of Agariciis and Coprinus have fully 90 per cent,
of water. As to nitrogen, although the proportion of this element in
the dry matter of different fleshy species varies from 2 to 6 per cent., it
has been found that much of the nitrogen is present in the form of non-
protein substance of a very low food value and some of it enters into
the composition of a substance closely related to cellulose. Thus, not-
withstanding the fact that Coprinus comatns contains 5.79 per cent, of
nitrogen, we find only 0.82 per cent, as available (digestible) proteins,
:
238 MYCOLOGY
SO that the food value of this form is less than had formerly been sup-
posed. The amount
fatty substances soluble in ether are present to the
of 4 to 8 per cent. The carbohydrates (cellulose, glycogen, trehalose,
mannite, glucose, etc.) make up the largest part of the dry matter of
the mushroom. Starch usually present in higher plants is absent in

these fungi. The ash varies greatly, varying from i,o8 to 15 per cent.
with potassium as the most
abundant element. Sulphuric
acid occurs in the ash of all fungi,
with 1.58 per cent, in the ash of

Hehella esculenta.
The poisonous substances are
alkaloids, such as choline,found
in Amanita muscaria, Hehella
esculenta and other fungi, neurin
(deadly), muscarin, the most
dangerous alkaloid found in toad-
stools, as in Amanita muscaria
(Fig. 93). Phallin, a deadly
poison, found in Amanita phal-
loides, is albuminous in nature.
Helvellic acid, a deadly poisonous
substance, occurs in Helvetia es-
culenta, especially in old decaying
specimens. The symptoms of
poisoning with muscarin are long
delayed. They may be summed
up words of Mr. V. K.
in the
Chestnut (Circular No. 13 Divi-
Fig. 95. — Deadly
amanita, Amanita
sion of Botany, United States
phalloides, showing death cup, or volva, at
base of stipe. {From Gager, after E. M. Department of Agriculture)
Kiltredge.)
"Vomiting and diarrhoea almost
always occur, with a pronounced flow of saliva, suppression of the
urine, and various phenomena beginning with giddiness,
cerebral
loss of confidence in one's ability to make ordinary movements, and
derangements of vision. This is succeeded by stupor, cold sweats,
and a very marked weakening of the heart's action. In case of rapid
recovery, the stupor is short and usually marked with mild delirium.
In fatal cases, the stupor continues from one to two or three days,
and death at last ensues from the gradual weakening and final stop-
page of the heart's action." Fortunately an antidote has been found
in the hypodermic injection of atropine in doses of one-hundredth to
one-sixtieth of a grain. Strong emetics should also be used to rid the
stomach of the offending food. The action of phallin from Amanita
phalloides (Fig. 95) for which no antidote is known except the adminis-
tration of emetics and the transfusion of blood into the patient, which
may be of little avail is best summed up in Chestnut's account: "The
fundamental injury is not due, as in the case of muscarin, to a paralysis
of the nerves controlling the action of the heart, but to a direct effect
on the blood corpuscles. These are quickly dissolved by phallin, the
blood serum escaping from the blood-vessels into the alimentary canal,
and the whole system being drained rapidly of its vitality. No bad
taste warns the victim, nor do the preUminary symptoms begin until
nine to fourteen hours after the poisonous mushrooms are eaten. There
is then considerable abdominal pain and there may be cramps in the
legs and other nervous phenomena, such as convulsions and even lock-
jaw, or other kinds of tetanic spasms. The pulse is weak, the abdom-
inal pain is followed rapidly by nausea, vomiting, and extreme diarrhoea,
the intestinal discharges assuming the rice-water condition characteristic
of cholera. The latter symptoms are maintained persistently, generally
without loss of consciousness, until death ensues, which happens in
from two to four days."
B. Gasteromycetes. —-The fungi known as the Gasteromycetes
{yaarrip = belly, sac -f /jlvktjs = fungus) have the basidial layers, or
hymenium, enclosed within a peridium, as in the common puff-ball.
The shell or hull enclosing the masses of spores is called the peridium,
which is a simple uniform layer in some genera (Scleroderma), or it con-
sists of two distinct layers, the exoperidium and the endoperidium.
The earth-star (Geaster) has a thick outer peridium, which splits in a

stellate manner, later becoming reflexed. The exoperidium in such
genera as Bovista and Lycoperdon is a loose pliable coat often having
spines and warts. Many of the genera are stalkless, but other genera,
such as Tylostoma, are stalked. Inside of an unripe puff-ball, we find a
white fleshy mass of soft cellular matter, the gleba. As the fruit
bodies grow they become chambered. The chambers, in countless
numbers, are narrow, irregularly curved and branched, separated from
240 MYCOLOGY
each other by curved plates of tissue which anastomose in every direc-

tion. The walls of the chambers consist of layers of branched hyphse
bearing the basidia which line the interior walls of the cavities and con-
stitute the hymenium. Each basidium usually bears four spores.
The way the spores are borne on the basidia is characteristic. They are
almost sessile in Geaster, in Bovista they are found on long sterigmata.
Mitremyces may have as many as a dozen basidiospores, which are
sessile and lateral.
When and ripens, the tissues become
the puff-ball reaches full size
moist, deliquesceand change in color. The tissues are absorbed and
disappear and the whole mass dries up, leaving the interior sur-
rounded by the peridium filled with a dry dusty mass usually con-
sisting of slender threads (the capillitium) and countless multi-
tudes of ripe spores. The threads of the capillitium are absent in many
genera, but when present they are characteristic and used as important
points in the classification. There are two distinct kinds of capillitial
threads. In one kind, the threads are long hair-like strands, simple,
more or less branched and interwoven, proceeding from the inner walls
of the peridium, or from the centrally placed columella. The second
type, characteristic of Bovista, Bovlstella and Mycenastrum, has rela-
tively short and branched threads entirely separate and distinct from
each other and are not connected with the peridium nor the columella.
The bird-nest fungi are characterized by the thickening of the walls of
the glebal chambers to form separate little seed-hke bodies enclosing the
spores. These are known as peridioles. The ripe spores in some are
smooth, some are spinulose, while in shape they are globose, oblong or
oval.
The most primitive forms of these fungi are probably the subter-
ranean forms included in the family Hymenogastrace^. In one
Gasteromycetes, the division of the famihes is
classification of the
based on whether the sporophore is borne above or below the ground.
The family Hymenogastrace^ with subterranean fruit bodies belongs
to one division, all of the other families to the other division.
Family i. Hymenogastrace^. —The subterranean fruit bodies of
these fungi suggest those of the families Terfeziace^ and Tuberace^
among the ASCOMYCETALES, but the spores of the two latter
families are borne in asci,and are known as ascospores, while those of
the former family are borne on basidia and are known as basidiospores.
1
Most of the forms are irregularly globose and grow under trees, some-
times their association with certain kinds of trees suggesting a para-
sitic attachment. They are often found in sandy places, where they
are exposed frequently by rain erosion. The mycelium of these fungi
is filamentous, or cord-like. The gleba is richly chambered and the
walls of the glebal chambers are lined with the hymenium. Cystidia
are often found between the basidia. The fruit bodies are variously
shaped. In Lycogalopsis, they are hemispheric; in Phyllogaster, pear-
shaped; in Cauloglossum, club-shaped; some are stalked and suggest
the shape of the Agaricace^.
Very few of the forms are known commonly, and of the dozen Cali-
fornian species, many are known imperfectly by a single collection.
and Sclerogaster have each a single species in California;
Gaiitieria
Hymenogaster and Octaviana are represented by two Californian species,
while Hysterangium and Melanogaster have three species in California.
Two species of Rhizopogon and one found in South
of Melanogaster are
Carolina. The climate probably has something to do with this
distribution.
Family 2. Tylostomace^. At first, the fruit body is subter-
ranean, later as in Tylostoma niammosa, a form found in heathland, it is
raisedon a stalk not prolonged as an axis. The peridium is double, the

outer one falUng off at maturity, the inner one is thin. The uncham-
bered gleba possesses well-developed capilUtial threads, which are con-
nected with the inner wall of the endoperidium. The basidia in
Tylostoma are unicellular, club-shaped and bear four laterally placed
spores, one above the other on well-developed sterigmata, thus differ-
ing from the other two basidiomycetous fungi.
—
Family 3. Lycoperdace^. -The fruit body from the beginning is
epigaeic. Its gleba is chambered richly and the inner walls of each
chamber are lined with a hymenium. The peridium is differentiated
into an outer and an inner peridium. The gleba, when ripe, breaks
down into powdery spores and richly branched capilHtial threads. This
family contains some of our most delicious and important food species,
if they are taken before fully mature. The genus Ly coperdon, in which
the true peridium opens by an apical mouth, includes over one hundred
species, which in America can be divided into the purple-spored series,
and the olive-spored series. Lycoperdon atropurpureum is found in sandy
pastures, woods and bushy places commonly in the months from August
16
242 MYCOLOGY
to October. It is an extremely variable species, Lycoperdon gemma-

turn, an olive-spored species, has a turbinate shape, its outer peridium
being marked with long, thick, erect spines, or warts of irregular shape
with intervening smaller ones, whitish, or gray in color. The larger
spines fall away first imparting to the surface of the peridium a reticu-
late appearance. It often grows cespitosely on the ground, or rotten
tree trunks in woodlands. Lycoperdon pyriforme is another common
species found in woods and clearings on the ground, or on decaying
wood. It is edible, tender and of second-class flavor when young.
Fig. 96. — Fruit-body of Calvatia cyalhiformis. {Photo, by IF. H. Wahnsley.)
The largest puff-balls are included in the genus Calvatia (Fig. 96),
which differs from Lycoperdon in the absence of an apical mouth and
a regular dehiscence. The fruit bodies are globose, or top-shaped, aris-
ing on the surface of the ground from subterranean, cord-like hyphae.
Calvatia cyathiformis (Fig. 96) which is edible, if eaten when white in-
side, grows in open grassy fields and lawns and reaches a diameter of
three to six inches. Calvatia gigantea, the giant puff-ball, grows in
pastures and meadows. Usually the fruit bodies are ten to twenty
inches in diameter and even larger. The genus Bovista has a fragile
exoperidium, and in the absence of a sterile base and the fact that the
fruit body separates easily from the place of attachment it is distin-
guished from Lycoperdon. Because they are readily detached and
Fig. 97. — Specimen of Geaster fornicatus from Carleton Rea, England. {After Lloyd,
J. U., and C. G., Bull. 5, Lloyd Library, June, 1902, Mycological Series No. 2.)
readilyblown about, they are called "tumblers." Catastoma has an

outer peridium which splits by a circular Hne of cleavage, so that the
upper part is dislodged carrying along with it the inner peridium which
244 MYCOLOGY
opens by a mouth that is situated at the actual base of the plant as it
grows. The lower part remains as a saucer-shaped body in the soil.

A capillitium is present. Catastoma circumscissum is the common species.
The earth stars are included in the genus Geaster, where the peridium
consists of three persistent coats, the two outer adhere and split into
leathery, stellate divisions exposing the parchment-like inner peridium,

which opens by an apical pore (Fig. 97). It has a columella. The spores
are dark brown and mixed with the simple capillitial threads. Geaster
hygrometricus is the common species. It grows in sandy soil and in dry
weather its segments are strongly recurved, but in wet weather they
expand, hence the plant is sometimes dubbed poor man's weather
glass. Astr(Eus, which resembles Geaster, is distinguished by the
absence of a columella and by the long capilUtial threads which are
much branched and interwoven.
—
Family 4. Nidulariace^. The following account of the family
of bird's-nest fungi is taken from Bulletin 175, United States Depart-
ment of Agriculture, on "Mushrooms and Other Common Fungi" by
Flora W. Patterson and Vera K. Charles. Dried material of these
fungi might be kept for use by the class in the systematic study of the
higher fungi with the following key at hand. The types should be
used as unknowns.
Members of the family Nidulariace^
are represented by small,
leathery, cup-shaped plants growing on old sacking, manure, earth, and
decaying or dried wood. The common name is suggested by the form
of the peridium, which is cup-shaped and contains many small, lenticu-
lar bodies (peridiola) resembhng eggs. The mouth of the peridium is at
first covered by a membrane (epiphragm), which later becomes ruptured
and exposes the peridioles. In Cyathiis and Crucibulum, the peridi-

oles are attached to the inner wall of the peridium by elastic cords
called funiculi. The spore-bearing tissue and spores are never resolved
into a dusty mass, as in many Gasteromycetes, but persist in the
form of peridiola which contain the spores, which are hyaline and
ellipsoidal to subglobose.
Key to Nidulariace^
Peridium with several to many sporangioles:

Peridium torn at the apex in opening—
Sporangioles not attached to the inner wall of the peridium. Nidiilaria.
— —
Peridium opening by a deciduous membrane

Sporangioles attached to the inner wall of the peridium
Peridium of three united layers and spores mixed with
filaments Cyalhua.
Peridium of a single layer and spores not mixed with
filaments Crudbnlmn.
Cyathus
In Cyalhiis the peridium is cup-like and composed of three layers.

The apex is covered by a white membrane, which bursts, disclosing egg-
like bodies, the peridiola, which usually fill about one-half of the cup.
The peridiola are attached to the inner wall of the peridium by an elastic
cord, which is attached to each peridiolum in a depression on one side.
Cyathus stercoreus
Peridium cylindrical, campanulate to infundibuliform, sessile or with an elon-

gated base, light brownish, at first with shaggy, matted hairs which disappear in age,
interiorsmooth and nonstriate; peridiola black.

Cyathus skrcoreusis an exceedingly common species and is to be found growing
on manure or in heavily manured places. It is subject to considerable variation in

size and form.
Cyathus striatus
Peridium obconic, exterior even, brownish, hairy, interior striate, lead-colored;

apex truncate, covered by a white membrane, which is at first strigose; peridiola
compressed, subcircular.
Plant one-half to three-fourths inch in height and about three-eighths inch in
diameter.
Cyathus vernicosus
Peridium bell-shaped, subsessile, base narrow, broadly open above, exterior at

firstbrownish, silky tomentose, becoming smooth, interior dull lead color, smooth.
Differs from Cyathus striatus in the even, non-fluted inner surface of the peridium and
in the larger peridiola.
Plant about one-half inch in height and about three-eighths inch in diameter.
Crucibulum
In Crucibulum the peridium is cup-shaped and consists of one thick

fibrous layer, Hned by a very thin, smooth, and shining layer. The
246 MYCOLOGY
mouth when young is covered with a yellowish tomentose membrane,
the peridiola are more numerous than in the preceding genus, and each is
Attached to the peridium by an elastic cord which springs from a pro-
jection on the peridiolum. The plants are smaller than in the genus
Cyathus.
Crucibulum vidgare
Peridium yellowish-brown, becoming paler with age, outer surface when young
velvety tomentose, inner surface smooth and shining; mouth at first closed by a yel-
lowish membrane, which ruptures and exposes the peridiola. Peridiola biconcave,
with a projection on one side from which originates the elastic cord which attaches
the peridiola to the peridium.
Plant about one-fourth inch in height and about the same in diameter.
Family 5. Sclerodermace^. —The fruit bodies of the fungi in-
cluded in this family are subterranean, or epigeic, globose, sessile, or

occasionally with a root-like stalk. The peridium is generally simple,
thick, rough, warty, or scaly, opening irregularly at maturity. The
gleba consists of rounded basidia-bearing parts, which are separated by
sterile veins or strands of hyphge. The individual basidia are pear-
shaped to club-shaped with spores which are often lateral in position.
The capilhtiiim is rudimentary. Scleroderma is the most common genus
with sessile fruit bodies and thick, hard, leathery peridium, frequently
warty. apex into stellate lobes. Scleroderma
It usually bursts at the
geaster grows in sandy woods, banks or along roadsides. S. vidgare
is common in dry situations, or hard ground, along cinder paths and
gravel walks.
Family 6. —
Sph^robolace^. The fruit body is on the surface of
the ground. The periphery of the gleba is furnished with a palisade-
like layer of radially arranged turgescent cells. The basidia-bearing
portion of the gleba is penetrated by sterile veins, or hyphal strands.
When ripe the gelatinous gleba is forcibly ejected from the fruit body by
the inversion of the pahsade-like layer. The family includes a single
genus, Sphcerobolus, of five species. The best-known species is S. car-
pobolus of cosmopolitan distribution.
C. Phallomycetes. —The carrion fungi, stink-horn fungi, or dead-
men's fingers, resembles the button stage of the Amanitas, and the puff-
balls when still young, but later the outer wall is ruptured and the stem
elongates carrying upward the sporogenous tissue as a terminal cap, or
enlargement. The subterranean mycehum is cord-hke and from it the
fruit body arises which has a pcridium of two or three layers. The
outer peridium is leathery and tough, while the inner peridium is gelat-
inous at maturity. The outer peridium remains at the base, as a
cup called the volva. The sporophore, pileus, or cap, is raised up on the
end of a stalk, or stipe, which is usually spongy in character. The
sporophore takes a variety of forms, but in all cases, its outer surface at
y^'"-
Fig. 98. -Clathrus cancellatus, fully mature fruit-body, natural size. {After Ed.
Fischer, Die naliirlichen Pflanzenfamilien I. lA**, p. 282.)
first represents the hymenium which deliquesces at maturity, so that the

minute spores are imbedded in a greenish, fetid sUme, which gives off a
penetrating, nauseating odor, attractive to blue-bottle flies, that lick
offthe malodorous slime with evident enjoyment and are the agents
by which the spores are distributed. In fact, it has been proved that
the basidiospores germinate better after passage through the alimentary
.
248 MYCOLOGY
canals of flies. The gleba is the fruiting portion of the phalloid and its
bulk appears considerable in the early egg-shaped stage of the fruit

body. As the carrion fungus matures, it forms proportionately less of
the fruit body, for it is converted into the greenish, mucilaginous mass
which is removed by the flies. Some forms like Didyophora have a veil
that hangs under the pileus and spreads out as a net around the stem.
Although it is called the veil, it is more correctly the indusium. The
sporophore in genera like Clathrus (Fig. 98) takes the form of a hollow
sphere, or of a basket-like lattice, while in other genera it resembles the
open iron framework of a lantern, a brazier, a crinoid, or storie-lily, an
octopus, or even a sea-anemone. One tropic form of Brazil has been
called Pilzblumen by the Germans. The species are not common in
temperate regions, but in the tropics they are richer in forms and more
abundant; for example, in Florida the species of Clathrus are common,
the writer finding four specimens within a quarter of a mile along a road
across the sand dunes at Ormond.
—
Development of the Carrion Fungi. Several authors have studied
the development of several forms of the Phallomycetes, notably
Burt and Atkinson. Burt^ has contributed three papers dealing with
the genera Anthurus, Clathrus and Mutinus, while Atkinson's studies^
are concerned with Ithyphallus and Didyophora.
Burt finds in the Clathrace^ that the egg consists of cortical and
medullary systems continued upward from the mycelial strand in the
earliest stage. The cortical layer gives rise to the outer layer of the
volva, the cortical plates and the pseudoparenchyma of the receptacu-
lum. The medullary portion gives rise to the gelatinous masses of the
gelatinous layer of the volva, to the gleba, and to the gelatinous tissue
of the chambers of the receptaculum. The elongation of the receptacle
in Clathrus columnatus (Fig. 98) begins at the baseand after its elonga-
tion the gleba hangs suspended from the arch of the receptaculum by
medullary tissue constituting the chamber masses of the receptacle.
In the earliest recognizable stage of Mutinus caninus, the egg con-
sists of the cortical and medullary tissues of the mycelial strand,
1 Edward A.: A North American Anthurus: Its Structure and Develop-
Burt,
ment. Memoirs Boston Soc. of Nat. Hist., 3: 487 (1894); The Development of
Mutinus caninus. Annals of Botany, 10: 343 (1896); The Phalloideas of the United
States. Development of the Receptacle of Clathrus columnaLus.
Atkinson, George F.: The Origin and Taxonomic Value of the Veil in Dicty-
ophora and Ithyphallus. Botanical Gazette, 50: 1-20, January, 1911.
continued directly upward from the strand. Of these tissues, the

medullary bundle spreads out at itsupper end and forms a dense
sheaf-like head by repeated branching and anastomosing. The
cortical layer of tissue becomes the outer wall of the volva; the sheaf-
like head gradually differentiates into all the other parts of the older
egg. In such differentiation the central
column first appears. The formation of
the gelatinous layer of the volva now begins
in the periphery of the head. A dense
dome-shaped mass Along the inner
arises.
surface of the dense zone and next to the in-

termediate tissue, the rudiment of the gleba
arises from the clustered swollen ends of
lateral branches of the tramal tissue. These
hyphal ends take position in a palisade
layer facing the intermediate tissues and by
the crowding in of new hyphal ends (basidia)
the surface of this layer becomes greatly
enlarged and thrown into folds and torn
from the intermediate tissue. The rudiment
of the stipe arises in the intermediate tissue
lying next to the central column by the forma-
tion of deeply staining tissue rich in proto-
plasm. Somewhat later, masses of tissue in
the dense and intricately interwoven rudi-
ment of the stipe show a tendency toward
gelatinization. These masses mark the
position of the later chamber-cavities in the
wall. Toward the upper end of the stipe,
such masses are in contact with the central
column, and they mark the position of the —
FiG. 99. Mature stink-
horn, Diclyophora duplicata.
pits which open into the main central cavity {Photo by W. II. Walmsley.)
of the stipe inmature stages of M. can in us.
The chamber thrown into folds through a more rapid growth
walls are
of the pseudoparenchyma than that of other parts of the egg. Final
elongation of the stipe and elevation of the gleba is brought about
through the straightening out of the folds in the chamber walls.
The studies of Atkinson deal with the origin of the veil of Dictyo-
—
2 50 MYCOLOGY
phoni (Figs, gg and loo), and llhyphaUiis. From such studies, he

confirms the making of two genera out of them. His plates show
Fig. 100. Diclyophora phallaidea. Fully developed fruit-body with veil 2/3 natural
size. {After Alf Moller in Die natUrlichen pflanzenfamilien I. lA **, p. 294.)
that three common forms were examined, viz., Ilhy phallus impudicus,
Diclyophora duplicata and the Phallus Ravenellii.
Two families are distinguished: Clathrace^ and Phallace^
which may be distinguished as follows:
"
Receptacle latticed or irregularly branched, sessile or stalked;

gleba inclosed within the receptacle. Family i. Clathrace^.
Receptacle tubular or cyhndric, capitate, with the gleba external.
Family 2. Piiallace.e.
Fig. ioi. — A, B, Dictyophora phalloidea. A, Longitudinal section of a fruit-body

fully stretched beyond volva (natural size); B, longitudinal section of a young fruit-
body (twice enlarged); G, volva mucilage; a, gleba; H, cap; /, indusium; Sw, stipe;
Pi, primordial layer between cap and indusium'; Pi, primordial layer between in-
dusium and stipe; S, S, tissue of stem; B, tissue of base of fruit-body. {After Ed.
Fischer in Die naliirlichen PJlanzejifamilien I. lA**, p. 295.)
The first family, according to "Die natiirlichen Pflanzenfamilien,

comprises eleven genera of which Clathrtis (Fig. 98), Sinihlum, An-
thurus are North American. Three species of Clathrus have been col-
lected in this country. Simhlum rubescens was collected originally on
252 MYCOLOGY
Long Island and later in Nebraska, while Anthurus horcalis has been
found in New York, Massachusetts and Pennsylvania.
The family Phallace^ is represented in the eastern United States
by three important and interesting genera, viz., Mutinus, I thy phallus,
Didyophora (Figs. 99, 100, loi). Mutinus is the simplest form with
the gleba Ijorne on the upper portion of the stipe without the hanging
cap. Mutinus caninus has a hollow, perforate stipe reddish in color
bearing the greenish bad-smelling spore slime over its upper end.
Ithyphallus impudicus, our commonest species, has a globose volva,
cylindric, hollow spongy stalk bearing a campanulate pileus, the spore-
bearing surface being reticulate pitted. Didyophora duplicata, which
resembles the Brazihan Pilzblumen, D. phalloidea in (Figs. 100, loi)
the possession of a long white indusium, which hangs down beneath the
cap like a spread-out hoopskirt. The terminal cap is campanulate
and after the removal of the malodorous greenish spore slime appears
reticulate-pitted. The volva is prominent.
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Farlow, William G.: Bibliographical Index of North American Fungi, vol. i,
part I, Abrothallus to Badhamia, Publ. 8, Carnegie Institution of Washington,
1905-
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2 54 MYCOLOGY
Ferguson, Margaret C: A Preliminary Study of the Germination of the Spores

of Agaricus campestris and other Basidiomycetous Fungi. Bull. i6, Bureau of
Plant Industry, U. S. Dept. Agric, 1902.
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23: 515-534, 1909-
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der Pilze Abhandl. d Senkenb. Naturf. Ges., v: 204, 1864.
Fischer, Ed.: Versuch eine systematischen Uebersicht Uber die bisher bekanntem
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Fischer, •
Ed.: Untersuchungen zur vergleichenden Entwickelungsgeschichte
und Systematik der Phalloideen. Denkschr. Schw. naturj. Gesellsch., 32:
1-103, pi. 1-6, 1890.
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1-51, pis. 1-3, 1893.

Fischer. Ed.: Phallinefe, Plectobasidiineae. Die natiirlichen Pflanzenfamilien,
I Teil, Abt. i**, 1900.
Gerard, W. R. List of United States Phalloidei. Bull. Torr. Bot. Club, 7:11, 1880.
:
Harper, R. A. Binucleated Cells in Certain Hymenomycetes. Botanical Gazette,

:
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20,
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:
and 20 figures, 1906: 1-32.

Lloyd, C. G.: The Phalloids of Australasia, 1907: 1-24.
Lloyd, C. G.: Concerning the Phalloids. Mycological Notes, No. 26, 325-337,
pis. 112-121, figs. 160-163, 1907.
Lloyd, C. G.: Concerning the Phalloids. Mycological Notes, No. 24, December,
1906; No. 26, May, 1907; No. 28, October, 1907; No. 29, January, 1908; No.
30, February, 1908.
Lloyd, C. G.: Polystictus (Section Pelloporus). Mj'cological Notes, Polyporoid
Issue, No. I, February, 1908.
Lloyd, C. G.: The Genus Favolus. Mycological Notes, Polyporoid Issue, No. 2,
August, 1909.
Lloyd, C. G.: Synopsis of the Genus Hexagona, 1910: 1-46.
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schaftliche Rundschau, xxi:
Marshall, Nina L.: The Mushroom Book, New York, Doubleday, Page & Co.,
1902: i-xxvi + 1-167.
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Fungi, Amer. Journ. Physiol., i: 225-238, 1898.
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den Tropen, 1895; i~"i52, with 8 plates.
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vol. 9, parts I & 2, conclusio.
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vol. 10, part I, July 28, 1914.
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256 MYCOLOGY
Central America, southern Florida, the West Indies, and other islands
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25, 1916.
CHAPTER XXII
FUNGI IMPERFECTI (DEUTEROMYCETES)
The life histories of the fungi belonging to this group are imperfectly
known, and hence, it happens that when it has been established, the
type is removed from the fungi imperfecti and properly classified with
some other group. The name Deuteromycetes, also applied to the
imperfect fungi, is derived from the Greek, devrepos = second. Many
important parasites are included here, and hence, it has been considered
important by mycologists to give the characters by which the fungi
imperfecti are distinguished.
—
General Characters. The mycelium consists of septate, hyaline, or
pigmented hyphas, or only of chain of yeast-like cells. The hyphas are
diffuse, or plectenchymatous (irXeKTos = woven). Stromata are fre-
quently present. The fructification is a single conidiophore, a layer
of conidiophores, or a conidial fructification (pycnidium). The
Fungi Imperfecti represent the accessory fruit forms of the ASCOMY-
CETALES, rarely those of other orders. The mycelium is practically
the same as found in the sac fungi. The septate hyphae spread over
the substratum, or penetrate its interior, and the fungi live sapro-
phytically, or parasitically. The arrangement of the hyphas in various
ways has suggested the segregation of species and genera. The under
layer (subiculum = felted stratum of hyphae) is of loose, entangled
threads, or disc-like bodies, or radially stretching fibrils, aggregated
loosely. The stroma on the contrary represents compact tissue, cor-
responding to similarly named structures in the ASCOMYCETALES.
The on the stroma.
fruit layer originates in or
Reproduction dependent on exogenously produced spores, known
is
as conidiospores. In the simplest cases, the mycelium gives rise at

indefinite places to outgrowths, which are separated as spores. There
arise from the mycelium, erect conidiophores which form conidiospores
in the different species. With an unbranched conidiophore, the conidio-
spores arise at apex followed by a second, a third, etc. When the
its
end of the conidiophore is globutar, the spores arise on the ends of

sterigma. By the branching of the conidiophores originate conidial
258
—
rUNGI IMPERFECTI (dEUTEROMYCETES) 259
Fig. 102. Phylloslicta pavice on horse-chestnut leaves. (Cold Spring Harbor, L.I.,
July 28. 1915-)
26o MYCOLOGY
strands, which suggest the inflorescences of flowering plants. One can

separate these into monopodial, or sympodial forms. A bundle of coni-
diophores is known as a coremium {Kopr^na = broom). If the conidio-
phores are arranged side by side, they form a conidial layer, which
ariseson the upper surface of a stroma. Such a conidial layer may be
folded, orit may be chambered, the irregular chambered spaces being
lined with the conidial layer. Finally, the conidial layer may be in-
closed in receptacles called pycnidium, which correspond to those of
the Pyrenomycetiine^. The conidiospores are of different sizes,
hence one can distinguish them as a micropycnidia and as macro-
pycnidia, and the spores as micro- and macropycnospores. Stylospores
are those spores borne on a filament {oTxiKos = a column). This term
is also superfluous. The number of fungi imperfecti surpasses the
ASCOMYCETALES.
Systematic Position. — Fuckel includes all those fungous forms
as fungi imperfecti which have no final fruit forms, such as asci
and basidia. The name Deuteromycetes of Saccardo is less fortunate
than that of Fuckel. That many fungi imperfecti represent accessory
fruit forms of ASCOMYCETALES is known, so that the group is not
a permanent systematic entity. It is a motley assemblage of hetero-
geneous forms. As with the large group, so it is with the genera.
Some of the genera inclose not always related forms, that is of the same
phylogenetic series. Schroeter calls such genera Formgattungen (
=
form genera). In the following classification of them, this point of
view must be kept prominently in view, for a natural classification of
Fungi Imperfecti is in the nature of things an impossibility. The great-
est number are saprophytes, useful in the destruction of dead plant
parts. Many are parasites and produce dangerous diseases in culti-
vated plants.
A. Conidia in pycnidia, or chamber-like hollows. I. SPH^ROPSI-

DALES.
B. Conidia in conidial layer formed ultimately wholly free. II.
MELANCONIALES.
C. Conidia on conidiophores. Single or in coremia. III. HYPHO-
MYCETALES.
I. SPH^ROPSIDALES.— The conidia are formed in pycnidia.
The receptacles are closed or open by a pore, or by a slit suggesting
FUNGI IMPERrECTI (dEUTEROMYCETES^ ?6l
groups of ASCOMYCETALES. Four families are included in this

order, and these number of important
families include a considerable
genera of fungi, which specifically are the cause of important plant
diseases. PhyUosticta is a genus, the species of which are confined to
leaves, and they produce characteristic leaf spots on a great variety of
plants. The specific name of the fungus is usually derived from that
of the host plant attacked, as for example, PhyUosticta catalpce, which
Fig. 103. — Six j3en Davis apples showing apple blotch (Phyllostica solilaria).
{After Scott, W. M., and Rorer, J. B., Bull. 144, U. S. Bureau of PlantJndustry,
March 6, 1909.)
grows on the leaves of the catalpa. The group has been monographed
systematically by J. B. Ellis. The spores are small, egg-shaped or
elongated, unseptate and in color pale green, or hyaline, produced in
pycnidia. The most important species of this genus are PhyUosticta
ampelopsidis on the Virginia creeper {A mpelopsis) ; on catalpa
catalpcB
leaves; labruscce on the leaves of the grape; pavi<^ on horse chest-
nut leaves (Fig. 102); PhyUosticta solitaria E. and E. (Figs. 103 and
104) is the cause of apple blotch, and violce on violets. The conidio-
262 MYCOLOGY
spores in Phoma are colorless and unicellular. The pycnidia are

black with a terminal pore and depressed in the tissues of the host.
The genus is arbitrarily limited to those species in which the spores
are less than 15^, for the larger spored forms have been placed in the
genus Macrophoma. The most important species from the pathologic
viewpoint are out of the iioo species recognized the following: Phoma
betcB is the cause of the heart rot and blight of beets. Phoma batata
produces a dry rot of sweet potato; while Phoma solani behaves much
nrn o
Fig. 104. — Microscopic characters of apple blotch fungus {Phyllosticta solilaria).
I, vertical section of pycnidium showing pycnospores; 2, 3, 4, 5, mature pycnospores;
6, 7, 8, germinating spores; 9, mycelium. {After Scott, WM.. and Rorer, J. B., Bull.
144, U. S. Bureau of Plant Industry, pi. Hi, March 16, 1909.)
like the damping-o£f fungus, attacking seedling egg plants near the sur-
face of the ground. The most destructive fungus of the genus Sphcerop-
sis is S. malorum which causes the decay of apples, quinces and pears
and attacks the stem of the apple tree producing characteristic cankers.
The genus includes about 180 species. The 150 species of the genus
C oniothyrium are widely spread geographically. The blight of rasp-
berry canes due to Coniothyrium Fuckelii, which has only recently
is
come into prominence in the United States. The genus Septoria in-
FUNGI IMPERFECTI (dEUTEROMYCETES) 263
Fig. 105. — Septoria leaf spot disease of celery, or celery blight. (After Coons, G. N.,
and Levin, Ezra, Spec. Bull. 77, Mich. Agric. Coll. Exper. Stat., March. 1916.
SPORES^
Fig. 106. — Section through leaf spot of celery blight (Septoria) î i-lis)
and
in leaf tissue and pycnidium with exuding pycnospores. (After Coons, G. 11.,
Levin, Ezra, Spec. Bull. 77, Mich. Agric. Coll. Exper. Slat., March, 1916.)
264 MYCOLOGY
eludes the fungi which c§iuse the leaf spot of the pear, Septoria pyricola,
the late blight of the celery S. petroselini (Figs. 105 and 106), the leaf
blight of thetomato S. lycopersici and the leaf spot of currants, S. rihis.
The pycnidia in this genus develop under the epidermis of the host
producing leaf spots. The center of the leaf spot is occupied by the pore
of the spheric, black pycnidium. Leptothyrium pomi is an imperfect
fungus responsible for the sooty blotch of the apple and other
plants. According to Floyd the same fungus causes the fly speck of
apples. The genus Entomosporium is a small one with closed half-
spheric, black pycnidia. The spores suggest an insect in being four-
celled, the cells being arranged cross-like with attenuated extremities
and swollen Entomosporium maculatum is the cause of the leaf
bases.
blight of the pear and quince.
II. MELANCONIALES.^ —
The mycelium is formed in the interior
of the host plants. The fruit is in the form of a conidial hymenium,
which is produced below the epidermis of the host, breaking through
clefts in the surface of the host as bright or black spots. The conidio-
phores stand closely together and are simple, or rarely branched, hya-
line, or rarely dark-colored. Pycnidia are unknown in this group of
imperfect fungi. The spores are of different shapes, single or in chains.
The order includes both parasites and saprophytes. The pustule, or
acervulus, which produces spores in Gleosporium may be extensive
The short conidiophore arise from or are inclosed within a cushion,
or stroma, of fungous tissue. The rupture of the epidermis of the host
is accomplished by the opening of the stroma. The ovoidal, fusiform,
slightly curved hyahne spores are discharged with the opening of the
stroma. Some species of Gloeosporium are connected with other genera,
viz., Glomerella (rufomaculans) Gnomonia and Pseud opeziza the im-
,
perfect stages of which were placed as species under the form genus
Glososporium, which is the important form pathologically speaking.
As examples form genus Gloeosporium, we have G. ampelophagum

of the
which causes the anthracnose of the grape; G. venetum which is re-

sponsible for the anthracnose of blackberry and raspberry, while other
species attack the linden, walnut, pine and Norway maple.
In Colletotrichum (Fig. 107) the conidial cushions have a bristly
border, while the conidiospores are in chains. Colletotrichum Lindemuth-
ianum causes anthracnose of bean, an important disease in gardens and
fields (Fig. 107). The cotton is attacked by C. gossypii, citrus fruits by
FUNGI IMPERFECTI (dEUTEROMYCETEs) !65
4 \
M
i
^k
Fig. 107. —Anthracnose cankers on bean pods (Colletolrichum Lindemidhianum)
(After Whetzel, H. H., Bull. 255, Cornell Agric. Exper. Stat.)
:
266 MYCOLOGY
C. glceosporioides, clovers andby C. trijolii and the snapdragon

alfalfa
by C. on these plants induced by species
antirrhini. Usually the diseases
of Colletotrichum are known as anthracnose (Fig. 107). Coryneum Bei-
jerinckii is a destructive fungus causing the peach blight. Pestalozzia
Guepini var. vaccinii is a fungus often found upon the cranberry leaves
and fruits. The conidiospores are three-celled, the terminal cells with
filiform appendages. The shot-hole disease of plum and cherry is due to
Cylindrosporium padi. The formation of the acervuli is followed by the
falling out of the disease areas of the leaf resulting in the formation
of the characteristic shot-hole.
The fruit spot of apples is caused
by C. pomii.
III. HYPHOMYCETALES.
—The hyphae are septate,
branched in or on the substra-
tum. They are dark, or hyaline,
separate, or bound into coremia,
or layer-like cushions. The con-
idiospores may exist as oidio-
spores through the separation of
the hypha. The conidiophores
are simple, or branched. The
conidiospores of different shapes
and colors are borne in a variety
Fig., 108. — Sweet-potato stem rot of ways on the conidiophores or
{Fusarium balatalis). Section through
sweet potato showing blackened ring just their branches. The genera may
below surface caused by the stem-rot fun-
be arranged in three series.
gus. {After Harter, L. L., U. S. Farmers'
Bull. 714, March ii, 1916.) MyceHum and spores light-
A.
colored: Oospora, Monilia,
Oidium, Sporotrichum, Botrytis, Cephalothecium, Ramularia, Cercos-
porella, Piricularia. B Mycelium
. dark-colored at least with age spores ;
generally dark: Fusicladium, Polythrincium, Scoletotrichum, Clado-

sporium, Helminthosporium, Macrosporium, Alternaria, Cercospora,
C. Conidiophores in the form of a tuberculate mass, or sporodochium
Fusarmm. As examples of common disease producing forms
Volutella,
of the above genera without enumerating all of the more important
species may be mentioned the potato scab fungus, Actinomyces chro-
mogenes, the early blight of potato fungus, Macrosporiums olani; the
FUNGI IMPERrECTI (dEUTEROMYCETES) 267
fungus which causes leaf spot of beets, Cercospora beticola. The form
genus Fusarium (Fig. 109), established by Link in 1809, is one which
has come into prominence recently as associated with the production
of serious plant diseases. At least eleven species are found on the
sweet potato (Fig. 108), and these have been investigated by H. W.
Wollenweber^ and other mycologists. He finds that the genus has a
number of vegetative and spore stages the variabihty of which has
caused confusion, as transfers of mycehum produce a growth quite
different in general appearance from that derived from spores from the
Fig. 109. —
Spores of two stem-rot organisms. A, Fusarium bataiatis and B.
F. hyperoxysporum, X500. {After Harter, L. L., U. S. Farmers" Bull. 714, March 11,
1916.)
same medium under conditions otherwise identic. 'yVollenweber and

AppeP have pubUshed a monograph of Fusarium and later Wollen-
weber has studied the Fusarium problem and similar studies should
be made of each one of the form genera of the Fungi Imperfecti.
The genus Fusarium is divisible into sections not only by physiologic
characters (pathogenicity) but also by morphologic characters (coni-
diospores, chlamydospores). The section, Elegans, comprises the
vascular parasitic Fusaria, which are serious enemies of plants, causing
1 WoLLENWEBER, H. W. Identification
: of Species of Fusarium occurring on
the Sweet Potato, Ipomaa hatatis. Journal of Agricultural Research, II: 251-286
July 15, 1914.
Âppel, Otto, and Wollenwebek, H. W.: Grundlage einer Monograph
der Gattung Fusarium Link Arb. Biol. Anst. f. Land. u. Forst., Bd. 8, Heft, i,
pp. 1-207; Phytopathology III: 24-50.
2 68 MYCOLOGY
—
Fig. 1 10. Violet leaf spot {Fusarium viola), i, Germination of microconidio-
sporeS; 2, formation of microconidiospores in hanging drop cultures; 3, germination
of macroconidia; 4, various forms of macroconidia. (.After Mycologia, 2: 19-21, pi.
xviii, January, 191 o).
FUNGI IMPERFECTI (dEUTEROMYCETES) 269
especially wilt diseases. Fusarium oxysporum and T. trichorthecoides

can produce both tuber rot and wilt of the potato plant. Fusarium
viola causes violet leaf spot (Fig. no). Ftisarium bat at atis is respon-
sible for sweet potato stem rot (Figs. 108 and 109).
The sterile fungus Rhizoctonia, represented in America by two para-
Rhizoctonia solani, which is found on at least 165 different
sitic species
hosts,and R. crocorum with a limited distribution on alfalfa and potato

tubers has through the discoveries of Rolfs and Burt been connected
with a basidiomycetous fungus, Corticium vagum var. solani.^
1 Peltier, George L. : Parasitic Rhizoctonias in America. Bull, 189, Agri.

Exper. Stat. University of Illinois, June, 1916.
PART II
GENERAL PLANT PATHOLOGY

CHAPTER XXIII
GENERAL CONSIDERATION OF PLANT DISEASES
The student who would become acquainted with the general
pathology of plants must have some previous knowledge of other sub-
jects, especially those which are concerned with the life of the plant.
To appreciate diseased conditions the normal state of the plant must
be understood. A phytopathology, which as a department of
study of
scientific inquiry concerns itself with plant diseases, therefore, presup-
poses that the would-be phytopathologist is acquainted with plant

morphology, systematic botany (fungi and flowering plants) histology,
cytology, embryology, genetics, physiology, with bacteriology, zoology
(especially entomology) chemistry and physics,^ as well as meteorology.
Plant morphology deals with the general form and gross structure of
plant parts, such as roots, stems, leaves, flowers, fruits and seeds. The
student should know the common fungi (see part I), the technique of
their study (see part IV), as well as the flowering plants, which act as
hosts to the bacteria and fungi causing disease. Histology is concerned
with the microscopic details of plants, while cytology treats of cell
structure and organization. Embryology, as a distinct subject of in-

quiry, embraces a study of their productive cells and organs. Genetics
is a new branch of inquiry. As Walter tersely put it, "The study of the
origin of the individual, which has grown out of the more general
consideration of the origin of species, forms the subject matter of
heredity, or, to use the more definitive word of Bateson, of genetics."
The functions of a plant are considered when we study physiology and
the chief divisions of that subject treat of the nutrition, growth and
1 Along these lines see suggestive papers by Ernest Shaw Reynolds: Plant
Pathology in its Relations to other Sciences. Science, new ser., xxvii: 937-940;
June 19, 1908.
271
272 GENERAL PLANT PATHOLOGY
irritability of the living plant organisms. A knowledge of insect life
is essential, as also the chemistry of the plant, of the soils, of the ferti-
lizers, of and fungicides. The physics of sap ascent,

the insecticides
of osmosis, of turgescence, and of the soil must be studied.^
The investigation of malformed organs and cells may be classified
under the head of Pathologic Morphology, and if the microscope is
used, it may include Pathologic Histology and Pathologic Cytology.
Disturbed conditions of the reproductive cells and organs bring about
anomalies in the offspring, so that genetically speaking freaks, bizarre
forms, or chimeras arise. Diseased conditions may be traceable to
disturbed nutrition, to excessive or retarded growth and to abnormal
irritability. Therefore to be a successful pathologist, one must be a
good morphologist, histologist, geneticist and physiologist.
Phytopathology is that phase of botanic inquiry which treats of the
diseases of plants. Its history dates from about 1850. Disease may be
looked upon as an unwholesome condition, derangement of, perversion
of, or departure from the normal in structure, in function, or in both com-
bined. It is a morbid One who studies phytopathology is con-
state.
cerned with the characteristic symptoms of disease (Symptomatology),

the interpretation of symptoms (Diagnosis), with the causes of diseases
(Etiology) and with the remedies (Therapeutics) and prevention of dis-
ease (Prophylaxis). Recently considerable attention has been given
prophylaxis, following out the old adage that an ounce of prevention is
worth a pound of cure. Curative agents are therapeutic agents.

ETIOLOGY. — At the outset it is important to consider the causes of
disease. These may be considered under two heads, predisposing and
determining.
Predisposing Causes of Disease. —The normal plant can to a cer-
tain extent ward off the attack of disease, but the power to do so varies
within wide limits, which may be conditioned upon racial, or individual
characteristics of resistance. The degree of this resistance determines

the degree of the immunity of the plant organism. It is well known
that the normal constitution of plants varies considerably in individ-
uals of the same variety and among different races and varieties of the
same species. Some individuals and varieties are constitutionally weak,
others are strong and resistant to external influences of every descrip-
iAppel, O.: The Relations between Scientific Botany and Phytopathology.

Annals Mo. Bot. Gard., 2: 275-285, February, April, 1915.
GENERAL CONSIDERATION OF PLANT DISEASES 273
tion. Such plants are designated as cast-iron, or hardy, while the

others are tenderand need constant care and attention on the part of
the cultivator. Such weakness of constitution, of histologic structure,
or absence of protecting chemical bodies in the cells of the plant may be
looked upon, other things being equal, as predisposing causes of diseases.
Such depend on the hereditary character of the plant, and in case of
varieties susceptible to disease may be designated hereditary predis-
position. Immunity, on the other hand, may be hereditary, as in the
case of the plants of strong constitution, or acquired. Resistance on the
part of certain plants may be due to the hereditary resistance of the pro-
toplasm, it may be
due to histologic structure, such as the presence of a
thick cuticle in the resistant form and its absence in the susceptible form,
for Sorauer has found that the resistance of different carnations was due
to the thickness of the cuticle. The habit of earliness, or lateness, may
be the determining factor in resistance. A late variety might be at-
tacked, because of growth in relation to the life history of some insect,
its
or fungous parasite, while for this reasonan early variety might remain
healthy. Morphologic peculiarities may be effective, for the investiga-
tions of Hecke and Brefeld have shown that in the varieties of wheat with
closed flowers, and which are close pollinated, therefore, the spores of
the loose smut fungus carried by the wind are unable to reach the stig-
mas, and hence, infection does not take place. Such varieties would be
smut proof for the simple morphologic reason that their stigmas are not
exposed to the smut spores. Osterwalder has indicated that varieties
of pears without an open channel from the calyx to the carpels are pro-
tected against infection by Fiisarium putrefaciens, while those varieties
with an open channel from calyx to t"he carpels are susceptible. The
habit of a plant, as to drying after a rain, may influence its disease
resistance, as shown by Appel.^ Infection of potatoes by the spores of
late blight, Phytophthora infestans is , due to the wind carrying the spores
to healthy plants where in the raindrops on the surface of the leaves
zoospores are formed.
The leaves of some varieties dry within half an hour after a rain,
while on others the leaves do not dry for several hours. Quick-drying
varieties are less susceptible than the slow-drying ones. In some mem-
bers of the pea family, the seeds are imbedded in a woolly outgrowth of
^ Appel, O.: Disease Resistance in Plants. Science, New ser., xli: 773-782,
May 28, 1915.
iS
the inner epidermis of the pod. It has been found that infection of the
seeds with Ascochyta pisi is by the presence of the hairs, for
facihtated
the fungus grows, as in a culture medium, and infects every seed, while
in the hairless forms infection takes place only where the seed actually
touches the infected spot of the pod.
The presence of certain chemic substances may explain immunity, for
the disease resistance of Vaccinium vitis idcea is supposed to be due to the
presence of benzoic acid. So, too, the presence of tannins may increase
the power of resistance to fungus and insect diseases, as indicated by
Cook and Taubenhaus.' Enzymes also play an important role in the
production of chemic substances, which increase disease resistance.
Such hereditary disease resistance may be made to play an important
partby breeding and growing the varieties which have been proved to
be disease resistant.
Immunity may be acquired by growing the susceptible form at a
different season of the year from its accustomed one. Grafting has
been used with success. The method is to graft a non-resistant variety
on a European vine on the American
resistant one, as in the case of the
vine,which resists the attack of the Phylloxera insect, which devastated
the European vineyards until this method was adopted. Crossing has
been resorted to as a second means of increasing disease resistance. The
weak variety is crossed with a disease resistant form to increase its
immunity. The third way to obtain immune forms is to select resistant
individuals and from them breed pure strains. This has been accom-
plished with some degree of success by Orton with cotton, by BoUey
with flax, by L. R. Jones with cabbage. It should be emphasized that
the inheritance of the unit characters and their behavior in the next
generation is one of the fundamentals of breeding resistant races.
Determining Causes. —Having considered the general reasons for the
predisposition of plants to diseases and the immunity of others, it is
important to describe next the causes which determine disease. These

may be divided into those of external origin and those ofinternal. The
external factors of disease are the chemical conditions of the soil, as a
determining cause, also the physical character of the soil. The influ-
ence of a superabundance of water, or its absence, is important. Cli-
^ Cook, Mel T. and Taubenhaus, J. J.: The Relation of Parasitic Fungi to the
Contents of the Cells of the Host Plants, i. The Toxicity of Tannin, BuU. 91,
Delaware College Agric. Exper. Stat., Feb. i, 1911.

matic and meteorologic conditions may be influential, when these dis-
turb the normal life of the plant. Light, heat, cold, rain, dew, hail,
frost, wind and Ughtning play an important role. The gaseous emana-
tions from gas pipes, smelter works, smokestacks, including soot, dust
from cement works, acids, poisons, and dye stuffs, which pollute streams,
all are determining causes of disease. Traumatism or mechanic injury
may be of various sorts and the effects are dependent upon the form and
severity of the injury, or wound.
Fig. III. —Rose-chafer {Macrodaclylis subspinosus). a. Adult or beetle; b,

larva; c, d, mouth parts of same; e, pupa, /, injury to leaves and blossoms of
grape with heetles at work. {From Marlalt in Quaintance, A. L. and Shear, C. L.,
U. S. Farmers' Bull. 284, 1907.)
Living organisms, whether animal or vegetal, may be the cause of

disease. All groups of animals may be considered, but the mammals,
worms and insects (Fig. iii) are of most importance and interest.
Insect depredations of plants are notorious and insects occupy first
place in their destructive effects on plants (Fig. 112). Various para-
sitic flowering plants are known, as well, as the bacteria and fungi,
for their disease-producing powers.
As an internal determining cause, the formation of enzymes under

abnormal conditions must be reckoned as causal, as well, as nutritive
disturbances which produce monstrosities and the like.
Having classified the chief causes of disease, a more detailed descrip-
tion of these factors should be put in a form available for student use.
Much of the information is and part of it is buried in foreign
scattered,
botanic and pathologic journals, which can be consulted only in the
largest scientific hbraries at home and abroad.
Fig. 112. — Oyster-shell scale {Lepidosaphes ulmi. After Quaintance, A. L., U. S.

Farmers' Bull. 723, Apr. 26, 1916.
The chemic condition of the soil, as a determining cause of disease,

may be considered from the standpoint of the normal influence of the
important soil ingredients, as contrasted with the absence or deficiency
of such elements. Potassium is usually found in young tissues and dis-
appears in the older ones. It is associated in some way with the for-
mation carbohydrates in the plant such as starch, sugar and cellulose.
of
The absence of potassium in the soil causes a cessation of growth, the
leaves fail to develop the power of forming starch within the greenr
coloring bodies, or chloroplasts. A plant which has failed to grow for
months will recover in a few days after potassium salts have been added
and after a few hours the formation of starch in the chloroplasts will
be detected.^ The storage of reserve materials is, therefore, inhibited,
and one finds in such plants, as the cereals, that the formation of green
parts is at the expense of the grain, and in the beet, the vegetative part of
the plant is at the expense of the fleshy roots. Potassium hunger causes"
in the potato and buckwheat a shortening internodes and a of the
convex bending of the leaf blades, which are spotted with yellow blotches.
Calcium is abundant in nature in the form of the carbonate which
forms the rocks known as marble and limestone. It is chiefly concerned
in the strengthening of the cell wall, where in such plants as Chara it is
deposited. It plays an important role in fixing the calcium oxalate

formed in the metabolism of the plant. Ecologists in Europe classify
many plants either as calciphile (calcium-requiring), or calciphobe
(calcium-hating). The apphcation of calcium to soils under certain
conditions promotes apparently the disease of beets called heart- or
dry rot. The chlorosis, or icterus, of the grape vine seems to be in-
creased in soils with a high calcium content. The accumulation of
oxalic acid in the absence of its fixation by calcium poisons the plant.
The formation of brown blotches on leaves, the yellowing, or brown-
ing of pine needles, the death of the root tips of water plants is associated
with the absence of calcium.
Magnesium is chemically allied to calcium, but it cannot replace
calcium in the economy of the plant. It apparently works together
with nitrogen in the formation of protoplasm, and has an influence in
the formation of chlorophyll, for plants grown without magnesium have
yellowish-green chloroplasts, and new cell formation does not proceed
readily. The absence of magnesium is shown in the pale-green color
of the chloroplasts, the yellow to orange-yellow blotches on the leaves,
and the brown spots on the stems. The amount of starch formed by
the chloroplasts is reduced, the internodes are shortened, the young
leaves do not unfold. These are symptoms associated with a
deprivation of magnesium.
Iron is necessary in the formation of chlorophyll, for if the plant is
grown in an iron-free remains permanently etiolated

solution, it
(blanched). The diseased condition which arises through the lack of

the requisite amount of iron is called chlorosis. Too much iron in the
soil acts poisonously.
^Hartwell, B. L.: Bull. 165, R. 1. Agric. Exper. Stat., May, 1916.
Sulphur and phosphorus are of some value in the production of

albuminous substances by the plant, and in the soil they exist mainly
as calcium sulphate and calcium phosphate. Phosphorus is in some
way associated with the formation of the crystalloids, globoids and
aleurone grains of the plant. Some soils are poor in phosphorus, so
that the agriculturist must supply phosphates. The_ deficiency_of
phosphorus is seen in the production of a red coloration in plarits. The
leaves are blotched with red and later the spots become dark brown.
The formation of flowers and seeds is partially inhibited. The absence
of sulphur is manifest in the poor development of the whole plant and
in the reduction in the amount of fruit produced.
Nitrogen enters largely into the living substance of the plant,

protoplasm. It is stored in the form of protein granules and aleurone
grains. In the life of the plant, it is concerned in the building of young
tissues, and in the metabohsm of plants, it appears in the form of aspar-
agin which in the soluble state is conducted through the bast portions
of the vascular bundles from one part of the plant to another part.
Some plants have a pecuhar relationship to nitrogen. Such are the
leguminous plants, which are provided with root nodules, where there
are nests of bacteria. These bacteria can utiUze free atmospheric nitro-
gen and later in the involution form as bacteroids, they are absorbed
by the green plant which is thus enriched with nitrogen. During the
period of entrance of bacteria into the root hairs, the young seed-
ling goes through a period of nitrogen starvation, when it appears
to flag, but growth and vitality when the
later, it regains its active
nodules have been formed. Contrasted with the same leguminous

species without nodules and when the root systems alone take up
nitrogen in the form of nitrates, the nodulated plant is larger and
stronger in every respect.
A deficiency of nitrogen in the soil can be detected in the case of
Indian corn and other agricultural plants by a general paling of the
green color, so that in some cases the plant becomes yellowish-green.
Klebahn^ indicates that the leaves of beets, buckwheat and potatoes
assume a yellowish color with a deficiency of nitrogen, and as the leaves
dry, they become yellowish-brown. The prothaUia of ferns in a nitrogen-
free nutritive solution do not form meristem or archegonia. Excessive
supplies of nitrates in their application to cultivated fields stimu-
1 Klebahn, Prof. D. H.: Grundzuge der Allgemeinen Phytopathologie, 1912: 11.
lates in the case of the crops grown upon such fields the development
of the vegetative organs and, therefore, delays the formation of flowers
and and the ripening of seeds. Such delay may mean the attack
fruit
of parasitic fungi. For example, a large field of winter wheat which
had been sown about the end of October was much attacked by stink-
ing smut (60 per cent.), while the adjacent fields belonging to the same
farmer, under the same variety of wheat and treated in a similar
manner, but sown early in October showed no sign of infection. With
fruit trees, one notices greater frost susceptibility in those plants which
have received an excessive nitrogen supply. Lipman (Science, new
ser. xxxix: 728-730, May 15, 1914) has suggested that the poor nitri-
fying power of soils is a possible cause of "die-back" (exanthema) in
lemons. It has been a serious disease with the citrus growers of
Florida and California.
Physical Character of the Soil.- —The physical character of the soil
is of great importance as a determining cause of disease. When we

speak of the physical character of the soil, we refer to the size of its
particles, the relation of these particles to each other, the presence
of colloidal bodies, the presence of air spaces associated with the air
content, the distribution of the water through the soil, the presence or
absence of organic matter, or humus, the color and temperature of the
soil. Of greatest importance to the life of the plant is the water
which is available for the needs of the plant. ^ A too plentiful sup-
ply of water causes the formation of a wet ball of roots with the
formation of alcohol. Frequently gardeners fearing that the soil is
dry, water potted plants with more water than the plants actually
need, so that the lower part of the soil is continuously saturated with
water. Alcohol is formed and decay of the roots sets in, because they
are gradually suffocated. Too little water on the other hand causes
a drooping or wilting of the plant, and if water is not supplied in
time permanent wilting and death of the foliage results. But a
diminished water supply may be decidedly beneficial to plants, as it
has been found that the formation of flower buds is best initiated by
preserving a period of rest following a diminished water supply.
Different plants have different water requirements and these require-
ments vary with the season of the year and the development of
* Cf. SoRAUER, Paul, Lindau, G. and Reh, L., transl. by Dorrance, Frances:
Manual i, parts i and 2.
of Plant Diseases, vol.
28o GENERAL PLANT PATHOLOGY
the plant. As an illustration of this may be cited the planting of the

Carolina poplar on the open porous sandy soils of New Jersey. About
Philadelphia, where the tree is largely planted, it grows rapidly with
a dense crown of dark-green, foliage leaves. In New Jersey, it grows
crown is more open by a wider spacing of the branches
less rapidly, its
and the leaves have a greenish-yellow appearance and drop off earlier
in the autumn than similar trees on the Pennsylvania side of the Dela-
ware River. This difference is without doubt associated with the water
requirements of the tree, for on the Pennsylvania soils, it can secure
abundance of water during the growing season, while in the New Jersey
sands, owing to their porosity and the rapid drainage of water through
them, the Carolina poplar does not receive sufl&cient amounts of water
for its most vigorous growth.
The experiments of Miinch^ throw important light on the content
of water and air in the tissues as a determining factor of disease of
woody plants, such as on forest and fruit trees. He has shown that the
greater number of the wood-destroying fungi require a large amount of
air and are able grow only when a maximum amount is present. The
to
air content of the tissues is dependent on the water supply and trees
with narrow annual rings are more resistant than those with broad ones,
because the former contain m.ore water and less air relatively. Differ-
ent annual rings of the same tree may be attacked differently.
The decayed rings of wood in such trees are always the broad ones.
The tissues of vigorous branches are rich in water and poor in air and
infections do not always penetrate to such regions. The healthy bark
of beech trees in winter-rest contains 19 to 20 per cent, of air and at
the time of budding the air diminishes to 11 per cent., rising afterwards;
This is correlated with the canker disease, Nectria ditissima, which in
Europe does its damage during the winter months, while during the
vegetative period itHence, we have opened here a very profita-
ceases.
ble line of investigation to determine the relative amounts of air and
water with respect to immunity, or its absence. Again, in the irrigated
districts of America, the fruit trees have only a few diseases due to
species of Valsa and other species of fungi. Defective irrigation may
bring about the prevalence of the die-back diseases, which may be reme-
^MtJNCH, E.: Untersuchungen iiber Immunitat und Krankheitsempfanglich-

keit der Holzpflanzen. Naturwiss. Zeitsch. f. Forst. und Landw., 7: 54-75, 87-
114, 129-160, 1909; Appel, O.: Phytopathology & Scientific Botany, loc. cit.
died by changing the system of irrigation. The land should be irri-
gated at the time when the trees contain small amounts of water and
much air, so as to prevent an excessive decrease of water in the tissues.
The condition of the humus has a rather remarkable influence on
the growth of plants. Ericaceous plants, such as the trailing arbutus
{EpigcEa repens), wintergreen {Gaultheria procumbens), bearberry
(Arctostaphylos iiva-iirsi), blueberry {Vaccinium corymhosum) flourish
in an acid humus and if the attempt is made to grow such plants
under other conditions, they languish and die. Other species like In-
dian turnip {Ariscema triphylla), blood root (Sanguinaria canadensis),
rue anemone {Anemonella thalidroides) grow best in a leaf- mould humus
which is neutral or slightly alkaline. Reverse the reaction of the soils
about these plants and they gradually die.
The presence of an impervious hard pan below the surface soil is
a condition which prevents the normal development of trees, as I have
shown in my book on the "Pine Barren Vegetation of New Jersey,"
where in the region known as the Plains, the pitch-pine trees are kept
dwarf owing to an impervious subsoil layer. There the trees flourish
for a number of years, then begin to suffer until unable to penetrate the
deeper layers of thesoil, they finally succumb to be replaced by younger
trees which meet the same fate.

Climatic and Meteorologic Factors. The most important —
climatic factors, which may be looked upon as in any way related to
disease production, are light, heat, precipitation (rain, dew, frost, snow,
hail and ice) wind and electricity (lightning, etc.).
Light is Carbon
essential for the life functions of all green plants.
dioxide and water are united by the energy form starch.
of sunlight to
The synthesis takes place in the chloroplast, starch being formed as
the first visible product. Ordinary sunlight of a bright, clear day may
under certain conditions of plant growth be too intense and it acts
prejudicially. The writer has frequently noted, that garden plants
suffer,if after a wet, cloudy spell during the rapid period of growth,
they are exposed to a bright sun without protection. It takes a few

days of bright light to sun-harden the plants. Trees, especially with
a smooth bark, which have grown in a very dense wood, and then
suddenly isolated in later life, suffer from scorching of the cortex.
They are sunburned. Plants grown in greenhouses, which have been
painted with whitewash to reduce the intensity of the rays of light, have
their leaves burned if part of the whitewash is removed. The light

passes through the opening thus made and the leaves on which it is
concentrated are scorched.

Several diseases of plants are caused by too brilliant sunlight. Such
are sunscald, sunscorch and bronzing.^ Sunscald may follow as a
result of too intensive sunlight, as, for example, when certain fruit trees
are stripped of their foliage in summer, such as sometimes results from
the ravages of the gypsy moth. In such instances the new unripened
wood sunscalds badly. Sometimes it is associated with severe and
abrupt changes in temperature on non-ripened wood. "Sunscorch"
is a term applied to the burning of foliage in summer during periods
when the soil is dry, and is also common to evergreens during warm
wiildy days in spring before the frost is out of the ground. Any
defects in the root system which prevent root absorption may cause
sunscorch. "Bronzing" of leaves is a form of sun scorch characterized
by the occurrence of a reddish-brown or bronze color of the leaf. It is
caused by a lack of soil moisture, or defective root absorption during

dry, hot periods.
Too much shade is also detrimental to plants, as is seen under the
dense canopy of beech trees on a lawn, where nothing will grow, not
even a blade of grass. The grasses, etc., die of inanition. The condi-
tion known as etiolation originates where a plant is grown in the dark,
or in subdued sunlight. Growth in darkness leads to important modi-
fications in the general habit and structure of a plant. If we take a
potato plant and raise it in the dark, we find the etiolated shoot has
a white stem and leaves which are at first pinkish, and subsequently
pale yellow, and the absence of chlorophyll is noteworthy. The inter-
nodes are long and slender and the leaves are small compared with the
green plant and there are corresponding anatomic differences. Morn-
ing glories raised in greenhouses in the winter do not twine. They
grow from four to five inches tall and have only one to two flowers.
Heat as a factor in the growth of plants is well known. Each plant
has its minimum, maximum and optimum degree of heat. The dis-
tribution of plants over the larger stretches of the earth's surface is
associated with the amount of heat that the different plants receive.
The absence of heat, where the plant is exposed to a temperature below
1 Stone, George E.: Injury to Vegetation Resulting from Climatic Conditions.
Monthly Weather Review, 44: 569-570, October, 1916.
freezing, is noteworthy. The death of cells rich in water, when exposed

to low temperatures, seems to depend upon the conversion of the water
extruded into the intercellular spaces into ice. The parenchymatous
tissues are ruptured and crystals of ice are formed. The water, there-
fore, which is in the cell reaches the surface and the cell sap diminishes
in amount and there may be chemic changes in the cell as a result of
freezing, for in some cases .the leaves assume a leathery brown color.
Long exposure to cold may lead to the actual disorganization of the
protoplasm. It, however, does not always follow that the formation
of ice in the intercellular spaces necessarily involves death. Slow
thawing may be followed by a return of the water to the cells until
the normal equilibrium is restored and the cells continue to live. A

rapid thawing, however, causes death of the cells, because the water is
not reabsorbed. Frost-killed twigs and branches are more susceptible

to the entrance of saprophytic fungi such as species of Nectria, Dasy-
scypha, and Valsa. The exposure of roots during a snowless winter
may lead to their disturbance by freezing. The anatomic changes
induced by freezing are frost blisters, such as appear on the leaves of
fruit trees and cereals, and frost cracks, which may ultimately heal
over, producing an external ridge or enlargement. The fruit-grower
can distinguish four kinds of winter injury to his trees. First, the
frosting of the blossoms after they begin to open; second, the freezing
of the buds in winter; third, the freezing of the twigs and branches;
fourth, root freezing. It may happen that early in the spring the
peach trees come into bloom. Then on a cold cloudless night with no
wind the temperature sinks below freezing and the partially open
flower buds are nipped by the frost. About twenty years ago the
upper Mississippi Valley was visited by an unusual cold wave. The
frost penetrated to great depths and the cold was so intense that the
tree roots were actually frozen in the soil.^
The formation of ice fringes upon plants has been investigated

exhaustively by Coblentz,- with the dittany, Cunila mariana. He
* Consult Waugh, Frank Jack Frost's Tricks. The Country Gentleman,
A.:
Feb. 6, 1915, p. 213. Wilson, Wilford M.: Frosts in New York. Bull. 316,
Cornell University Agri. Exper. Stat., June, 191 2. Chandler, W.H.: The Kill-
ing of Plant Tissue by Low Temperatures. Research Bull. 8, Coll. of Agric,
Univ. of Mo., Dec, 1913.
2 Coblentz, Wm. W. The Exudation of
: Ice from Stems of Plants. Monthly
Weather Review, 42: 490-499, August, 1914.
found that the ice fringes are formed when the temperature falls to
freezing. They are formed on the outer surface of the plant. The
growth of the ice fringe ceases when the ground is frozen to a depth of
2 to 3 cm. and when the moisture in the stem is frozen. The dimen-
sions of the fringe depend upon the rate of evaporation of water from
the stem up which it rises by capillary action and upon the amount
of moisture in the ground. Clouds and fogs in some regions have an
important effect on vegetation.^ The two forms of foliage leaves on
the branches of the redwoods of California are conditioned upon the
height of the fogs which drift in from the Pacific Ocean. The leaves
on the fog-exposed branches are flat and divergent, while those on the
sun-exposed branches above the fog level are scale-like and appressed.
The London fogs work detrimentally to outdoor and greenhouse plants,
and in Egypt, the cotton capsules long exposed to fog are more in-
fested with black moulds. Dew, which lodges on the margins of leaves,
is responsible for the entrance of fungi by their spores lodging in the
dewdrops and germinating there.

The weight of snow and ice breaks off the limbs of trees, breaks down
herbaceous plants, and this opens up the way for the entrance of
various parasitic fungi. Ice or sleet storms are especially severe at times
to trees. The year 1902 was noted for two exceptionally destructive
ice storms which visited the Philadelphia region. One of these storms
occurred on Friday, Feb. and the other on Saturday, Dec. 13.^ The
21,
storm of Feb. 21 was accompanied by high winds and did an irreparable
damage to the fruit, forest and shade trees. Meteorologically speaking,
regions of strongly variable temperature are subject to occasional
winter storms in which the precipitation occurring as rain, freezes as
soon as it touches any solid body, such as the branches of trees, telegraph
wires or the ground. This happens when the ground and the lower
air have been made excessively cold during a spell of clear anticyclonic
weather, when a moist upper current in advance of an approaching
cyclone brings clouds and rain. All our meteorologists prefer to call
such storms ice storms; locally near Philadelphia they are denominated
sleet storms. The weight of ice which such limbs carry is astounding.
1 Weiss, F. E., Imms, A. D., Robinson, W.: Plants in Health and Disease,
1916; 54-56.
" Harshberger, John W. : Relation of Ice Storms to Trees. Contrib. Bot.
Lab. Univ. of Penna., II: 345-349, 1904.
The author found the weight of a branch of Liriodendron tuUpiJera with

iceupon it to be 50 grams, without ice 9 grams; so that the ice weighed
41 grams, giving a ratio of i : 4.5. Juniperus virginiana with its ice
load weighed 310 grams, without ice 13 grams, making the weight of ice
297, a ratio of i : 23. Beginning with Dec. 5, 1914, a combination

rain, snow and ice storm swept across the Eastern States doing much
10
12
13
'^ H@ '5® '^®
17
Z2ZZ2
Fig. 113. —
Sectional view of twigs and leaves of various plants showing load of
ice carried during the ice storm of Feb. 12 and 13, 1916. i, Acer plalanoides; 2,
blade of grass; 3, Chionanthiis virginicus; 4, Diervilla florida; 5, Forsythia suspensa;

6, Liguslrum vulgare; 7, Liriodendron lulipifera; 8, Platanus orientalis; 9, Populus
alba; 10, Populus delloides; 11, Quercus paluslris; 12, Syringa vulgaris; 13, Tilia
americana; 14, Tecotna radicans; 15, xanthoceras sorbifolia; 16, Spiraea Thunbergii;
17, leaf of Rhododendron maximum; 18, icicle on tip of Rhododendron maxifnum,
leaf hanging down.
local damage^ and again on Friday, Dec. 31, a severe ice storm visited
the mountain region of Pennsylvania contiguous to the Juniata Valley
and Susquehanna River. During the afternoon of Saturday, Feb. 12,
191 6, a cold rain began which continued well into the night, coating the
pavements, streets, and trees with hard ice. On Sunday morning,
Feb. 13, men, boys and girls took advantage of the icy streets to skate
Illick, J. S.: A Destructive Snow and Tee Storm.
1 Forest Leaves, xv: 103-
107, Fehriiari', tqi6.
286 GENERAL PLANT TATHOLOGY
upon them and this unusual sight was stopped by a snow storm, which
followed on Sunday morning. The trees were loaded to the breaking
point. During the continuance of the storm, small branches were
taken oflf thirteen trees and shrubs and a blade of grass growing in West
Philadelphia, and the thickness of ice upon them measured with a
pair of compasses. The accompanying figures drawn life size show the
relative thickness of the load of ice borne by the twigs, whose thickness
is shown in the drawings (Fig. 113).
Fk, Mi t liii I h)nt\ iiDii h tiiui ]iIl ntifully supiilicd with ground
I
water ni-ai the builacc dcprcbbiuu ot the glacial uutwash plain at Westbury,
L. I., July. 1915.
The fall of hail stones may, if they are large enough, cause the
decortication of twigs, or the abrasion of other plant parts, thus per-
mitting the entrance of destructive bacteria and fungi to the interior
of the plants.
Wind is an active agent buds and limbs and
in the breaking off of
the formation of dangerous wounds. In such situations, as high moun-
tains, sand dunes and rocky shores, where trees are exposed to the
forcible action of the wind, they assume a windswept, bisected, or
prostrate form, which is characteristic and picturesque (Fig. 16).
Fig. 115. —
Unhappy vase-shapei.l white elm, I'ini:, ,,<,..>i. luo yards south
', . ,
of ahappy larger elm both growing on the outwashed plain, Westbury, L. I., July,
1915-
Fig. 116. — Wind-swept white poplar, Populus alba, Nantucket, Mass., August, 1915.
Strong winds increase the amount of transpiration, so that fre-

quently we find there is a balance established between the absorbing
root system and the transpiring amount of
leaf system, so that the
foliage is amount of water lost by
determined accordingly. If the
transpiration exceeds the amount absorbed by the roots the plant
usually succumbs. Happy trees are those in which the amount of
water available exceeds the amount transpired, while unhappy trees
are suffering physiologic drought through the action of the wind in
moving water faster than it can be supplied (Figs. 114, 115, 116).
Such trees are seen in planted specimens in Long Island, Nantucket
and along our seacoasts. With tornadic winds, trees are uprooted in
general and irreparable damage is done.
The effect of lightning is a marked one, as a determining factor in
disease. Recently Jones and Gilbert^ have published a paper on the
lightning injury to potato and cotton plants. One case occurred in a
field at Monetta, S. C. in the summer of 1913. The cotton plants
were fully grown and after a severe electric storm on Aug. 3, all the
cotton plants were killed over an area three rods in diameter. The
leaves wilted, died and blackened, but remained attached to the plants.
The most pronounced effect, however, was on the stem and root system.
Other cases are cited of a similar nature in Europe and America.
The action of lightning on trees is variable. The tree may be
scorched, it may be stripped of its leaves, it may be cleft longitudinally,
or, more rarely, severed horizontally. Sometimes the bark is stripped
from only one side, occasionally without a trace of burning: at other
times, it may be riddled, as by worms, with a multitude of Httle holes.
The lightning furrows may be single, double, oblique or spiral. If the
tree is inflammable a firemay be started. Such tall trees, as the big
trees of California, have been struck repeatedly by lightning and their
leaders broken and their tops stunted as a consequence. From early
times, there has been a current belief that certain trees attract the light-
ning, that others are not struck. The elder Pliny beheved that "Light-
Jones, L. R. and Gilbert, W. W. Lightning Injury to Potato and Cotton

^ :
Plants. Phytopathology, 5 94-101, with plate, April, 1915; Jones, L. R.: Light-
:
ning Injury to Kale. Phytopathology, 7: 140-142 with i fig., Apr., 1917;,

Stone, George E.: Electrical Injuries to Trees. Bull. 156, Mass. Agric. Exper.
Stat., Oct., 1914.
ning never strikes the laurel.'' In certain parts of the United States,
it is held that the beech tree is never struck.
"Avoid the oak, flee from the spruce, but seek the beech," yet in
the Garden Magazine for January, 19 16, is given a photograph and an
account of a fine beech tree which was struck by lightning in Pennsyl-
vania about the middle of June. Plummer^ sums up his investigations
on the relation of lightning and trees, as follows:
1. Trees are the objects most often struck by lightning because:
{a) they are the most numerous of all objects; (b) as a part of the ground,
they extend upward and shorten the distance to a cloud; (c) their
spreading branches in the air and spreading roots in the ground present
the ideal form for conducting an electrical discharge to the earth.
2. Any kind of tree is likely to be struck by lightning.
3. The greatest number struck in any locality will be of the domin-
ant species.
4. The likehhood of a tree being struck by lightning is increased:
(a) if it is taller than surrounding trees; (b) if it is isolated; (c) if it is
upon high ground; (d) if it is well (deeply) rooted; {e) if it is the best
conductor at the moment of the flash; that is, if temporary conditions,
such as being wet by rain, transform it for the time from a poor conduc-
tor to a good one.
5. Lightning may bring about a forest fire by igniting the tree itself,
or the humus at its base. Most forest fires caused by Ughtning proba-
bly start in the humus.
Experiments on the electric conductivity of various woods shows
that this conductivity depends upon the water content of the wood.
When absolutely dry none of the specimens showed conductivity, but
the resistance of all was practically infinity.
Effect Smoke, Soot, Gases and Smelter Fumes on Plants. The
of —
smoke, which is destructive to vegetation under our modern conditions,
is derived from four sources of supply: (i) smoke from manufacturing
plants, or from large buildings; (2) smoke from special concerns, such
as the electric of electric trolley lines; (3) smoke from rail-
power plants
road locomotives; smoke from the chimneys of dwelling houses.
(4)
Smoke belts have been drawn by students of the problem to determine
the area influenced by the smoke. From a survey made for the City
^Plummer, Fred G.: Lightning in ReUition to Forest Fires. BulL in, U. S-
Forest Service, 191 2.

19
of Des Moines, Iowa, by A. L. Bakke,^ it. has been found that conifers
are more susceptible than deciduous trees. The direct injury is seen
in the deposit of the tarry matters of the smoke in the stomata of
nearby plants; leaves and leaflets are shed, or assume abnormal shapes,
and the formation of foodstuffs is hindered. The sulphur dioxide and
acetylene as constituents of smoke act toxically upon the plant. The
work which has been done in the United States may be summed up as
follows: Burkhart states that injury from gases is the result of the
chemical constituent of the smoke and is not due to the clogging of the
stomata. The investigation of J. K. Haywood^ in the vicinity of
the famous smelter at Anaconda, Mont., is of importance. He finds
that trees are injured at a considerable distance; that very small
amounts of SO2 are toxic to plant growth; that water used for irrigation
purposes often has sufficient copper in it to be toxic to plant growth
and that certain trees, as the juniper, are more resistant than others.^
Officials of theForest Service are watching with interest the develop-
ments in the matter of the fumes from copper smelters in the southern
Appalachian Mountains. The service has been interested for years,
but since the acquirement of land in that section under the Weeks law
for forestry and watershed protection purposes, it has been felt that
the destruction of forests by the action of the fumes should be stopped.
W. L. Hall, forest supervisor of the seventh forest district, has
recently submitted to the bureau a report upon the subject. It seems
that one or more of the purchase areas established in the southern
Appalachians are endangered by the fumes, which are of a sulphuric
nature.
iBakke, a. L.: The Effect of City Smoke on Vegetation: Bull. 145, Agric.
Exper. Stat. Iowa State Coll. Agric. & Mech. Arts., October, 1913; The Effect
of Smoke and Gases on Vegetation. Proc. Iowa Acad. Sci., xx: 169-187, with
bibliography; also Anderson, Paul J.: The Effect of Dust from Cement Mills
in the Setting of Fruit. The Plant World, 17: 57-68, March, 1914.
2 Die Beschadigung der Vegetation durch Rauch. Handbuch zur Erkennung
und Beurteilung von Rauchschaden von Professor Dr. E. Haselhofe, Vorsteher
der landwirtschaftlichen Versuchsstation in Marburg i. H., und Professor Dr. G.
LiNDAu, Privatdozent der Botanik und Kustos am Kgl. Botanischen Garten in
Dahlem. Mit 27 Textabb.
3 Haywood,
J. K.: Bull. 89, Bureau of Chem., U. S. Dept. Agric, 1905; In-
jury to Vegetation and Animal Life by Smelter Wastes. Bull. 113 revised, Bureau
of Chem., U. S. Dept. Agric, 1910.
''
The Southern Lumberman, xxix: 27, Nov. 6, 1915.
GENERAL CONSIDERATION OF PLANT DISEASES 29T
The fumes are apt to destroy any vegetation within a radius of

several miles of the southern copper smelters. They are also working
destruction in the forests of Montana, California and other states. The
action of the fumes is peculiar and variable. Some trees succumb
quickly to their deadly effects, notably white pine. Other trees are
more resistant, including spruce, it is said. Nor does the gas act uni-
formly. Its effects vary with topographic conditions. The fumes
will travel long distances up a canyon or narrow valley, destroying the
woods in it, but leaving trees uninjured on either side. Again, it is
said, the sulphur fumes collect in globular form something like soap
bubbles, and drift away, doing no damage until the globular mass dis-
perses, sometimes at quite a distance. To a greater or less extent,

forests at a distance of several miles from copper smelters may be
damaged by the fumes.
It is admitted that the fumes can be controlled by condensation or
consumption, but the commercial practicability of the process is the
pending question. The fumes can be and are to a certain extent con-
verted into sulphuric acid, but the smelter people claim that the market
for this product is it does not pay to produce more than
Hmited, and that
a certain quantity of an oversupply sends the price down, which
it, as
would make it not worth while to control the fumes further.
Just now considerable trouble is being experienced in Tennessee
and Georgia on account of the sulphur fumes from copper plants.
In 1905 the State of Georgia took action against these companies,
alleging that they permitted a discharge of gases, which destroyed
vegetation, including forest trees, in that state. The companies were
forced to install plants to utiUze a considerable percentage of the sul-
phuric acid gas. These plants, however, have been unable to utilize
a sufficient quantity of the gas, and last spring the supreme court de-
cided to have a special expert ascertain the amount of gas released,
and the amount which ought to be utilized in order to make the
fumes harmless.
The time is close when the pathologist will have to take up this
question of fume damage, since large sections of the Cherokee area are
subject to such damage, and it is reported that the injury has extended
to the Georgia area.
The injurious effect of illuminating gas and ethylene upon flowering
carnations has been investigated by Crocker and Knight.^ The best
1 Crocker and Knight: Botanical Gazette, 46: 256-276, 1906.
work been done by Brizi,in England by Crowther and Rus-

in Italy has
ton.^ Recently in America J. F. Clevenger has published a bulletin
(No. 7), on "Smoke Investigation" for the Mellon Institute of Indus-
trial Research and School of Specific Industries, University of Pitts-
burgh, 19 1 3, with plates showing the effect of the smoke on the struc-
ture of the woody specimens examined by him.
Illuminating gas absorbed by the soil from nearby gas pipes is
injurious to trees and has frequently killed them outright, as instance

a group of street trees in Merchantville, N.
J., a few years ago, which
were killed in way, and for which the owner, Edwin C. Nevin,
this
received damages from the gas company for $1500, as a result of a
successful lawsuit. All the ordinary gases used for lighting and
heating are injurious and act much in the same way. Such are water
gas, coal gas, gasoline, acetylene and others. The first effects of gas
poisoning, may be seen in the foliage. The leaves turn yellow and in
some cases drop off, while the leaves of other trees fall while still green.
The water containing the gas in solution passes into the stem and the
wood and the cambial portion becomes abnormal. The underlying
tissues, cortex, bast and cambium die. Soon various species of fungi
gain access to the tree and cause its decay. With the Carolina poplar
especially, the bark, cortex, etc., on the trunk towards the source of
absorption showed three or four vertical cracks, or lesions, one-half to
two and a half feet long. The bark on the sides of these cracks bulged
out considerably, and an investigation showed a thix:k layer of soft
parenchymatous tissue extending to the wood and derived from the
cambium zone. Later this tissue turned brown, disintegrated and
became slimy in appearance, the sUme exuding from the cracks.
.
Illuminating gas dissolved in water in which willow cuttings were kept

stimulated the opening of the foliage buds several days earlier than plants
grown in water not charged with the gas. Stone^ found that the effect
of gas on lenticels was to increase their size, especially under water
charged with the gas. This appears to be a general response on the
part of the plant tissue to a demand for oxygen.
That the trees, shrubs and flowering plants in our large cities and
1 Journal of Agricultural Science, 4: 25, 1911.

2 Stone, G. E.: Effects of Illuminating Gas on Vegetation; 2sth Annual Rep.
Mass. Agric. Exper. Stat., January, 1913; Shade Trees, Characteristics, Adapta-
tion, Diseases and Care. Bull. 170, Mass. Agric. Exper. Stat., Sept., 1916, p. 220.
GENERAL CONSlDERyVTION OF PLANT DISEASES 293
in the country along our trunk-line railroads are subjected to conditions

which cause unhealthy growth and disease has been proven abundantly.
Large factories, power plants and railroad locomotives are pouring out
volumes of smoke, which alone is highly injurious, but in addition the
acid which is formed in the combustion of coal, when dissolved in rain
water, has injurious effect upon fohage and other plant parts. Its
action is seen in the corrosion of tin roofs, rain pipes and ornamental
iron work about city houses.
The following note is of interest to the plant pathologist and plant
physiologist.^ During the night of Sept, 19, 1913, a light rain fell,
followed by a fine drizzle in the early morning of Sept. 20. The wide-
open campanulate flowers of the common morning glory (Ipomcea
purpurea Roth), growing on a lot in West Philadelphia, four or five
blocks from the Pennsylvania Railroad, had their usual quota of rain-
drops studded over the upper, inner surface of the purple corollas.
Wherever the drops touched the surface of the corolla, the purple
color was changed to a pinkish red, and in the process of evaporation
of the raindrops the acid of the drops was concentrated, so that after
the complete disappearance of the drops a brown spot was left in the
center of the pinkish red circles of discoloration. The explanation of
the alteration of color is found in the change of the sap of the corolla
cells,where touched by the acid raindrops, from an alkaline to an acid

reaction. A similar change can be induced in blue violet petals by
bruising them slightly and placing them in an acid liquid. The petals
change, hke blue alkahne litmus paper, from blue to red, and this re-
action with violet petals has proved useful in the physiologic laboratory
in the absence of litmus paper. In nature a reverse change, which
illustrates the same chemic principle, takes place in many flowers of
plants belonging to the family Boraginaceae. For example, in
Symphytum and Mertensia, the red flower buds, the cells of which have
an acid cell sap, gradually change to blue as the flowers open. That
this is a chemic change is proved by treating the red buds with an
alkaline fluid and the blue flowers with an acid one.

Similar spotting, but less clearly discernible and demonstrable, as
the delicate reaction with morning-glory flowers, undoubtedly occurs
on leaves and fruits, and the suggestion is made here, that such spots
1 Harshberger, John W. The Acid Spotting

: of Morning Glories by City Rain.
Science, new ser., xxxviii: 548, Oct. 17, 1913.
caused by the acidity of raindrops serve repeatedly as the points of

entry of parasitic fungi, for there are many leaf spots and fruit spots
that show concentric rings of diseased tissue in the earliest lesions pro-
duced. A fungus, which is stimulated to growth by an acid condition
of the cell sap, would find ideal conditions for the commencement of
growth by entering areas influenced by acid raindrops.
—
Traumatism. Traumatism, or mechanic injury, may be of various
sorts and the effects are dependent upon the form and severity of the
injury. Mechanic injury to the plant usually takes the form of wounds,
which may be divided into natural and artificial. Natural wounds are
those which are produced on plants living in a state of nature, or in a
cultivated state in which other natural agents are concerned in their
production, man's activity not being considered. Insects and worms
may make burrows in the organs of plants. For example, bark boring
is accomplished by species of beetles, so also are tunnels through the bark
and the wood. Pith flecks are minute brown specks, or patches, found
in the wood layers of trees. They consist of holes formed by boring
insects filled with dead parenchyma cells, or dead empty cells filled
with fungous material. Eroded and skeleton leaves, and shot-holes
in the leaf tissue are directly traceable to the work of cutting insects.
Frost cracks are longitudinal wounds produced by the rending action

of the frost on the bark and wood of the trees. Sometimes this takes
place with a loud report. The attempt on the part of the plant to
heal the crack generally produces a frost ridge. Rents made by light-
ning also occur. Strangulations are lesions formed by woody vines, by
telegraph wires, or the like pressing on the outer surface of stems which
grow about the compressing object and create additional pressure, so
that the compressed tissue dies. Callus forms above the wounded
areas formed by compression. Large hailstones sometimes produce
bruises on the bark of young trees, as also the bombs shot out of vol-
canoes. The abrasion of a tree by the branch of a neighboring tree
rubbing against it or the cutting of large lateral roots in laying curb-
stones must be classed as wound phenomena. Wounds are also
formed by the teeth and horns of various mammals. Rodents, such
as mice, rats, beavers and squirrels, are responsible for wounds pro-
duced by gnawing with their chisel-shaped incisors. Bark is rubbed
oft", or scratched by the horns and antlers of animals of the cow and
deer tribes. Wounds are formed by the breaking off of branches

under the tearing action of the wind, or by the breaking action of the
weight of the ice and the snow of winter. The repair ofwounds
will be discussed with the consideration of the pathologic anatomy
of plants, which will form a separate chapter of this treatise.
wounds are due to the influence of man. The ploughing,

Artificial
discing, harrowing and cultivation of the soil frequently abrade roots,
break them off, or seriously wound them. Limbs are broken off and
bark removed by farm implements. Knife and axe wounds are easily
recognized by their sharp character, where the cut may have been
made vertically, obliquely, or horizontally. The stripping off of
pieces of bark opens up the inner tissues of the stem to the attack of
the agents of disintegration and decay. The removal of twigs and
branches in the ordinary operations of pruning opens up wounds, some-
times of a gaping character. The ringing, girdling, or scarification
of trees for various purposes, if not properly performed, opens up
wounds, so do nails, or spikes driven into the tree for various purposes
and the placing of electric cables and telegraph wires along our streets
and roads results in the removal of tree tops. The habit of cutting
initial letters and monograms in smooth-barked trees, such as the
beech, or the removal of sheets of birch bark, opens up wounds of vari-

ous menace to the health of the tree. Injuries due to man-created
environment may be of a thousand and one kinds too numerous for
even a brief mention.
Animate Agents of Disease. —These may be divided into two
groups, namely, animal and plant. Many animals are responsible for
the production of wounds and the destruction of plant parts. Man,
cattle, herbivorous animals, rodents (mice, rats, squirrels, rabbits), and
birds do great injury to plants by their horns, teeth, claws and beaks
(woodpeckers). Among the invertebrates are to be included the in-
sects, mites and worms. Certain nematode worms attack the roots of
a large variety of plants and produce galls of characteristic form and
appearance. Phylloxera, an hemipterous insect, winters on the roots
of the grape, mostly as a young wingless form. Wingless individuals
then abandon the roots and crawl up the stems to the leaves, where they
form galls. Formerly introduced into Europe, it was very destructive
toEuropean grape vines until it was found that it could be controlled
by grafting the European vine on the roots of American varieties.
Insects injurious to plants may be roughly divided into two groups:
those with mandibulate, or bithig mouth parts, and those with hausti-
late, or sucking mouth parts. The first group includes the insects
that bore into wood, those that bite off the leaf surface (Fig. in) and
thus skeletonize leaves and those which tear or bite pieces out of leaves
and other plant parts (Fig. in). The sucking insects include those
like the bugs, aphids, or plant lice, and scale insects (Fig. 112), which
cannot be destroyed by stomach poisons. These
by suck- latter insects
ing the plant juices do irreparable damage to all kinds of fruit and
shade trees, and reduce materially the yield of agricultural and
horticultural crops.
Of the mites, the most destructive is the red spider Tetranychus
niytilaspidis. The red spider is probably identic with the insect
known throughout Florida as the Purple Mite. It is quite a small
insect, yet distinctly visible to the naked eye. They appear during
summer in great numbers and damage the oranges by causing the
fruit to drop and injure the foliage leaves so that they cannot perform
their functions properly. The leaves become spotted and lose their
glossy green color. The males and females are protected by stiff hairs
and their color is purplish, or reddish-purple in the old insects, but of a
lighter red when young.
Animal galls are of various kinds. Those due to insects are charac-
teristic and will be described, when the pathologic anatomy of plants
is considered in detail.
The field of Economic Entomology is a special one and there are
bulky treatises dealing with various phases of it. A useful book, and
written in an easy style is one by John B. Smith, late Entomologist of
the New Jersey Agricultural Experiment Station, and is entitled
"Economic Entomology for the Farmer and Fruit Grower." etc.
Although published in 1896, it is still a useful book. A few American
classics on the subject may be mentioned, as follows:
Crosby, C. R. and Slingerland, M. V.: Manual of Fruit Insects,
1915-
Forbes, S. A.: Several Reports of the State Entomologist on the
Noxious and Beneficial Insects of the State of Illinois.
Harris, T. W.: Insects Injurious to Vegetation (several editions).
Insect Life, seven volumes (a mine of information on American
economic entomology).
Packard, Alpheus S.: Insects Injurious to Forest and Shade
Trees. Fifth Report of the United States Entomological Commission,

1890.
Riley, C. V.: Several Reports on the Noxious, Beneficial and other
Insects of the State of Missouri.
Saunders, Willi.^m: Insects Injurious to Fruits (several editions).
United States Bureau of Entomology: Popular and Technical
Bulletins on Insects.
CHAPTER XXIV
PLANTS AS DISEASE PRODUCERS, EPHIPHYTOTISM,
PROPHYLAXIS
Vegetal Agents of Disease. —The plants which are known to
be injurious to other plants fall naturally into two large groups, namely,
the Phanerogamic and the Cryptogamic. The latter includes injurious
algae, and fungi.
slime moulds, bacteria
The phanerogamic parasites belong to four families of plants.
Their morphology and physiology is fairly well known, so that in their
discussion, we are entering well-trodden fields of investigation.
The flowering plants, which lead a partially or wholly dependent
life upon a host plant, may be considered as belonging to two distinct
groups: the green parasites and the chlorophylless parasites. The
plants of the first groupby gradations how the conditions of
illustrate
life of the second group have arisen. The seeds of the first series of
green parasites begin their growth in the soil and there develop into
seedlings with cotyledons and root system, without any connection
with a host plant. The root branches supplied with suckers then
become attached to the roots or underground stems of other plants.
About one hundred plants of the sandalwood family, Santalace^,
belong to this series, including the true sandalwood, Santalum album
of India, where its roots live attached to the roots of a species of Acacia
leucophcBa and Pride of India, Melia azidarachta^.
The bastard toad-flax of Europe, Thesium alpinum, is another
member of this family. It develops relatively large suckers, which
become attached to the roots of other plants. These suckers are con-
stricted near their point of insertion. The swollen part spreads itself
over the root of the host as a plastic mass, while the central cores per-
forate the root and grow into the wood of the host where they spread
out. Comandra umbellata is a santalaceous parasite found in the pine-
" Wilson, C. C: Sandalwood. Indian Forester, xli: 248, August, 1915.
298
PLANTS AS DISEASE PRODUCERS 299
barren region of New Jersey. The family Scrophulariace^ includes

a number Such are the eyebright {Euphra-
of these root parasites.
sia), yellow-rattle {Rhinanthus), cow- wheat (Melampyrum), lousewort
(Pedicularis) and others. The suckers of the yellow-rattle are of

considerable size their margins are swollen and they spread around the
:
roots of the hosts. Those of the cow-wheat resemble in general those

of the yellow-rattle. In America species of Agalinis (old genus Gerardia
in part) are known to have parasitic attachments to the roots of various
plants. This plant is a member of the family Rhinanthacece (Scroph-
ULARiACE^, tribe Rhinanthae).
The second series comprises the chlorophylless root parasites, such
as Lathrcea squamaria, the toothwort. The young seedling lives at
first upon the reserve substances of its seed, sending out roots in all
directions. These finally fasten to the roots of ash, hornbeam or

poplar, by means of a sticky sucker, which develops a central core
that penetrates into the roots of its host. Colorless shoots covered
with whitish scale leaves are formed and the flowering shoot which
develops above ground has a purphsh hue.
The third series of parasitic flowering plants includes those of the
families Orobanchace^, Balanophorace^ and Hydnorace^. One
genus, Orobanche, the broom-rape genus, is sufficiently common to merit
attention (Fig. 117). The embryo of Orobanche shows no trace of root
and stem and is without cotyledons. It is a spiral filament of delicate
cells feeding on the stored reserve food. In its downward growth, its tip
traces a spiral line until it finds the roots of a congenial host, when it
not only adheres firmly to a root, but swells in such a way as to assume
a flask-shaped appearance. The thickened part becomes nodulated
and papillose and some of the papillae form conic pegs, which penetrate
into the root of the host until the vessels of the parasitic attachment
of the broom rape reach the A bud is formed at
vessels of the host.
and parasite and a strong thick flower-
the point of union between host
bearing stem grows above ground. Closely and intimately associated
with a host, such as a clover plant, the broom-rape does considerable
damage. Conopholis americana ,(Fig. 118) and C. mexicana live as
parasites on oak roots, developing large swelhngs out of which the
flowering shoots grow.
The writer collected Conopholis mexicana in 1896 on the roots of
an oak, Quercus reticulata, on the mountains at Eslava (10,000 feet)
Fig. 117. — Broom-i-apu (Orolnniche minor) upon greenhouse geranium. {After

Halslcd, B. D.. Rep. N. J. Agric. Exper. Slat., 1905.)
above the Valley of Mexico. Cf. Wilson, Lucy L. W., Observations

on Conopholis americana. Cont. Bot.-Lab., Univ. of Pa., II: 3-19.
The fourth series of phanerogamic parasites comprises plants of
the family Rafflesiace^, to which a number of genera belong. Raf-
Hesia is a genus confined to the islands off southeastern Asia, Java,
Borneo, Sumatra and Philippines. The whole plant is reduced to a
Fig. 118.— Cancer-root, Conopholis americana of the broom-rope family, Oroban-

chea; parasitic on roots of other plants. {From Gager, after Elsie M. Kiltredge.)
gigantic ill-smeUing flower, one meter across, with parasitic attach-

ments suggesting fungous hyphae, which penetrate the roots of vines
of the genus Cissiis. Brugmansia and Cytinus are two other genera
of this family. Cytinus hypocistus lives on the roots of shrubs of the
genus Cistus in Mediterranean Europe.
The fifth series of parasitic phanerogams includes epiphytes of
bushy habit belonging to the family Loranthace^. The genera
Fig. 119. — Distorted branch of mulberry caused by mistletoe {Phoradendron

flavescens), Austin, Texas. {After York. H. H., Bull. 120. Univ. of Tex., pi. ix,
March 15, 1909O
Loranthns, Phoradendron and Viscum include the well-known mistletoes.

The American mistletoe, Phoradendron flavescens (Fig. 119), extends
from southern New Jersey, Maryland, Ohio, Indiana and Missouri
to Texas. It is a slow-growing green parasite, which on account of its
chlorophyll is not entirely dependent upon its host for its carbohydrates
(Figs. 1 20 and 121). It is essentially a water parasite, and consequently,
its parasitic roots or sinkers grow into the woody cylinder of its host,
Fig. 120. — Cross-section of a live oak branch showing five stems of mistletoe
parasitic upon it. Note sinkers on parasitic roots penetrating into oakwood. {From
Gager.)
where they spread out circumferentially (Figs. 120 and 121). The
white berries, which are sticky, are carried by birds as the sticky
mass containing the seeds adheres to the bill and is only removed
by rubbing the beak against the bark of a tree, for example.
Mistletoe does not kill the trees directly, but it often causes them to
become very much dwarfed and their branches distorted greatly.
Parts of trees, however, may be killed. The larch mistletoe, Razou-

^
mofskya Douglasii laricis, is one which lives on the western larch in

Idaho and Oregon and in the open places interferes seriously with the
development of some of the more valuable timber trees.
The sixth series includes the climbing parasites, which are destitute
Fig. 121. —Sectional view, partly diagrammatic, of a branch infected with

mistletoe, showing relation of parasite and host, a, branch of host tree; b, mistletoe;
c, primary sucker; d, sucker from cortical root; e
f, cortex; g, cambium; h, wood
of branch. (After Bray, W. L., Bull. i66, U.S. Bureau of Plant Industry, Feb. 2, 1910.)
1 The student should consult the following for more detailed information about
mistletoe. Sorauer, Dr. Paul: Handbuch der Pflanzenkrankheiten (2d edition,
1886, ii: 25-32; Peirce, George J.: The Dissemination and Germmationoi Arceidho-
lium occidentalis. Annals of Botany, xix:99-ii3, January, 1905; York, Harlan H.:
The Anatomy and some of the Biological Aspects of the American Mistletoe.
Bull. Univ. of Texas, Scientific Series 13, March 15, 1909; Meinecke, E. P.: Parasit-
ism of Phoradendron juniperinum, Proc. Soc. Amer. Foresters, vii: 35-41, March,
1912; Mistletoe Pest in the Southwest, Bull. 166, Bureau of Plant Industry;
Weir, James R.: Larch Mistletoe, do. Bull. 317.
of chlorophyll and whose seeds sprout in the soil and send up a filiform
stem which brings itself by its movements into contact with some
host plant, which is penetrated by parasitic roots which enter, as
far as the bast regionand extract elaborated food. When estabhshed

on the host the parasite severs its soil connection. Leaves- have been
Fig. 122. — Dodder (Cuscuta) in flower and parasitic on a golden rod, Solidago ultni-
folia. {From Gager, after Elsie M. KlUredge.)
reduced to a few scales located near the clusters of small flowers and the
twining stem assumes a yellow, or orange-yellow color. The dodder,
Cuscuta (Figs. 122 and 123), belonging to the bindweed family, is
illustrative of these parasites.

Related in habit are species of the genus Cassytha. Most of the
species of Cassytha inhabit Australia, but some are found in New
Zealand, Borneo, Java, Ceylon, the Philippines, the Moluccas, South
3o6 GENERAL PLANT PATHOLOGY
Africa, the West Indies and Florida. In Florida/ Cassytha filiformis

is abundant on the dunes and in the rosemary scrub, where it spins its
yellow, or reddish-orange stems from bush to bush.
Fungous Organisms as the Cause of Disease.— The first part
of this book dealt with the morphology, physiology, and taxonomy, of
Fig. 123. — Photomicrograph of the section of a dicotyledonous host plant para-

sitized by dodder, Cuscula sp. At D
and Z>' note haustoria entering host plant as
far as the bast region of the stem. (After Gager).
the slime moulds, bacteria and true fungi. General reference was made
to the diseases induced by them and in the third part will be given an
1 Harshberger, John W. : The Vegetation of South Florida. Trans. Wagner
Free Inst, of Science, October, 1914; 86; Cf. Boewig, Harriet: The
vii, part 3,
Histology and Development of Cassytha filiformis. Cont. Bot. Lab., Univ. of

Penna., ii: 399-416, 1904.
account of the fungi which cause specific diseases. It remains for this
discussion to consider fungi as the causes of diseases in general. Fungi,
using the word in the broadest sense to include the bacteria and slime
moulds, are responsible for an extraordinary number of diseases. The
entrance of the organism into another is known as infection. Nothing
like the infection of animals where the microbe, or its poison, circulates
in the blood, and lodgment in most of the organs is found with
finds
plants. Infection follows, when a fungous spore germinates and pro-
duces an infecting hyphae, which grows into the cells^ or between the
cells of the host, it may be reaching to the ends of the plant. As disease
is induced by parasitic fungi, the parasite which enters the host and
spreads through it must absorb and utiHze the plastic and other sub-
stances of the plant, which is Thus, we can divide the
parasitized.
endophytic hyphae into the intercellular hyphse such as we find in the
oomycetous fungi and Puccinia simplex. With such hyphae ^he
protoplasmic and other contents of cells are utilized by the formation
of haustoria of different forms and kinds, which penetrate the interior
of the cells. The second kind are the intracellular hyphae, which as in
the disease of the plane tree, Gnomonia veneta, grow lengthwise and
crosswise from cell to cell.
The growth of the hyphae between and through the host cells is
accompanied by the formation of soluble ferments. These dissolve the
substance of the cell walls of cellulose, or woody walls with lignin and
pigment deposits. The hyphae live on the products of solution.^
Hence timber may be damaged in two ways: by the formation of minute
pores and apertures through it or by a solution of the cell- wall materials.
;
The wood loses in strength and in weight and becomes "rotten."

This rotten condition, however, is reached in a multiplicity of ways, for
every parasitic fungus that lives in the wood of growing trees destroys
the wood in a manner peculiar to itself. Starch grains are decomposed
and tannin, for by the disappearance
also in the cells, likewise crystals
sound wood is lost. Hartig has described
of the latter, the smell of
the several methods in his ''Text-book on the Diseases of Trees."
Then too, we have the epiphytic fungi which live on the surface
^ Sometimes the h3^hae grow toward and surround the nucleus as the nucleus
exerts a chemotactic influence. Such hyphae may be termed nucleotropic as in
Puccinia adoxce.
^ Consult Smith, Erwin F.: Bacteria in Relation to Plant Diseases, ii: 76-89.
of the host, as with the common mildews, and send short haustoria
into the epidermal cells of the host on which they grow. Some fungi
have mycelial hyphae that grow both ways, intracellularly and inter-
in
cellularly. Others, as a number of wood-destroying fungi, grow down
through the tissue of the host and ultimately kill it. Apical growth
is shown by some. The haustoria, as they enter a cell, may flatten out
against the cell wall, as in Piptocephalis. Such flattenings are known
as appressoria. The haustorium, which enters a cell, may become
branched, or dendritic, it may enlarge into a haustorial knob, or re-
main an haustorial tube. Internal sclerotia are formed sometimes
as
in certain parasitic fungi. These are consolidated or hardened masses
of hyphae, which are associated with a resting period.
Ordinarily when a spore falls on the surface of the plant, it produces
a germ tube, which by the action of a secreted ferment bores its way
through the epidermal and thus enters the host. Sometimes
cell walls
it penetrates the cuticle, grows between it and the cell wall and grows
down between the membranes of the cells, as in Botrytis parasitica.

Occasionally, but not commonly, it enters through the stomata, or
sometimes through nectaries and stigmatic surfaces. However, there
are certain bacteria, such as those which cause the black rot of the
cabbage, which fall upon the drops of water excreted by water stomata
and by following the water back into the plant infect the cabbage
leaves. A cork layer is protection against infection. Fungi, however,
gain access to the interior of the plant in a variety of ways. Some
years ago^ the writer considered the way which fungi enter living
in
trees and a restatement of the facts presented in that paper is
apropos.
Occasionally the planted seed contains a dormant fungus (but not
as a mycoplasm in Eriksson's sense), which begins its growth, as soon
as the seedling plant emerges. The oat- or wheat-smut spores are
produced in the grain and consequently infect the cereal plant when
it is small, and at or near the surface of the ground. In other cases the
fungus penetrates the underground parts or the twigs of trees. Fungi
gain entrance to plants, through injuries caused by mechanic, meteoro-
logic, chemic, or other agents. Mechanic injuries are due to man,
animals, or other causes, such as the weight of snow, the rubbing of
1 Harshberger, John W.: How Fungi Gain Entrance to Living Trees. Forest
Leaves, viii: 88-90, December, 1901.
PLANTS AS DISEASE PKODUCERS 309
two branches together. Squirrels in search of food bite off the twigs
of trees. Deer and moose browse upon the tender branches and bark
of various trees, the moose especially upon Acer pennsylvanicum and
Sorbus americana. Grizzly bears rub their backs against the bark of
trees and sometimes in this way decorticate them. Rodents peel off
the outer protective layers of roots as food, or as material with which to
line their burrows. The mycelia of Rhizocionia, or the oak-root fungus.
Fig. 124. — Street tree injured by use as a hitching post. ( I//1 \V. C, Rep.
Conn. Agi-ic. Expcr. Stal., pi. iii, igou )
RoseUinia quercina, which live in the soil, penetrate into roots through
wounds produced by field mice and gophers. The honey agaric,
Armillaria niellea, forms strands of hyphae known as rhizomorphs,
which grow through the soil and find an easy entrance into roots
decorticated by rodents. Beavers are active agents in cutting down
trees and removing the bark therefrom. Woodpeckers drill holes into
trees and in their case it has been definitely proved that they carry the
viable summer spores of the chestnut-bHgtht fungus, Endothio para-
3IO GENERAL PLANT PATHOLOGY
sitica, a single downy woodpecker carrying 757,074 spores.^ Wood-

boring insects (Family Scolytid^) of the genera Dendroctonus,
Scolytus,Tomicus are responsible agents in the destruction of trees

opening up holes through which fungi may gain entrance. Horses
do considerable damage to trees by stripping off the bark with their
teeth, and street trees cannot be too
soon or too carefully protected from
such ravages, for a tuHp tree planted in
the afternoon in front of the house of
the writer in West Philadelphia had a
strip of its bark removed by the curb-
stone horse of a delivery wagon before
nightfall of the same day (Fig. 124).
Telegraph wires stretched in every
direction rub against the trunks and
limbs of trees, and do mechanic injury
in this way, but, if the insulation is
rubbed off the tree may be badly burned,
or even set on fire by the electric cur-
rent, especially on rainy days when
there is a direct grounding of the cur-
rent through the water running down
the crevices of the bark. Many trees
in our cities are planted too close to the
curb and the wheels of passing wagons
Fig. 125. —Decay following un- tear off pieces of bark (Fig. 141).
skillful pruning. {Slurgis, W. C. Farmers in plowing, hoeing, mowing
Rep. Conn. Agric. Ex per. Stat., pi
Hi, 1900.)
and cultivating the soil injure the
and stems of cultivated plants
roots
and open the way for the entrance of destructive fungi. The blazing
of treesby surveyors, the careless system of lumbering, careless trans-
planting of young trees, are fruitful sources of injury to trees. Careless
pruning (Figs. 125 and 126) of trees by inexperienced men, such as was
prevalent in Philadelphia before the Park Commission undertook to
properly care for the trees, caused the death of many fine shade trees.
1 Heald, F. D. and Studhalter, R. A.: Preliminary Note on Birds as Carriers

of the Chestnut Blight Fungus. Science, new ser., xxxviii: 278-280, Aug. 22, 1913.
Stubs were left which never healed over and through the exposed sur-
face the fungi of wood decay gained easy access.
The produced by meteorologic causes are important.
injuries
Entire forests have been levelled by tornadoes. Cracks are produced
by wind action. Lightning opens a way by cracks to the interior.
Snow and ice snap off large limbs and hail stones bruise the bark and
leaves of trees so that fungi can readily enter. Chemic substances are
rather exceptional destructive agents to which reference has been called
26. — Black walnut, Juglans nigra. Cold Spring Harbor, L. I. Note large
open-branch stub (July, 1914).
in a previous page. Besides these agents, it occasionally happens, that

fungi enter healthy plants through diseased grafts which are inserted.
Robert Hartig mentions such a graft union of diseased and healthy
roots in the case of the red-rot fungus, Trameies radiciperda. Here
contact of the diseased root containing the fungus with the sound one
of a neighboring tree and the partial natural graft union of these two
roots explains how such infection occurs. An enumeration of the
way in which fungi can gain entrance to plants follows:
A. By means of spores, or h>i)luc, into stoinata and

water stomata.
Infection by natural B. By rerment action of a fungus on the epidermis of
growth of the fungus the host.
f By developing from a dormant state in the seed into
[
an active state in the seedling.
[
Beasts
I. Mechanic injuries )
Man
induced by Fall of fruit
Combined weight action of fruit
Wind
Snow
Ice
II. Meteorologic in-
Hail
juries induced by
Lightning
Sun
Infection through Frost
Factory gases
Sewer gases
III. Chemic injuries Locomotive gases
induced by Chemicals at roots.
Alkali soils
Gases and chemicals in geysers, etc.
IV. Non-classifiable
Natural grafting and budding
injuries induced by
Incubation.^ —The period of incubation is the time between ex-

posure to the cause of the disease and the first appearance of the symp-
toms, or physical signs of the disease. This period in plants is quite as
variable as in animals, and
dependent on the nature of the organ-
it is
ism, whether it is virulent, or its virulency attenuated, on its food re-

quirements, on its temperature requirements, the volume of infectious
material, the stage of development, or age of the host plant, the amount
of water and air in the invaded tissues, and individual or varietal re-
sistance. The period of incubation may be as short as a few hours,
or as long as three to four weeks. Presumably on seedling tissues the
period of incubation of the damping-off fungus, Pythium de Baryanum,
is only a few hours. Experiments performed by Erwin F. Smith^
1 Smith, Erwin F.: Bacteria in Relation to Plant Diseases, ii: 66.
with Bacillus trachciphUus and young cucumbers where the organ-

ism was inoculated from young cultures, and on susceptible plants by
needle-pricks, showed that signs of disease rarely appeared in less than
three to four days, and that signs of wilt and change of color usually
were visible in five to seven days. In the case of the white pine bhster
rust, Cronartium ribicola, the period of incubation in the pine is from
one to six years.
Duration of Disease. —The resistance of plants to disease is various
even after the fungus has obtained an entrance into the tissue of the
Fig. 127. — Chestnut, Caslanea denlaia, killed by blight fungus, lindolhia payascaili,
Cold Spriiig Harbor, L. I., July, 1914.
host. In the case of large trees like the white oak, a number of years
may elapse before the tree finally succumbs to such fungi, as Fomes
{Poly poms) applanatus. A chestnut tree, a few miles outside of
Philadelphia resisted the chestnut-blight disease for over four years
from the time of first infection before it finally succumbed. Smith
{loc. cit.) describes how a good-sized potato tuber was half rotted in
five days at ordinary autumn temperatures when inoculated with
Bacillus phytophthorus by means of a few needle-pricks.
The final outcome of the disease may be a complete destruction of

the host (Fig. 127), or its complete recovery. The simplest cases are leaf
spots, or fruit spots,which are removed from the plant when the leaves
and fruits fall without in any way jeopardizing the general health of the
plant. Sometimes the plant recovers from bacterial, or fungal diseases,
but such recovery does not protect the plant from subsequent attacks
of the same disease, as is the case with some diseases of animals. Old
and slow-growing cabbages are rather resistant to Pseudomonas cam-
pestris while young and rapidly growing plants are apt to be destroyed.
Vaccination of plants to ward off diseases has never been successful,
and it is doubtful whether this means of protection is available for
plants. It is, however, a wholly unworked field. Some experiments
which Smith, Townsend and Brown performed in 1908 and 1909 seem
to show that after Paris daisies have been inoculated several times with
Pseudomonas tumefaciens with the production of tumors, that subse-
quent inoculations with cultures of the same virulence are without
effect, but owing to the possibility that the results were due to loss of
virulence, the experiments were inconclusive. For the student, who

may be interested in pursuing this line of important research work
further, the following bibliography is here given, taken from Smith.
Shattock, Samuel G.: The Healing of Incisions in Vegetable Tissues. Journ.

Path, and Bact. Edinburgh and London, v: 39-58, 1898.
HiLTNER, L. and Stormer, K.: Neue Untersuchungen iiber die WurzelknoUchen
der Legurainosen und deren Erreger. Arb. a.d. Biologischen Abt. fur Land-
und Forstwirthschaft am Kaiser. Gesundheitsamte iii, heft 3: 151, 1903.
Brullowa, J. P.: Ueber den Selbstschutz der Pflanzenzelle gegen Pilzinfektion.
Jahrb. f. Pflz. Krh. K. Bot. Garten Petersb., Nr. 4, 1907.
Alten, H. von: Zur Thyllenfrage. Callusartige Wucherungen in verlezten
Blattstielen von Nuphar luteum. Bot. Ztg., 68, part ii: 89-95, 1910.
Smith, Erwin F.: Bacteria in Relation to Plant Diseases, ii: 93-94, 1914.
DISSEMINATION OF FUNGI
Fungi are usually reproduced by spores, which are minute and light
and easily carried about by various agents, such as on seeds, by the wind,
by water, by insects, by other animals, by agricultural and commer-
cial practices and by railroads, cars and other vehicles. The black-leg,
or Phoma wilt of cabbage of recent introduction, was introduced from
Europe undoubtedly with imported seed, and as we have seen various
smuts are carried by the single fruits of various grains. In the aecial
stage of the cedar-apple fungus, Gymno sporangium juniperi-virginiance,

the spores are set free during dry weather at a time when they are most
likely to be wind-carried.^ The spores of the water molds are carried
by currents of water and those of the cranberry gall due to Synchy-
trium vaccinii. The motile zoospores of the damping-off fungus need
water for their dissemination. The spores developed during the
Sphacelia stage of the ergot fungus on rye are carried by insects. The
formation of the conidiospores is accompanied by a sweet substance,
the so-called honey-dew, which is much relished. Birds, especially
woodpeckers, disseminate the spores of the chestnut-blight fungus,
Endothia parasitica, and in a great many different ways man is active.
EPIPHYTOTISMS (EPIDEMICS)
When a plant disease becomes virulent, rampant and aggressive,
spreading rapidly from place to place, it is said to be epiphytotic
(epidemic). A number such epiphytotisms (epidemics) have oc-
of
curred and the destruction due to some particular plant disease has
been enormous. The potato crop in the British Isles during the
summer of 1845, owing to a high temperature and abundant rains,
suffered entire destruction in the short space of a fortnight. This was
due to the ravages of Phytophthora infestans, an oomycetous fungus,
whose spores in wet weather produce numerous infecting motile
zoospores. The destruction of the potato crop led to the repeal of the
corn laws of England, and as a sequence, the inauguration of a free trade
policy. The Irish famine was the direct result and thousands of the
natives of the Emerald Isle emigrated to America. With respect to the
disease known as peach yellows Dr. Erwin E. Smith writing in 1891^
says: "Formerly was confined to a small district on the At-
this disease
lantic Coast, but during the
last twenty years it has invaded distant
regions hitherto free, and has entirely ruined the peach industry over
very considerable areas. Within ten years the disease has taken fresh
1 Heald, V. D.: The Disseminations of Fungi Causing Disease. Trans.

American Microscopical Society, xxxiii: 5-29, June, 1913.
2 Smith, Erwin F. Additional Evidence on the Communicability of Peach
:
Yellows and Peach Rosette, Bull, i, Div. of Vegetable Pathology, U. S. Dept.

Agric, 1891.
3l6 GENERAL PLANT PATHOLOGY
very strong hold upon the orchards Delaware and Chesapeake and
in the
and has destroyed thousands

region, the north portion of the peninsula,
and thousands of trees, rendering a great industry unprofitable and
precarious." The recent spread and virulency of the chestnut-blight
fungus, Endothia parasitica, from the neighborhood of New York City,
where it was probably first introduced, is so recent and fresh in the
minds of the public, that an extended account of the epiphytotism
(epidemic) need hardly be made here. The disease has practically
destroyed the native chestnut trees of the forested areas of the east-
ern states east of a line running northeast and southwest through
the central part of Pennsylvania . There have been a few sporadic cases
west of that line removed through the heroic efforts of the men em-
ployed by the Pennsylvania Chestnut Blight Commission, who with a
big appropriation of state money tried to find a way of heading off the
disease and finally controlling it but without success. Introduced in all
probability from China, where it has been found recently, the ravages
of this disease have been without precedent.
As to the epiphytotic diseases of plants due to animals, we have a
number of instructive illustrations. The account of the introduction,
spread and final control of the cottony cushion scale forms one of the
most interesting chapters in the history of American phytopathology.
Having been introduced from Australia to California in 1868, it
spread so rapidly during the next twenty years that its ravages proved
a very serious menace to the citrus industry of the southern part of
California. The Australian ladybird beetle, which was introduced
into California from Australia in 1889 for the purpose of controlling
this scale, was so successful, that except for occasional outbreaks it
ceased to be considered a serious citrus pest.

All of these epiphytotisms (epidemics) and others that might be
cited have been possible in all probability because the climatic condi-
tions of temperature, moisture, rainfall, wind and soil conditions have
been favorable during the period of most active virulency, when the
diseases became firmly established. As an important contributing
cause may be considered the unhealthy, abnormal, or susceptible condi-
tion of the host plant owing to the methods of cultivation which have
reduced the disease-resisting capacity of the plant. In the case of
the chestnut, the restoration of the trees by sprouting from the stump
was undoubtedly one of the contributing causes of the rapid spread of
the disease. Altogether, these epiphytotisms (epidemics) result either

when the conditions are favorable for the spread of the parasites, or
when the general tone and health of the plant has been lowered by
improper methods of handling, so that its disease-resisting capacity
has been reduced. Recognizing the possibility of the introduction of
other virulent fungous, or animal diseases, a stricter quarantine has
been instituted by both the individual state and national governments
with a careful inspection of nursery stock designed for shipment from
place to place.
PROPHYLAXIS
Prophylaxis may be defined as the means taken to prevent disease.
It includes a consideration of the methods of protecting plants from
disease, of preventing the spread of disease, and of the methods of
breeding by which the disease resistance of plants is increased until in
some cases absolute immunity is reached and the plant is made proof
against disease. Some diseases are preventible by the observance of
proper care in the cultivation of plants,^ and by habits of cleanliness,
when no refuse which might harbor insect or fungous disease is per-
mitted to remain, but is either destroyed, or rendered innocuous.
For example, vegetable and agricultural crops should be rotated, so
that the same crop would not follow upon the same piece of soil where
the animal or fungous parasite may be lurking. Neither should the
farmer attempt to cultivate certain crops in acid soils, or in low situa-
tions subject to frost action. Nor should seeds be placed in beds rife
with the spores of the damping-off fungus, Pythium de Baryamim. By
proper care on the part of the grower diseased plants should not be
sentaway from an infected locality, and vice versa, he should be careful
about the introduction of nursery stock and plants from other localities
without a careful inspection. The national and state quarantine
regulations are designed to help the grower in these respects, and he can
refuse to purchase new plants without they are accompanied by a
certificate setting forth that these plants are free from animal and
fungous diseases. Orton- in two suggestive papers, has shown that
^ BoLLEY, H. L. Cereal Cropping: Sanitation, a New Basis for Crop Rotation,
:
Manuring Tillage and Seed Selection. Science, xxxvii: 249-250, Aug. 22, 1913.
^
Orton, W. A.: International Phytopathology and Quarantine Regulation,
Phytopathology, 3: 143-151, June, 1913. The Biological Basis of International
Phytopathology, Phytopathology, 3: 325-333, February, 1914.
3l8 GENERAL PLANT PATHOLOGY
this problem is not only of national, but of international and inter-

continental importance. These papers should be read by every
serious-minded student.
Plant protection may be secured by the use of spraying materials.^
The principal rules to be observed in their use are: (i) the poison em-
ployed must be sufficiently strong or concentrated to kill the parasite,
but not sufficiently powerful to injure the host; (2) it must be applied at
the right time, as suggested by a knowledge of the life history of the
fungus, or insect in question. Such sprays may, therefore, be divided

into two kinds, and fungicides. Applications of these
viz., insecticides
to healthy plants serve to protect the plant from the attacks of its
fungous and insect enemies. Vast possibilities of controlling disease
have been opened up by the treatment of seeds with hot water and other
substances before the seeds are planted.
iMcCuE, C. A.: Plant Protection. Bull. 97, Del. Coll. Agric. Exper. Stat.
June IS, 1912; Rees, Charles C. and Macfarlane, Wallace: A Bibliography of
Recent Literature Concerning Plant Disease Prevention. Univ. of 111.: Agric.
Exper. Stat., Circular 183, May, 1915.
CHAPTER XXV
PRACTICAL TREE SURGERY^
The object of tree surgery is to repair the damage done to trees by
the various causes previously described (page 274). The principles
involved in all such remedial work are the removal of all decayed, dis-
eased, or injured wood and bark, the cauterization, sterilization, and
waterproofing of the cleaned, or cut, surfaces, and the putting of the
treein a condition for rapid healing. Such treatment should be
watched from year to year, so that any defects will receive immediate
attention.
As the work requires the apphcation of scientific principles, no
ignorant laborers should be employed. The men who act as tree sur-
geons should have some knowledge of the structure of trees, their
physiology and their habits of growth. A knowledge of the general
would not come in amiss, such as
principles of horticultural practice
the tenets of grafting and pruning. Such workmen would be still
better prepared, if acquainted with the structure, growth and life
histories of the common destructive fungi and insects. If a town or
municipality is unable to obtain such skilled labor, then the appoint-

ment of a superintendent, or town forester, who is acquainted with such
matters, should be made. Such a man should know the right thing
to be done and all the details of the work.
Preventive Measures. —As means Oi" preventing injuries to trees,
various things may be done. The placing of an open tree box or fence
of iron, or wire netting, is important, because it protects the tree from
the gnawing of horses and the rubbing action of passing vehicles, or the
viciousness of street arabs. Proper attention to the insulation of
telephone, telegraph and electric wires will prevent a lot of damage to
shade trees. Electric linemen, unless properly supervised, have no
' A by J. Franklin Collins will be found
detailed account of practical tree surgery
in the Yearbook of the United States Department of Agriculture, 1913; also con-
sult Stone, George E.: Shade Trees, Characteristics, Adaptation, Diseases and
Cure, Bull. 170 Mass. Agric. Exper. Stat., Sept., 191 6.
319
regard for shade trees, as they look upon them as obstacles to the
prosecution of their work. Improper pruning, when large stubs are
1
PRACTICAL TREE SURGERY 32
a rope and ladder are needed. The cuts should be made close to the
main tree trunk, so as to reduce the surface exposed to the action of
the elements. Cut surfaces should be cauterized and water-proofed.
The best antiseptic dressings are some of the creosotes, which destroy
and prevent the growth of wood-destroying fungi, because it penetrates
the wood better than a watery antiseptic. The antiseptic treatment
with creosote should be followed by painting the scar with coal-tar.
Lead paint sometimes more available. It is useful, but not as
is
satisfactory, as a heavy coat of coal-tar.

—
Cavity Treatment. The removal of all decayed and diseased parts
of the tree should be accomplished first by the use of gouges, chisels and
scraping tools. The use of the chisels is assisted by a wooden mallet.
These cutting instruments should have keen edges for the cambium
may be injured by dull tools. After properly clearing away all decayed
material, the freshly cut surfaces should be treated with creosote and
heavy coal-tar which should coat the surface of the sound and healthy
exposed surfaces of the wood. The excavation should be so made as
to provide drainage at the bottom of the cavity, but the undercutting
should be done in such a way as to hold the filling material. Before
the filling material is added to the cavity, it may be necessary to place
one or more bolts in position to hold the tree shell firmly together.
Iron rods and wire netting are also sometimes placed in the hollow to
help reinforce the concrete, or cement, when it is mixed and ready
for use. The tree surgeon learns by experience the best methods of
procedure in the use of bolts, wire netting and the placing of the filling
substance.
—
Mixing and Placing the Cement. A good grade of Portland cement
and clean, sharp sand free from loam (i part of cement to 3 or less of
sand) should be used. The mixing can be done in a mortar bin, a
wheelbarrow, a pail, or in any other available receptacle. A mason's
flattrowel and an ordinary garden trowel with a curved blade will be
found convenient in placing the cement. A tamping stick, one or two
inches thick and one to three feet long, according to the size of the cavity,
will be needed, also some rocks to help fill the cavity and a pail of
water. As the cement begins to harden, the surface should be carefully
smoothed, so that it conforms with the general contour of the tree trunk.
Sometimes cloth, or wire dams are used. These are stretched across
the opening and a more liquid cement is poured into the space behind
Fig. 131.— Cement cavity fillings, showing different types and successive stages.
I, Alarge cavity in an elm filled with cement blocks separated by layers of tarred
paper; a patented process. 2, An excavated cavity ready for treating and filling.
3, The cavity shown in 2, which has been nailed and partly filled with cement. The
ends of the rods for reinforcing the concrete are sprung into shallow holes in the wood.
The wire dam is sometimes allowed to remain embedded in the cement, though it
is usually removed as soon as the cement has partially set. 4, A later stage of the
work shown in 3. The height of the wire dam has been increased. 5, The same
cavity shown in 2, 3, and 4, several days after the filling was completed. (After
Collins, F. L., U. S. Yearbook Dept. Agric, 1913.)
the dam which is removed when the fiUing has hardened. Asphalt and
asphalt mixtures promise much for the future, when the proper methods
of applying liquid asphalt have been discovered (Fig. 131).
Defects in cement work are due to the use of cheap materials,

carelessness in the mixing of the cement, splitting of the tree by the
action of intense cold, dislodgment of the cement by the swaying action
of the wind. Cracks appear in the cement, if the wood of the tree
contracts away from the Ming, or by the spread of the decayed tissue
behind the cement work due to lack of care in excavating rotten wood
These defects may cause lots of trouble.
prior to the filling operation.
Metal-covered Cavities. —
Sheet tin, zinc and iron have been used
extensively to cover cavities. These coverings often serve to exclude
rain, fungous organisms and destructive insects for some time. If not
properly applied, such tin-covered cavities are a greater menace to the
tree than open cavities. If such covers are used at all, the excavated
cavity should be thoroughly sterilized and waterproofed. The metal
is nailed fast with a light hammer and its center should be allowed
to curve outward, so as to conform to the general shape of the tree
trunk. The tacked edges should be as nearly air-tight and water-
proof as it is possible to make them, and this can be assisted by paint-
ing the surface of the tin. Sometimes fumigation of the cavity is
resorted to as an added precautionary measure.
Where the tree is not of sufficient value to fill with cement, an open
cleaned cavity may be left after cauterization of the cleaned wood
surface and waterproofing. A layer of burned wood sometimes a
is
if the burning is
sufficient protective covering, accompHshed by one
of theblow lamps, such as painters use for stripping the paint off
woodwork.
Guying. — Closely associated with the work of tree surgery proper,
and often an indispensable adjunct is the guying of limbs to prevent
the spHtting of the crotches, or to check further splitting. Experience
demonstrates the best methods of applying the hook bolts, chains or
other braces to the trees to be treated. This varies so widely in dif-
ferent trees that it is impossible to give specific directions for this
kind of work.
In conclusion, it should be stated that tree surgery can be under-
taken safely at almost any season of the year, especially well when the
sap is not flowing actively, and the weather is not too cold, to freeze
the cement, and destroy such expensive filling work. Most ornamental
and shade trees having only a few dead limbs are unquestionably worth
attention. Others which have many dead limbs, or numerous decayed
areas may not be worth the expense. Trees of large size, rare trees,
historic trees and trees which fill a peculiar place in the landscape are
probably worth saving by the most expensive methods of tree surgery,
if necessary. Another phase of tree surgery is the commercial side,
where ignorant men and tree fakers have undertaken to make a business
of pruning and treating trees. The sad appearance of excessively
pruned trees in all of our large American cities are living spectacles of
the zeal of such men, who should be driven out of the business, as they
have in Philadelphia by the municipal authorities undertaking to do
the work by the employment of skilled tree surgeons.
Bailey, L. H.: The Pruning Book. The Macmillan Co., New York, 1907.
Blakeslee, Albert F. and Jarvis, Chester Deacon: Trees in Winter. Their
Study Planting Care and Identification. The Macmillan Co., New York, 1913.
Collins, J. Franklin: Practical Tree Surgery. Yearbook of the United States
Department of Agriculture, 1913: 163-190.
Gaskili, Alfred: The Planting and Care of Shade Trees.
Forest Park Reservation Commission of New Jersey, 19 1 2, with papers on Insects
Injurious to Shade Trees by John B. Smith and Diseases of Shade and Forest
Trees by Mel T. Cook.
Start, E. A. Stone, G. E., and Fernald, H. T.: Shade Trees. Bull. 125, Mass.
Agric. Exper. Sta., Oct. i, 1908.
It has been a matter of general knowledge that a disease may be

controlled by a change in the time of planting, for with smuts the very
different climatic conditions prevailing at the time of the various
sowings have influenced the rate of infection. Early sowing of winter
wheat has been found beneficial in the reduction of the amount of
stinking smut, for wheat sown early in October showed no sign of infec-
tion, while plants sown at the end of October were much attacked
(about 60 per cent.) by the smut. By experiment as a problem in
prophylaxis this matter of sowing as a means of controlling disease
should be established for all of our important cultivated crops.
Then too, a study of the cells and tissues which protect plants
against the entrance of insects and fungi is a matter of prophylactic
interest. The formation of cork, of bark, of callus, of how in response
to the attack of fungi, the multiplication of protecting, or outer cells,
is accomplished, should receive the attention of the student of phyto-
pathology. The presence of tannin and other protective chemical

substances in the plant may explain immunity or non-immunity.^
Disease resistance and disease susceptibility are understood imper-
fectly. The determination of the cause of the inherent differences in
the tendency of this or that variety to suffer from disease is a matter
of great importance. Breeding for disease resistance is a promising
field of research. 2 Something has been accomplished along this line,
but the amount which we do not know vastly exceeds the knowledge
which we now possess. Rustproof varieties of wheat have been ob-
tained. At the Ohio Experiment Station by selection of hills of
potatoes that withstood attacks of the early blight fungus and planting
tubers therefrom with subsequent repetition of this line of work, early
blight resistant strains were secured. Progress has been made with
cotton resistant to wilt and with musk melons resistant to leaf blight.
Recently Jones and Oilman^, Wisconsin, have undertaken to con-
trol the disease known as yellows caused by the parasitic soil fungus,
Fusarium conglutinans, by breeding cabbage plants that show disease
resistance. By repeated selection of the occasional sound heads in
fields of diseased cabbages, strains of winter cabbage of the Hollander
type have been secured which have proved in a high degree resistant
against the attacks of Fusarium. The chances for research along these
lines are practically unlimited and full of promise for the future of
agriculture and horticulture.
1 Cook, Mel T. and Taubenhaus, J. J. : The Relation of Parasitic Fungi to

the Contents of the Cells of the Host Plants, (i. The Toxicity of the Tannins)
Bull. 91, Del. Agric. Exper. Stat., February, 1911.

2 The Development of Farm Crops resistant to Disease. Year-
Orton, W. a. :
book United States Department of Agriculture, 1908: 453-464.

of the
3 Jones, L. R. and Oilman, The Control of Cabbage Yellows through
J. C. :
Disease Resistance. Research Bull. 38, Agric. Exper. Stat. Univ. Wis., December,
1915; Norton, J. B.: Methods used in Breeding Asparagus for Rust Resistance,
U. S. Bureau of Plant Industry, Bull. 263, 1913.
CHAPTER XXVI
INTERNAL CAUSES OF DISEASE
During recent years attention has been called to diseases which are
evidently due to the action of an enzyme, or ferment in the plant,
which renews itself perhaps as a catalytic agent in the tissues of the
host. As it is filterable through a Berkefeld filter, it may be a soluble
enzyme pure and simple, or it may be one of the extremely minute,
ultra-microscopic organisms to which attention has been called recently.
All the evidence seems to point to its enzymatic nature. Such diseases
are caused by the excessive activity of the oxidase and peroxidase
enzymes in the plant and the loss of function of catalase, another en-
zyme, which carries off some of the residual products of the others
mentioned. Such diseases due to a Contagiimi viviim fluidum affect a
number of plants, notably the tobacco, and all of these diseases seem
to be more or less related, as to their nature and origin. Recently
Kiister in the second edition of his "Pathological Plant Anatomy"
(191 6) has grouped many of the enzyme-produced conditions under
the head of "Panaschiering." He distinguishes several types. The
first is when the green parts contract sharply under the pale parts.
Under this head he considers: (a) marginal panaschiering, when such
terms as "albo-marginatis" would be applicable, as in such cultivated
plants as Pelargonium zonale, Hedera helix and Weigelia rosea, (b)
In sectional panaschiering, the white and the green colors are dis-
tributed sectionally over leaves and stems, as in Chamaecyparis pisi-
fera plumosa argentea. (c) He distinguishes marbled and pulverulent
panaschiering. His second group includes cases where the border

between green and pale parts is not sharply marked and this group
includes {a) Zebra-panaschiering, as in the banded leaves of Etdalia,
and (b) flecked panaschiering, where white specks are distributed over
a green background and blend with it. It is clear that "Mosaic,"
"Brindle," "Calico" or "Mottle Top" of tobacco is a physiologic,
not a fungous or bacterial disease.
326
INTERNAL CAUSES OF DISEASE 327
It is infectious, and to a certain extent contagious. As calico is an

important disease of tobacco and tomato a description of it in these
plants will serve to show what enzyme diseases are like in general.
The leaves present a mottled appearance, being divided into smaller, or
larger, areas of light-green and dark-green patches. In the tomato, the
light-green areas become yellowish, as the disease progresses, and in
very badly affected plants become finally purplish-red in color. The
leaves are much distorted, stiff, and badly curled. It attacks other
plants, notably the poke weed. Phytolacca decandra, ragweed, Am-
brosia Jamestown weed. Datura stramonium.
artemisicBfolia, It is
probable that peach "yellows," aster "yellows" are more or less similar
to the true "mosaic." Calico is primarily a disease of the green color-
ing matter (chlorophyll) of the infected plants; hence it disturbs the
normal nutrition of the plant. To this destruction of the chlorophyll
the name of chlorosis has been given and calico is, therefore, a state of
chlorosis. The contagious nature shown by experiments
of calico is
which prove that it can be communicated at least in some cases by

mere contact of calicoed plants with the healthy. Juice on the hands
from calicoed plants when handling disease-free plants will spread the
disease in nearly all cases, and this infection is due to the chlorotic juice
on the hands of the experimenter. Chlorosis, or calico, usually takes
ten to fourteen days to make
appearance after infection and a plant
its
once infected remains permanently so, and all new growth usually
becomes calicoed. Calico, or mosaic, can be transferred to other species
and varities of Nicotiana than the common N. iabacum, also to potato,
egg plant, peppers, petunia, etc. The dried leaves of calicoed tobacco
retain their power of infection for at least a year or two, to some degree,
but if wetted they lose this power. The virus, if it is permissible to
use this word, can be apparently extracted from calicoed leaves by
and alcohol without destroying its infectious qualities.
ether, chloroform
Bunzel has measured the oxidase content of plant juices, because of the
importance of oxidase in chlorotic diseases of plants, in their causal
relationship to color production in plants, their importance in the dark-
ening of tea and in the production of the smooth, black and hard
lacquer of the Japanese, from the white, fluid, soft secretion of the
lacquer tree, Rhus vernicifera. The
on oxidizing enzymes
literature
is a copious one. The following papers and books can be consulted,
as well as the bibliography which each includes:
BuNZEL, Herbert H. The Measurement of the Oxidase Content of Plant Juices.

:
Bull. 238, Bureau of Plant Industry, U. S. Dept. Agric., 191 2.

Chapman, G. H. Mosaic and Allied Diseases with Especial Reference to Tobacco
:
and Tomato. 2sth Annual Report Mass. Agric. Exper. Stat., 1913: 94-104.
Clinton, G. P.: Chlorosis of Plants with Special Reference to Calico of Tobacco.
Report Conn. Agric. Exper. Stat., New Havên, 1914: 357-424, with 8 plates.
Kastle, J. H. The Oxidases and other Oxygen Catalysts concerned in Biological
:
Oxidations. Bull. 59, U. S. Hygienic Lab., 1910.

Klebahn, Professor Dr. H.: Grundziige der Allgemeinen Phytopathologie,
191 2: 124-127.
Woods, Albert F.: Observations on the Mosaic Disease of Tobacco. Bull. 18,
Bureau of Plant Industry, U. S. Dept. Agric, 1902.
Nutritive disturbances may also be included as internal causes of

disease. any reason, such as the inability of the living cells of
If for
the root to take up water through a change in the osmotic power of the
protoplasmic membrane of the root hair cells, the leaves above owing
to active transpiration cannot secure sufficient quantities of water
and the whole plant wilts. A disturbance in the formation of starch
in the chloroplast results in a deficiency of the plastic carbohydrates,
and the active cells of the cambium during this period of starvation
form less wood and, therefore, fewer conducting vessels. This reacts
on the tissues everywhere in the plant by reducing the available water
and food and, therefore, the plant is dwarfed and perhaps sickly.
Intumescences are trichomatous outgrowths not associated with
insects or fungi which are due to some disturbance of the balance
between transpiration and assimilation.
Mutations which result in the sterility of an annual species would
lead to the extinction of the plant with such non-seed production.
(Enothera albida is a pale-green, rather brittle and very delicate form
with narrow leaves; never attaining anything like the height of (E.
Lamarckiana. It bears pale flowers and weak fruits which contain
little seed. It appears every year in most of de Vries's cultures in
larger or smaller numbers. The plants are so weak that de Vries
imagined them to be diseased,^ and after much difficulty he secured
seeds from them. Enough has been given on these points to show that
mutations may be along the line of plants constitutionally weak.
The absence of amygdalin and prussic acid in the Sweet Almond
may make such a form more susceptible to disease, as also the absence
of quinine from cinchona trees kept in European hot houses.
^DE Vries, Hugo: The Mutation Theory (English edition), I: 229, 1909.
internal causes of disease 329
Malformations and Monstrosities
Hugo shown that malformations and monstrosities

de Vries has
do not but may be looked upon as muta-
arise as a result of variations,
tions. His tricotylous, hemisyncotylous, syncotylous, and amphi-
syncotylous races are proof of this statement. Fasciation in its
simplest form consists of a flat, ribbon-like expansion of stem,

branch, flower clusters, flowers and fruits which may be cylindric
below, but flattened above. This is one of the most common of all
malformations and by numerous experimental cultures the fasciation
has been found to be heritable. Spirally twisted plants are more
striking than fasciations. Valeriana officinalis is
malformations
one best-known examples displaying spiral torsion. It is also
of the
displayed in a teasle. Dipsacus silvestris torsus, twisted sweet william,
Dianthiis barbatus, dark-eyed Viscaria, Viscaria oculata. Such mal-
formations de Vries has shown to be truly heritable. (Pleiphylly is
that condition where two or more leaves arise in place of a single one.}
Such we find in the ever-sporting races of clovers, where four, five,
six,seven, or even eight leaves appear instead of the normal three.
The presence of three leaves in a whorl, or of three cotyledons, as above
noted, is called polyphylly. Shull has shown that the ascidial
leaflets of the white ash, Fraxinus americanus, are heritable. Pistil-
lody is demonstrated in the appearance of imperfect pistils in place of
stamens, as in the poppy. When become green,
colored flower parts
this condition is known and is illustrated
as antholysis, or chloranthy,
in green roses and green dahlias. This condition and petalody and
sepalody are transmitted. Peloria, where a normally zygomorphic
flower, as in the toad-flax, Linaria vulgaris, is transformed into a regular
flower with five spurred petals instead of one spurred petal, is
another example of monstrosities which are heritable.

The history of Cytisus Adami which originated as a graft hybrid
is of interest in connection with the study of Chimaeras. Hybrids that
arise by vegetative reproduction, where scion and stock are mutually
affected, are known as graft hybrids. The origin of Cytisus Adami
seems to have been as follows: a shoot of Cytisus purpureus was
grafted on a stock of Cytisus laburnum; from this were produced many
shoots, one of which grew vigorously, and developed larger leaves
than those of C. purpurcus and from this shoot plants were propagated
constituting Cytisus Adami. It was found, that on flowering, this

form had dingy red flowers. Winkler believes that graft hybrids and
chimaeras are the result of the process by which cells of two distinct
kinds or species are united vegetatively instead of by sexual methods,
and that this serves as the point of departure for an organism which in
a single growth shows bound together the peculiarities of both species.
Hence, a graft hybrid is a complex chimgera. Baur thinks that the
union between CratcBgus and Mespilus {Crafa gomes pilus) is a periclinal
chimgera, and refers this and the graft hybrid to the development of a
mixed vegetation point, where the periclinal chimaera originates in
the development of an apical region with a periclinal arrangement
of cells. ^
Branches of shrubs and trees originate as mutants with a dififerent
combination than the rest of the shrub, or trees. Such

of characters
mutants probably arise in the change of some single cell. The shoot
which arises from tissue formed by mutating cells develops into
something new which is called a bud variation, or sport variety. If
the shoot arises from the mutating cells alone, then the resulting
shoot will consist only of the new cells an^ the sport can be propagated
true without any reversion. If the tissue which gives rise to the shoot
combines both old and new cells, then there arises a mixed branch,
which is known as a "sectorial chimaera." Citrus treess how such
"sectorial chimaeras" not infrequently when a Valencia orange tree
bears typical Valencia oranges and a small rough and worthless muta-
tion. A twig here and there produces oranges in which certain sectors
of the fruits show mutant tissue," forming what may be called mixed
oranges. These have probably arisen because the mutant tissue is
scattered or mixed with the tissue of the original form thus constituting
a "hyper chimaera."
"Mutations often occur in the cells which begin the formation of
the minute ovaries in the blossom buds. As the ovary grows in size,
the mutation appears as a sector of the fruit which differs in color,
ripening season, or thickness of skin from the rest of the fruit. Such
curious fruits have been called spontaneous chimaeras" (Coit).
1 Winkler, H.: Ueber Pfropfbastarde und Pflanzliche Chimaren. Ber.

Deutsch. Bot. Gesellsch., 25: 568-576, 1907; Baur, E.: Pfropfbastarde, Periklinal
chimaren und Hyperchimaren, Do., 27: 603-605, 1909.
2 Coit,
J. Eliot: Citrus Fruits, 1915:
121-122.
CHAPTER XXVII
CLASSIFICATION OF PLANT ABNORMALITIES
The older botanists prior to the pubHcation of the important work
of Maxwell T. Masters in 1869 gave little attention to abnormalities in
plants. Linnaeus treated of them to some extent in his ''Philosophia,"
but mainly to Augustin Pyramus de Candolle that the credit is
it is
due of calling attention to the importance of vegetable teratology, as

throwing light upon normal structure and functions. Until the epoch-
making work of de Vries on plant mutations drew attention to the
absolute necessity of experimental methods in the study of normal and
teratologic plants, the field of vegetable teratology was the concern of
the plant morphologist and the different abnormalities were studied by
comparative morphologic methods. Hugo de Vries and several of
his co-workers pointed out that many abnormal forms are heritable and
this suggested that the line of approach in their study was through
experiments in breeding these forms to discover their origin and
true character. This has been done with a few forms, but the whole
field who would devote
should be worked by some competent geneticist,
his life to the undertaking. Without further discussion, it has been
thought advisable to put in a form accessible to American college
students, a glossary of the more important terms used in teratology.
With the exception of a few additions the terms given in first volume
of " Pflanzen-Teratologie" (1890) by Dr. 0. Penzig are here translated
from the original, as serving as an outline of teratology for American
students.
Abortion (Masters and English authors; Abortus, German Avortion
or Avortement, French)— Stunting of an organ, that is the exceptionally
small formation of the same, whereby the form remains unchanged.
The German and French authors use the same expression very fre-
quently for the cases where a certain organ is entirely suppressed and
does not make an appearance.
Acaiilosy. —Acaulosia is the diminution in the size of the stem, for
absolute suppression of the stem, as the terms acaulescent and
331
—
acaulosia would signify, is an impossibility in a typic plant. The term

is purely a relative one.
Acheilary (Ch. Morren). — The suppression of the labellum in such
flowers as the Orchidace^.
Adesmy (Ch. Morren). — Congenital separation of organs which
are normally united together, therefore, often included as atavism.
Morren distinguishes between homologous adesmy as the separation
ofmembers of one whorl and heterologous adesmy the separation of the
members of one whorl from those of another.
—
Adenopetaly. Formation of a nectary in a former nectarless petal.
—
Adhesion.^ Normally used for the union of parts of different whorls
in the flower, for example, the union of a sepal with a petal, or of a
stamen with a carpel, and also for fusion in general (of a branch with
the main axis, of a leaf with a branch, etc.).
—
Adherence (Moquin-Tandon). Fusion of organs which normally
are separate.
Anaeretic (Schimper, 1854).^ Vnder foli alio anceretka, C. Schimper
obviously understood the abnormal arrangement of leaves on an axis
in a single row, a condition sometimes produced by a torsion, or twisting
of the axis.
Antherophylly (Ch. Morren).- — Formation of anthers upon leaf
blades.
Anthesmolysis (Engelmann). — Central or lateral metamorphosis
of an inflorescence, especiafly of heads as in the Dispaceae and
Compositse.
Antholysis (Spenner in Flor. Friburg). —A solution of flowers,
particularly applied to the condition in which the axis becomes elongated
and the flower whorls separated from each other.
Aphylly.^ —The condition of the plant in which leaves are suppressed.
Apilary (Ch. Morren). — Suppression of the upper in normallylip
bilabiate flowers, as in Calceolaria.

Apogamy.- —Vegetative reproduction of plant individuals instead
of by the usual method with sex organs, especially used with reference
to ferns where the antheridia and archegonia are suppressed or not
functional, the young plant arising directly from the prothallium. If
is also used for the non-sexual formation of embryos in the embryo sac
of the phanerogams.
Apophysis. —Vegetative, central proliferation of an inflorescence.
—
CLASSIFICATION OF PLANT ABNORMALITIES ^^^
Apostasis. — The monstrous disunion of parts normally united as

which the whorls are
in the elongation of a flower axis, as a result of
transformed into spirals. One, however, uses the term for the sepa-
ration of single floral phyllomes, for example single sepals from the
calycine whorl.
Atrophy. —Wasting away; degeneration of organs; abortion.
Autophyllogeny (Ch. Morren). —
The budding of one leaf from
another, as from the midrib.
Balance Organic (Moquin-Tandon).Ône uses this expression for
cases thatby atrophy of single organs of a plant is compensated by
hypertrophy of others.
Biastrepsis (C. Schimper). —This is analogous to the torsion, or
twisting of other authors.
Blastomany (A, Braun). —Abnormal tendency of single plant
individuals to develop an unusual number of leaf buds (axillary or
adventitious).
Calycanthemy (Masters). —Transformation sepals petaloid of to
structure.
Calyphyomy (Ch. Morren). —Adhesion one or the sepals
of all of to
the back of the petals.
Cenanthy (Ch. Morren).- Kevds = empty + avdos = flower: Abor-
tion, or suppression of the stamens and pistils of a flower, leaving the
perianth empty.
Ceratomany. —Abnormal formation of horn-like, or hooded, fre-
quently nectariferous structures in a flower. Clos has employed the

same term for the increase in the spurs in many families (Orchidace^) .
—
Chellomany (Ch. Morren). The doubling of the lip, or labellum,
in orchids, as in Orchis morio.
Chloranthy. —The transformation, or change of all or most of the
floral parts into leaf-like green parts; frondescence.
Chorisis. — The separation of a leaf or phylloid part into more than
one; dedoublement, doubling.
Cladomany. —An abnormally richly branched plant.
Cohesion. — A union between the members one and of the same
whorl (particularly in flowers), or between the parts of a composite
organ.
Coryphylly. —An abnormality in which a leaf ends the axis. This
leaf is sometimes colored.
Crateria.^ — C. Schimper uses this term for a leaf blade which de-
velops ascidia, as the ascidial white ash discovered by George H. Shull.
Cyclochorisis (Fermond). — Division of an axial organ in two direc-
tions, so that in place of a simple axis there arise whole clusters of
secondary axes.
Dedoublement (chorisis, doubling) .^
— Congenital division of an
organ in which several parts arise out of a single primordium. Lateral
and serial dedoublement are distinguishable.
Fig. 132. —Twin cherries due to dialysis, or disjunction, of the pistil of the flower
into two carpels, each of which matures into perfect drupe joined at the base with
its fellow. Philadelphia Market, May 25, 1916.
Deformation.^ —A malformation, or alteration from the normal

kind. A general expression for the irregular formation of an organ,
or a complex of organs.
Degeneration (Masters) —
Stunted formation of an organ with
.^
which changes of form are associated. An alteration for the worse.

—
Dialysis (Ch. Morren, Masters). The separation of parts normally
in one, especially parts of the same whorl. Scarcely distinguishable
from adesmy (Fig. 132).
Diaphysis (Engelmann).—A central proliferation of flowers. If
the flower axis elongated beyond the carpels bears another flower, we
CLASSIFICATION OF PLANT ABNORMALITIES 335
speak of Diaphysis floriparous; if leafy shoots arise, it is Diaphysis

frondiparons; if a cluster of flowers, it is known as Diaphysis
racemiparous.
Diplasy (Fermond). — The division of an axial organ into two
parts.
Diremption.^ —The occasional separation, or displacement of leaves.
Diruption. —A term used by Germain de St. Pierre for different
appearances (division of leaves, axes, fasciation).
Discentration (C. Schimper).^ —A term applied to fasciation of an
axial organ, but used occasionally for the multiple division of a
phyllome.
—
Displacement (Masters). The abnormal position of a plant organ.
—
Distrophy (Re). The dissimilar formation of the homologous
organs of a plant.
Divulsion (St. Germain de — See diruption.
Pierre).-
Ecblastesis (Engelmann).^ —^Lateral proliferation, that is bud for-

mation in the axils of flower parts (sepals, petals, stamens or carpels).
There can be distinguished floriparous, frondiparous and racemiparous
kinds of ecblastesis.
Enation.^ —The formation of excrescences of different kinds on the
upper surface of other organs. We find scales projecting from petals,
small lamina on foliage, leaves, etc.
Epanody (Ch. Morren). —Abnormal reversion of an organ to a
simpler form than it normally shows.
Epipedochorisis (Fermond).- —A manifold division of an axial
organ in one plane. Frequently not distinguishable from fasciation.
Epistrophy (Ch. Morren). —A reversion of an apparently constant
monstrosity to the normal form of single organs, for example, the
development of branches with normal leaves in place of those with cleft
leaves.
Etiolated. — Blanched, or lengthened abnormally by the absence of
light.
Expansivity .^
—A term used by Germain de St. Pierre with a similar
sense to Diruption and Divulsion.
Fasciation (Olaus Borrich, i67i).Â flat band-like, or ribbon-like
expansion of a normal cylindric axis, or stem, associated with departure
from the normal leaf position. If flowers are developed they are
generally altered in structure (Fig. 133).
—
Fission. — A division of a normally simple organ.

Frondescence.^ —The prolifer-
ation of a normally reduced petal
to a foliage leaf with lamina.
Gamomery (Engelmann).
The condition in which the
normally distinct petals are
united into a gamopetalous
corolla.
Gemmiparity. —The condtion
of leaves which develop adventi-
tious buds.
Gymnaxony (Ch. Morren).—
The condition which the
in
placenta protrudes through the
ovary of the flower.
Gynophylly (Ch. Morren).
The transformation of a carpel
into a foliage leaf. Phyllomor-
phy of the ovary.
Hemitery. —An abnormality
of elementary organs, or of axial
appendages.
Heterogamy (Masters). — An
alteration in the position of the
sexual organs.
Heteromorphy (Masters).
Irregular formation of an organ.
—
Heterotaxy. This term is
used by Masters for the cases in
which a new organ, or structure,
appears in unusual places, as leaf
buds and flower buds on a root.
Later authors (Freyhold) use the
word in an entirely different sense
Fig. 133.— Fasciated stem and fruits o^ for the inversion of the floral plan.
the poppy {Papaver). {Drawing by Alice M-
Russell.)
Homotypy. The — develop-
ment of an organ, or of any
structure in the same place, where normally another one originates.
Hypertrophy. —An abnormal largeness, strong formations of any

plant part.
Idiotery.—A monstrosity by which a plant departs from the normal
type and from related forms.
all of its
Lepyrophylly (Ch. Morren). — The transformation the integu- of

ments the ovule into
of or scales, leaves.
Meiophylly. — The diminution the number leaves
in a whorl, as of in
compared with those of the preceding whorl.
Meiotaxy.— The suppression of entire whorls.
—
Metamorphosis. The transformation of an organ into another one,
that is morphologically equivalent to it, but it may be has a wholly
different appearance and other functions.
Metaphery (Ch. Morren) .^
—The displacement of organs, as when
become opposite.
alternate
Metastasis (Moquin-Tandon). —The shifting of an organ to some
unusual position.
Mischomany (Ch. Morren). —An increase in the number of pedicels
Muscari comosum.
or the branching of the inflorescence, as in
—
Monosy (Ch. Morren). Separation of floral parts from one another
with which they normally are in Cohesion, or Adhesion. The abnormal
isolation of parts due to a desmy or dialysis.
Multiplication. —The division of an order into many homologous
parts.
Oolysis. —A greening (viridescence) which shows conspicuously
in the carpels and ovules of the flowers.
Peloria (Linnaeus). —The radial (actinomorphic) regular formation
of anormal zygomorphic (irregular) flower.
—
Periphyllogeny (Weinmann). The formation of numerous leaflets
about the border of a leaf blade.
—
Permutation (De CandoUe). An enlargement of the floral envelopes
with corresponding abortion of the sexual organs.
Petalody. —The metamorphosis of stamens, or other organs into
petals with their usual form, color and consistence.
Petalomania. — An abnormal multiplication of petals.
Phyllocally (Lemaire). —The budding new of leaflets on the surface
of foliage leaves.
Phyllody (Masters). —The appearance of foliage leaves in place of
floral ones.
Phyllomania. —An abnormal production green leaves. of

Pistillody— The transformation of parts into carpels.
floral
Pleiomorphy (Masters). — An abnormal or excessive development.

Pleiophylly (Masters) — The appearance
.^
many leaves place of
of in
a single part.
Pleiotaxy (Masters). — The increase in the number of whorls in a
flower.
Plesiasmy (Fermond). —An abnormal shortening the stem of inter-
nodes, so that the leaves are arranged closely together.
Pollaplasy (Fermond). — The division a theoretic simple organ
of
into many analogous structures.
Polyclady. —An unusual development branches and of twigs.
Polyphylly. — The abnormal increase the number partsin the of of
floralwhorls.
—This term used with a number
Prolification. is of different
meanings. One is the central, or lateral, outgrowth from a flower, or
an inflorescence. The different kinds are designated as median, axil-
lary, extrafloral, while each kind is again divided into foliar and floral,
depending upon the nature of the adventitious bud. The axillary

prolification is known as echlastesis (Engelmann) and the median as
diaphysis.
Rachitism (Touchy). — Hypertrophy of the floral envelopes, as in
JUNCACE.E, CyPERACE^, GrAMINACE.E.
Recrudescence.- —The production of a leafy, or flowering, shoot from
an axis of inflorescence after the formation of ripe fruit on that axis,
Rhizocallesy (Ch. Morren). — The union of two plants of the same

species solely by their roots.
Salpinganthy (Ch. Morren).— The transformation of ligulate or
ray florets of Compositae into conspicuous tubular florets.
Scyphogeny (Ch. Morren). —The formation of ascidia from leaf
blades.
Sepalody.^The transformation of petals into sepals, or sepaloid
parts.
Solenoidy. (Ch Morren). — The metamorphosis of stamens into
tubular structures.
Solution (Masters) —Abnormal
.^
separation of the members .of a
whorl from those of another (similar to the Adesmia heterologous of
Morren).
Sphaerochorisis (Fermond). — Multiple division of an axis in all
directions producing a witches'-broom-like arrangement of branches.

Speiranthy (Ch. Morren). —The anomalous condition in which the
flowers develop into a twisted form.
Spiroism (Ch. Morren).— An elongated snail-like development of
an organ.
—
Staminody. The transformation of a petal into a stamen.
—
Stasimorphy (Masters). The arrest in the development of an
organ, or an organ complex, and the stoppage of development at a lower
stage.
—
Stesomy (Ch. Morren). A term with similar usage to stasimorphy.
—
Strophomany (Schimper).^ A term used in the same sense as
biastrepsis for twisting, or torsion.
Suppression. —The complete abortion of an organ.
Synandry.^— The abnormal union of stamens.
Synanthy.^—Lateral union of two or more flowers. This condition
can arise in a number of ways; for example, by the approach and fusion
of two floral fundaments, or through the partial forking of a receptacle,
or through floriparous.ecblastesis, etc.
—
Synanthody. ^Lateral union of two floral buds on the same stalk,
or on two pedunfles which have become fasciated.
—
Syncarpy.- ^Lateral fusion of two or more fruits. This condition
is the natural result of synanthy.
Synophthy (Ch. Morren). —The union of two leaf buds, or foliage
shoots with each other.
Sjmspermy.^ —The fusion of several seeds.
Taxitery (Gubler). —A modification which is so slight that it admits
of comparison with the normal form. Contrast Idiotery.
Torsion. —A spiral twisting, or bending, or parts or organs.
Triplasy (Fermond). —The separation of an organ into three analo-
gous structures. Trifurcation.
Virescence. —The abnormal development of flowers in which all
organs are colored green and more or less wholly transformed to small
foliage leaves. If the metamorphosis is complete, there result foliage
leaves with distinct lamina and this condition is known as frondescence.
In concluding might be well
this glossary of teratologic terms, it
to add that a recent work on plant teratology has appeared. It is
designed to bring our knowledge up to date. The first volume of
Worsdell's^ "Principles of Plant Teratology" includes a consideration

of the fungi and bryophytes as non-vascular plants and with vascular
plants he goes as far as a consideration of the teratology of roots, stems,
leavesand flowers. It is issued by the Ray Society, as was that of
Maxwell T. Masters in 1869.
^ WoRSDELL, Wilson Crosfield: The Principles of Plant Teratology, vol. i.,
London, printed for the Ray Society, 1915; vol. ii, 1916.
CHAPTER XXVHI
SYMPTOMS OF DISEASE (SYMPTOMATOLOGY)
The preceding pages have dealt with the causes of plant diseases,
that is their etiology. It remains to discuss the symptoms of disease
as that is a very important matter in deciding as to the nature of the
disease, and the harm that the various diseases may do to our agri-
cultural crops. It is easy to determine that there is something wrong
with the plant, because such well-known symptoms as withering,
as yellowing, as abnormal growth are evidences of it, but it is quite
another thing to decide as to the specific nature of the disease, its cause
and probable amelioration. Even to the trained plant pathologist, it
is not an easy problem to decide what the trouble is. It requires some-
times two or three years of research work with all the refined methods
of modern science to reach a satisfactory conclusion, and at times even
then the solution is baffling. To call a pathologist, or a botanist, an
ignoramus, because he cannot by a study of the symptoms name the dis-
ease, is unworthy of people who claim to be cultured, and yet it fre-
quently happens that the farmer's opinion of the book scientist is based
upon just such a flimsy pretext. General conclusions are reached in
this field of inquiry, just as in other fields, by the process of exclusion.
The pathologist puts questions to himself about the plant and gradually
he eliminates the impossible conditions, gradually narrowing himself
down to a few possibilities. For example, he might ask himself
whether the cause of the disease is external or internal. If external,
then whether it is due to climate, to animals, or plant parasites. If
plant parasites are concerned, then are they flowering plants or fungi.
We will suppose that he finds that the disease is of fungal origin. Then
with the cultural means at his disposal, the fungus must be obtained
in pure culture, and its pathogenicity tried out upon healthy individuals
corresponding racially, or specifically, with the diseased ones. If the
inoculation of the healthy host is successful, then the recovery of the
fungus from the tissues for comparative cultural study will follow.
341
.
Knowing the specific fungal organism, a great stride has been made
toward a comprehensive knowledge of the disease.
The plant pathologist, who would be successful in his profession,
must be acquainted with the normal, or healthy, conditions of plants,
or how can he study the unhealthy states? Any departure from the
healthy state is indicated by a certain behavior of the plant, or reac-
tion to the causes of disease and certain peculiarities of structure, form
and color are also manifested. An investigation of these character-
istics of disease concerns symptomatology. The most common symp-
toms of plant diseases may be classified according to the outline pre-
sented by Heald in Bulletin 135 of the University of Texas, Nov. 15,
1909, entitled "Symptoms of Diseases in Plants."
1. Discoloration or change of color from the normal.

(a) Pallor. Yellowish or white instead of the normal green.
(b) Colored spots or areas on leaves or stems.
Whitish or gray: mildews; white rusts, etc.
Yellow: many leaf spots.
Red or orange: rusts, leaf spots, etc.
Brown: many leaf spots.
Black: black rust, tar spots, etc.
Variegated: leaf spots, etc.

2. Shot-hole: perforation of leaves.
3. Wilting: "damping-off," "wilt," etc.
4. Necrosis: death of parts, as leaves, twigs, stems, etc.
5. Reduction in size: dwarfing or atrophy.
6. Increase in size: hypertrophy.
7. Replacement of organs by a new structure.
8. Mummification.
9. Change of position.
10. Destruction of organs.
1 1 Excrescences and malformations.
Galls: pustules, tumors, corky outgrowths, crown galls, etc.
Cankers: malformations in the bark generally resulting in an open

wound.
Punks or conchs and other fruits of fleshy fungi. .
Witches' brooms.
Rosettes and hairy root.
SYMPTOMS OF DISEASE (SYMPTOMATOLOGY) 343
12. Exudations.
Slime flux.
Gummosis: especially for stone fruits.

Resinosis: especially for coniferous trees.
13. Rotting:
Dry rot and soft rot: "the gangrene" of plant tissue.
Root rots: alfalfa, cotton, beets, cherry, etc., generally woody or

fleshy roots.
Stem or trunk: dry rot of trees; rot of modified stems like rhi-
zomes, bulbs, or tubers. >
Buds.
Fruits: fleshy fruits of various kinds.
It will be profitable to discuss the symptoms of disease under the

above heads.
I. Discolor ations.
—
^The unnatural, or false color which plants
assume under diseased conditions may be included under the head of
discolorations. Sometimes, as in woods, the discoloration may appear
as a stain. AbnormaUty of color usua|y accompanies other symptoms
of plant disease. Pallor, or chlorosis, vêre the plant assumes a yellow-
ish to white, or sickly-pale hue, is due to a number of causes. Promi-
nently, one form is due to the absence of light, whereby the plant be-
comes etiolated, or suffers etiolation. It is considered that the laying of
wheat and other cereals is one form of this etiolation where, through
lack of carbohydrates, the cellulose which forms the strengthening of the
cell wall does not form properly.;;,' Sometimes the gardener induces
etiolation in his celery, endive and asparagus plants, where the blanch-
ing is secured by covering such plants with soil. True chlorosis is due
to an enzyme which destroys the chlorophyll pigment of the chloroplasts
which are fully developed. Icterus is the condition where the organs
are only yellow; chlorosis, where they are white, such as in the mosaic,
or calico disease of plants formerly described. Yellowing may be in-
duced experimentally by an excess of carbon dioxide, in fact yellowing
accompanies wilting, the attack of wire worms, the presence of poisons,
or acid gases.
Variegation and albinism may be apparently normal conditions of
some varieties of plants, for gardeners and horticulturists grow such
plants by preference for decorative uses. This variegation, or albinism,
is induced in all probability by the presence of oxidizing enzymes in
patches of cells where the chlorophyll pigment is destroyed and not in

other adjoining areas.
The formation of spots on leaves (Fig. 134), stems, flowers, or fruits is
due to a variety of causes. The grayish or whitish spots on the under

surface of grape leaves are due to mildews, on the stems of cruciferous
plants to white rusts and on the leaves of the parsnip are found white
spots due to a fungus, Cercospordla. Grayish spots on the prickly pear
Fig. 134. —
Apple leaves showing leaf spots produced by natural infection with
Sphaeropsis malorum. {After Scott, W. M., and Rorer, J. B., Bull. 121, U. S. Bureau
of Plant Industry, 1908.)
and on the leaves of the box trees are occasioned by a disease known as
anthracnose. Many leaf spots are yellow as in violets, oaks, cucumbers
and melons. The red or orange spots on plants usually suggest the
presence of rusts as on wheat, rye, alfalfa and a host of other cultivated
and wild plants. The so-called tar spots of the maple leaves are bla'gk
in color and such discolorations of the leaf surface are traceable to the
attack of a fungus, Rhytisma acerinum. Apples are frequently marked
by fly specks which are usually clustered as small circular black spots..
A fungus is the causal agent.
2. Shot-holes (Fig. 135). —The perforations and the forma-

of leaves
tion of what are form of fungous

called shot-holes illustrate another
attack, where circular patches of dead tissue killed by the fungus drop
out leaving a hole. The Enghsh morello cherry trees in some sections
of our country have been killed during the past few years by this " shot-
FlG. 135. — Shot-hole disease of the plum due to Cylindrosporium {.After

Heald, F. D., Bull. 135 {Sci. Ser. 14), Univ. of Tex., Nov. 15, 1909.)
hole" disease. When the funguses belonging to the genera Cercospora

and Phyllostida attack the leaves of Virginia creeper perforations may
be formed.
3. Wilting. — Wilting in general is due to the lack of sufficient water
to supply that lost by transpiration, for wherever the amount of water
transpired exceeds that absorbed by the roots wilting occurs. Wilting

may result, if the normal ascent of the sap is interfered with by the
growth of fungi into the water-conducting tissues, the entrance of bac-
teria into the woody vessels of the plant, whereby they are literally
plugged with such organisms, or some injury which cuts off the ascend-
ing current of water. Damping-off is a form of- wilt in which an oomy-
cetous fungus enters the collar of seedling plants, or where a Rhizoctonia
species invests the roots of the growing plants and interferes with the
regular water absorptive processes.
4. iVecro5M.— Necrosis is the mortification, or death, of the tissues.
The term is usually applied to the death, or loss of vitality, of one part of
a plant, while the other parts remain alive. When the fungus, Fusa-
rium trichothecoides is inoculated into Green Mountain potato tubers,
,
in about three weeks' time it will be found that a portion of the tuber,
usually the central part directly beneath the point of inoculation, has
undergone necrosis. The surface of the potato tuber becomes sunken
through the death and collapse of the starch containing cells and the
lesions may involve half of the tuber. The black rot of the navel
orange is due to a fungus, AUernaria citri, which gains entrance to the
fruit through slight imperfections about the navel end. A black
decayed area is found under the skin. This decay does not spread im-
mediately through the entire fruit, but remains for weeks- as a small
black necrotic area with a mass of the fungus present. The decayed tis-
sue does not always extend to the surface, but remains beneath the skin.
Necrosis often follows the action of frost in killing the cortex cells of
fruit trees in patches with a blackening of the tissues. Fire bhght may
be the cause of necrosis, for the cambium which is killed dries up in
black patches.
5. Dwarfing. —A reduction in the size of a plant is very often asso-
ciated with disease. This may be true of the whole plant, or some
particular organ only may be dwarfed. Apples are frequently reduced
in sizeby the attack of the scab fungus, sometimes not reaching one-
fourth the size, and the same is true of apples affected by the cedar rust.
Dwarfing of the whole plant may be a symptom of malnutrition. It
may be evidence of a poor soil, or the repeated maiming, or nipping off of
the buds by cattle, or purposely by man, as is the case with the minia-
ture trees of the Japanese. Dwarfing, or nanism, may be the result
of climate, as is the normal case with alpine plants. Prostrate forms of
trees of great age are formed by the action of the dimate of high
mountains, or by growth in porous sand on exposed sea dunes. Atro-
phy, or the non-formation of parts, or organs, is a phase of dwarfing.
It is seen in the dwindling of organs in size, as the result of various
causes, such as the attack of fungi. The carpels of Anemone are
atrophied in plants infested by Mcidium and the whole flower is sup-
pressed when the cherry is attacked by Ex-
oasciis cerasi. Exoascus pruni is responsible
for the absence of the stone in plum fruits, etc.
6. Hypertrophy. — The undue excessive de-
velopment of a plant part is a symptom of a
diseased condition of that part. The bladder
plums formed in the plum pocket disease are
good illustrations of hypertrophied tissues, as
the replacement of the rye ovary by the ergot
sclerotium, following the entrance of the spores
of Clamceps purpurea. The attack of Gym-
nosporangium biseptatum (Fig. 136) results in
the massive enlargement of the stem of the
white cedar. A rust fungus is responsible for
the increase in size of the twigs and petioles of
our common ash and elder.
7. Replacement: — Â new structure takes
the place of organs.
8. M ummifi cation. —The drying and
wrinkling of fruits and other plant parts Fig. 136. — Swelling of
where the general shape of the part is pre- main stem of white cedar
caused by Gymnosporangium
served, but in a reduced size, is an evidence biseptatum. (After Harsh-
of the unhealthy condition of that organ, or berger, Proc. Acad. Nat. Set.,
Phila., May, 1902.)
part. The attack of the black-rot fungus,
Sphceropsis malorum, brings about a slow desiccation of the fruit which
may remain hanging on the tree over winter and in a shriveled condi-
tion. Frequently, the mummies produce a crop of spores, which spread
the disease.
9. Alteration of Position. —
The change of position of an organ from
itsnormal one'is a sure symptom of disease, usually the attack of some
fungous parasite. The normal position of the leaves of the house leek,
Sempervivum tectorum, is that of a rosette with the spirally arranged
leaves approximately horizontal. When attacked by a rust fungus,

Endophyllum sempervivi (Fig. 137), the diseased leaves grow erect.
The same is true with our native American hepatica, Hepatica triloba.
Infrequently, it is attacked by a rust fungus in the aecial condition,
Tranzschelia punctata, so that (Fig. 138), the rusted leaves develop a
larger, stiffer petiole, stand erect .with a smaller, stifYer leaf blade on
which the aecia are found. The common garden purslane Portiilaca
oleracea, usually grows in a prostrate position, but when attacked by
the white rust, Cystopus (Albugo) portulacce, many of the diseased
branches become erect or ascending. The stems ofVaccinium vitis-
idcBa become erect the second year after infection by Melampsora
Goeppertiana.
Fig. 137. —
Two plants of house-leek, Sempervivum. Left one affected by Endo-
phyllum sempervivi. Right one, a healthy plant. {After Grove, W. B.: The Brilish
Rusl Fungi, 1913: 54.
10. Destruction of Organs. —The destruction of plant organs by the

attack of fungi is well illustrated by the cereal smuts, which attack the
flower parts reducing them to a black powdery mass of spores, which are
carried away, leaving nothing but the bare axis on which the flowers
were originally situated.
11. Excrescences and Malformations. —
These will be treated of in
detail in another chapter. Here it may be said that galls, pustules,
tumors, corky outgrowths, crown galls, cankers, burls, or knauers,
(Fig. 139) witches' brooms (Fig. 140), etc., are evidences of diseased
conditions. The nature of these excrescences and malformations can-
not be discussed here, but it may be said that they are specific and
usually associated with the attack of some fungus, as for example the
plum knot due to Plowrightia morbosa, the cedar apples formed on the
—
Fig. 138. Hepatica triloba parasitized by a rust fungus, Tranzschelia punctata,

which causes some of the leaves to stiffen and grow erect. Left figure shows ascia,
April 29, 1915.
35° GENERAL PLANT PATHOLOGY
red cedar by Gymnosporangimn juniperi-virginiatKE. The crown galls,
or possible vegetal caricers, are another illustration of such excres-

cences, while malformations are represented by peach leaf curl and
the witches' brooms on trees.
12. Exudations. —
The formation of slimy substances, which flow
from trees and plants, the diseased conditions known as bacteriosis,
gummosis^ and resinosis, illustrate the character of the exudations from
Fig. 139. -Burl, or enlarged base of an oak tree in the forest on Gardiner's Island,
New York, July 17, 1915.
plants under abnormal conditions. The production of clear amber-

colored secretions, which accumulate on the surface of the diseased parts,
isknown as gummosis and is seen in cherries, apricots, almonds and
many other trees. It follows wounds or the attack of fungi. The
same condition in coniferous trees is known as resinosis and in a few
trees it is of economic interest because, as in the spruce, the exudation of
1 Wolf, Frederick A.: Gummosis. The Plant World, 15: 49-59, March, 191 2.
Butler, O.: A Study on Gummosis of Prmiits and Citrus. Annals of Botany,
25: 107-153, 1910-
1.
Fig. 140. — Branch-knot or witches'-broom of the Hackberry ,(C>//J5 occidentalis)

{After Kellerman, W. A., Mycological Bulletin, Nos. 61-72, July, 1906.
gum rosin known as ''spruce gum" is collected and sold at from two
dollars to two dollars and fifty cents a pound. ^ Where due to the attack
of bacteria it is called bacteriosis. Tumescence is the over-turgescence
of plant tissues due to the excess of water. It sometimes indicates
pathologic changes and was formerly called (edema, or dropsy. Flux
is another name applied to the issuance of fluids from wounds in trees,
while slime flux issuing from wounds may be frothy, owing to the fer-
mentative activity of yeasts and other fungi, which live in such slimes.
Manna flux is found in such trees as the manna ash and species of
tamarisk. Cuckoo spit is a frothy material found on grasses and
g
The wet rot of potatoes is probably due to putrefactive bacteria. The

tissues become soft, then mushy, and finally become a liquid mass with
a vile smell.
BIBLIOGRAPHY OF PLANT DISEASES IN GENERAL

Heald, Frederick D.: Symptoms of Disease in Plants. Bull. 135, University of
Texas, Nov. 15, 1909.
Klebahn, Prof. Dr. H.: Grundziige der allgemeinen Phytopathologie. Berlin
Gebriider Borntraeger, 191 2.
KtJSTER, Dr. Ern.st: Pathologische Pflanzenanatomie. Gustav Fischer in Jena,
1903, Zweite Auflage, 1916.

KtJSTER, Dr. Ernst: Pathological Plant Anatomy. Authorized translation by
Frances Dorrance, 1913-1915.
Smith, John B.: Economic Entomology for the Farmer and Fruit Grower, and for
Use as a Text-book in Agricultural Schools and Colleges, J. B. Lippincott Co.,
1896.
Stengel, Alfred: A Text-book of Pathology, W. B. Saunders Co., Philadelphia,
1906.
Ward, H. Marshall: Disease in Plants, Macmillan Co., & 1901.
23
CHAPTER XXIX
PATHOLOGIC PLANT ANATOMY
With the multiplicity of higher plant forms, in which the same end
is attained in a diversity of ways, the terms normal and abnormal
become in one sense merely relative terms for what apparently is the
normal method of procedure in one group of plants, may be decidedly
different, or abnormal, in other uncommon groups. The words normal
and abnormal are, therefore, variable terms, but useful ones. Speci-
fically, when we use the word abnormal, we mean the departure, or
deviation, from the normal (average) structure or function of the mem-

bers of any group selected for investigation. Pathologic plant anatomy,
therefore, has to deal with abnormal, but not necessarily diseased
organs, and yet a study of diseased tissues is an important subject of
investigation for the plant pathologist.
The material which forms the substance of our inquiry naturally
falls into two principal groups.
1. The differentiation, number or size of the cells of pathologic
tissuesremain more or less below the normal, so that the tissues in one
or more ways remain in a stage of incomplete development. The term
Hypoplasia designates those abnormal processes of formation, which
compared with the corresponding normal processes of development
appear retarded as it were and prematurely.
2. The pathologic cells and tissues exceed the conditions of differen-
tiation and growth characteristic of normal plants, so that a treatment

of such necessitates a consideration of several independent groups.
(a) The abnormal from the normal ones only in their
cells differ
and for the processes of

internal structure (contents, mechanics, etc.)
differentiation by which the tissue cells supplement their normal
qualities, or exchange them for new ones, the term Metaplasia is
used.
(6) The increase in size of abnormal cells over normal ones is
termed Hypertrophy (v-rrep = over, excessive; rpepoi = to nourish),
and it is not important fundamentally whether the histologic structure
354
PATHOLOGIC PLANT ANATOMY 355
of the cells concerned remains similar to that of the normal ones, or is
altered in some way.

(c) The by an increase in the number of its indi-
increase of a part
vidual structural elements known as Hyperplasia (wep = over,
is
excessive; TrXoo-ts = formation, structure), and this depends on cell

division following cell growth. A large number of abnormal formations
arise through hyperplasia and the histology of the newly formed tissues
is exceedingly varied.
3. The
processes of Restitution consist in the restoration of
structures, which resemble those lost in injuries and mutilations of the
plant body. Although the tissues thus formed are like the normal
ones yet their formation following injuries, or mutilations, comes within
the realm of pathologic anatomy.
Hence we shall treat of morbid anatomy under the five heads
suggested in the above considerations. Naturally the material for
our investigation and treatment arranges itself into five chapters, on
"Restitution," "Hypoplasia," "Metaplasia," "Hypertrophy" and
"Hyperplasia."
RESTITUTION
Following a wound or other injury or the removal of a plant part,
the organs are stimulated to renew the lost part, or to repair the damage
to the cells or tissues. The regeneration of lost or injured plant cells,
tissues, or organs, is called specifically in pathologic plant anatomy

restitution, wHile the word regeneration, although implying restitution
(L. restitutio (-n), < restitutus, pp. of restituo, restore, < re-, again, +
statuo, set up, < sto, stand), is used in a somewhat different sense.
The process of restitution, it is conceivable, includes a number of
distinct operations.^ The newly formed parts are formed at the place
ofamputation and are like the lost portion (as the regeneration of root
newly formed parts, which resemble the lost ones, are not
tips) or the
produced at the injured place, but some distance away from it, or the
new parts arise on the cut surface, but are unlike the lost part (hetero-
morphosis), and finally the new parts do not resemble the lost ones, nor
do they arise at the surface of the amputation.
It will be profitable to discuss the two most important forms of
* Consult Studien iiber die Regeneration v. Professor Dr. B. N^mec. Mit 18
Textabb.
and that of the tissues. The experi-

restitution, viz., that of the cell
ments Tittman have shown that the waxy cuticle of the castor-oil
of
plant, Ricinus communis, may be restored after removal. Exposure
of the protoplast results in many cases in the formation of a new cell
membrane, as is illustrated in some of the large-celled algae belonging to
the Siphoned. Frequently, it is possible to demonstrate the restitu-
tion of the cell membrane by the process of plasmolysis in which the
protoplasm is made to retreat from the cell wall. The time varies for
its formation under conditions of plasmolysis. In Conferva, it takes
place in one to two days, in Zygnema in three to four days. When the
root hairs of dicotyledonous plants are plasmolyzed new membranes
are formed about the protoplast.
Wounded siphonaceous algal cells {Caulerpa, Valonia, Vaucheria),
where the cell wall has been injured, are capable of restoring the cell
wall. Some fungi show such restitution also, while the injured cells
of the higher plants lack this power. A few exceptions are known where
nettle hairs of Urtica dioica may imperfectly replace the broken-off tip.
Pricking the turgid cell of Valonia utricularis, as I have done with

fresh specimens in Bermuda, is followed by the escape of a liquid jet
and later the opening is closed by a gall-like, protoplasmic, chloro-
phylless plug.
It has been demonstrated that the important cell wall can be regen-
erated on fragments of protoplasm provided the influence of the nucleus
is felt in such formation. Klebs has shown that, with the removal of
the nucleus from the cell, that cell has lost all its power to produce new
cell walls, but a distant nucleus may extend its wall-forming influence,
when removed several millimeters away in an adjoining cell.
In the restitution of tissues, we will consider those cases in which the

injured cells remain unhealed, but in which the uninjured neighboring
cells bring about the restitution. The removal of the rhizoidal hairs
on the thallus of Marchantia is followed by the appearance of other
hairs in a few days, which may grow out through the cavity of the
mutilated one as described so carefully by King. The mutilated
tip, or growing point, of many multicellular algae is replaced by the
development of the uppermost intact cell. Brefeld found in the

sclerotia of Coprinus stercorarius the inner cells are able to regenerate
the outer black cuticularized coat, if that is removed.
The number of cases of tissue restitution known in the higher plants
are few. The peridium, or secondary tegumentary tissue of stem or

root, is easily regenerated, as is seen in the formation of new cork layers
in the cork oak after the removal of older ones. The epidermis is not
always replaced but Massart found that removal of the epidermis of
Lysimachia vulgaris resulted in the regeneration of a new hair-bearing
epidermis. The regeneration of the vascular bundles has been studied
in monocotyledonous plants and in dicotyledons. The regeneration
of roots in monocotyledons consists in the replacement of epidermis,
phloem and xylem. In dicotyledons before the wood and bast are
replaced there is a regeneration of the endodermis, so that the restora-
tion of central cylinders, that have been destroyed, is not unusual.
HYPOPLASIA
The condition of hypoplasia in plants is one of arrested develop-
ments. The organism, or one of its parts, does not reach
normal devel-
opment, but that development is arrested, or stopped prematurely.
Hypoplasia is, therefore, defective development. The plant morpholo-
gists and plant anatomists are chiefly concerned with the problems
of arrested development and recently awakened interest has been
taken in its study, because it has been found that the interpretation of
certain phenomena is subject to experimental treatment, and hence,
there has arisen a coterie of experimental plant morphologists. Such
investigators have found that the processes of growth and differentia-
tion are not always equally arrested, which are associated in time and
place in the normal course of development. For example, leaves differ
from the normal by their small size. They may be retarded in their
form, as the narrow leaves of Sagittaria produced under water, or the
form may remain entirely undeveloped. We will treat of hypoplasia as
to thenumber of cells, as to the size of the cells, as to the differentiation
and the tissues.
of the cells
—
A. Number of Cells. It has been found in a study of the dwarf
forms of plants such as occur on high mountain tops that the condition
of nanism is not so much due to a decrease in the size of the cells over
those of the normal plant, but is chiefly conditioned on a reduction in the
number of cells. The internodes of plants may be shortened, the size
of the leaf blade may be reduced, the thickness in the leaf may be re-
duced, and this reduction in size is usually associated with a loss in the
358 •
GENERAL PLANT PATHOLOGY
number of cells, as for example, the omission of one of the palisade

layers of the leaf. External factors are important in determining the
structure of the leaf tissue, for the leaf more than any other plant
organ an index of the influence of climate. This fact is empha-
is
sized by a work entirely devoted to this subject and given the

appropriate title of "Phyllobiologie." There is a marked difference
in the thickness of beech leaves, for example, which have developed
under different environmental conditions, as I have proved satisfacto-
rily by the use of calipers and microscopic measurements, which show
an accurate coincidence. The thickness, or thinness, of such a leaf de-

pends essentially on the number of rows of cells. The thickest leaves
with the largest number of palisade layers which I have studied, grew
in the bright sunlight in exposed places along the edge of a salt marsh
at Cold Spring Harbor, Long Island. Sun leaves back from the influ-
ence of salt water were thinner and broader, while those, growing in
the dense shade of the forest in an inland situation near Philadelphia
were the broadest and thinnest of all. Not only was the mesophyll
modified in these leaves, but a marked difference was found in the shape
of the epidermal cells in the sun and shade leaves.
The number of cells which arise from the cambial layer suffers a
marked diminution in trees which grow under unfavorable climatic life
conditions. Drought, strong winds, pressure, unfavorable light and
nutrition are disturbing factors. Growth activity of the cambium may
cease entirely, if become too intensive. Huntington has
these factors
proved abundantly by his study of yellow pines of New Mexico and
the big trees of California that climatic cycles of wet and arid conditions
in the past history of North America can be determined from a study of
the size and character of the annual rings due to the cambial activity of
those trees, and he has plotted curves showing this relationship for a
period approximately 3500 years in the case of the big tree. Sequoia
gigantea}
B. Size of Cells. —
The size of ceUs must be considered also in dis-
cussing the phenomena of hypoplasia. Abnormally small cells may
be produced in different ways: A fresh division of the cells may take
place before the cells have reached the average size which they as-
sume under normal conditions. Klebs recites a case where he culti-
1 Huntington Ellsworth: The Climatic Factor. Publ. 192, Carnegie Institu-

tion of Washington, 1914: 153.
vated Euaslrum verrucosum, a desmidiaceous alga, in lo per cent, cane

sugar. The daughter cells formed by a previous division of those
cells divided again before they had attained their normal The
size.
conditions in the higher plants where hypoplasia is shown by the

production of abnormally small cells are such that the period of elon-
gation, which normally follows the last cell division, does not take
is stopped part way.
place, or Abnormally narrow tracheal tubes are
found in dwarfs, in etiolated and poorly nourished plants, or in in-
dividuals infected by fungi, or gall-producing animals. Disturbances
in nutrition reduce the size of the wood elements produced by cambial
activity.
In the study of the differentiation of cells and tissues, those cases
should be considered first which concern the individual cells, where the
formative process may stop prematurely. An investigation of Udotea
Desfontainii shows the arresting action of unfavorable life conditions
upon the development of the cell form. The leaf-like part of this alga
is composed of elongated sacs, which run lengthwise and parallel, with
numerous side branches of limited growth, which interlock to give

the thallus its characteristic firmness. If artificially cultivated, the
parallel sacs show undiminished growth activity, but the side branches
no longer show limited growth, but unlimited, and the thallus loses its
wonted form.
Arrestment of the development of the cell wall is indicated in the par-
tial, or entire cessation of the secondary growth in thickness, and as a
result, the elements normally thick-walled have walls of only moder-
ate thickness. Weak, or insufficient, transpiration acts pari passu in a

poor development of the cuticle of epidermal cells. Dwarfed plants
frequently show weakly developed cell membranes, as a sign of disturb-
ances in the nutritive processes. Chemic changes may be associated
with hypoplasia. Lignification is rarely excluded in the formation
under disturbing influences of the woody elements of plants. The cells
of themedullary parenchyma in thorns {CratcBgus) remain unhgnified,
when infected with a rust fungus, Roestelia. Finally, the formation of
cross walls may remain incomplete, thus giving rise to chambers,
sometimes communicating with each other.
Hypoplasia, as it affects the cell contents, may be seen in the
reduction in the number of chloroplasts in variegated leaves, in plants
with pale-green leaves and in plants which grow in places saturated with
vapor. The individual chlorophyll grains may not attain their normal
size, remaining small. The formation of chlorophyll presupposes a cer-
tain temperature, the action of light, the presence of iron and certain
organic food materials. Low temperature may reduce chlorophyll for-
mation, as is seen in grain seedlings and bulbous outgrowths or with
yellowish color grown under a low temperature. Deficiency of light and
iron causes etiolation, more especially chlorosis, or icterus in the absence
of normal pigment due to the lack of iron, while in vines unable to
absorb iron chlorosis may take place with abundance of iron in the soil.
Sometimes happens, on the other hand, following the attacks of an

it
insect that ripening lemons remain green-flecked. This condition

isdue to arrested development of the chloroplasts, which normally
would be transformed to yellow chromatophores.
Light also seems to influence the development of the red pigment,
anthocyanin, as is especially noticeable in varieties of Coleus, while
other parts, such as rhizomes, bulbs and roots, which remain under-
ground, are richly provided with anthocyanin. Chromogenic bacteria
may power of producing pigment, as is illustrated by Micro-
lose the
coccus prodigiosus grown at the high temperature of 4o°C. A.F.W.
Schimper and other botanists have shown that the formation and dis-
tribution of crystals of calcium 'oxalate in plants is to a large extent
dependent on external factors. Shade leaves contain fewer crystals
than sun leaves and plants grown in moist air, or without light, are also
poor in these crystals.
C. Tissue Differentiation. — The arrestment of tissue differentiation
can be illustrated in simple where the cells are united into colo-
algae
nies. When the green alga, Scenedesmus caudatus, the end cells of
which have gelatinous horns, is subjected to abnormal life conditions
the horns do not form. In the consideration of tissues of multicel-
lular growths it may be said that there is no organ in which homo-
plasia may not appear. Examples have been found in the hepatic and
true mosses.
The best illustrations of the developmental arrest of tissues are
found among the flowering plants, where as one case the guard cells of
the stomata may be arrested by a lowered transpiration and weak illumi-
nation. Stapf in his experiments with the potato, Solanum tuberosum,
showed that under normal conditions there was one stoma for every
forty-six epidermal cells, and in specimens matured by him in gaslight,
PATHOLOGIC PLANT ANATOMY 36]
there was a pair of guard cells for every 204 epidermal cells. The for-
mation on the edge of the ocrea of Folygonum amphibium
of the hairs
is entirely suppressed in the form natans, which is grown under water,
while they are present in the form terrestre. The modification of the
mesophyll tissue in homoplasia is .due to the character of the environ-
ment. Plants cultivated in places saturated with moisture, or after
infection by fungi or animals, show a homogeneous development of
the mesophyll.
In homoplasia, the vascular bundles decrease in number, the
mechanic tissue degenerates and the collenchyma sometimes does not
A B
Fig. 142. —
.4, Cross-section of a normal thalloid shoot of Lunularia. {After
Nestler,Die natiirlichen Pflanzenfamilien 1. 3, p. 17.) B, Cross-section of a thalloid
shoot grown in the absence of light. {After Beauverie in Ktister Pathologische PJlanzen
Anatomie, 1903: 42.)
form. Thouvenin by the use of mechanic pressure retarded the

development of the woody tissues in the stem of Zinnia. The stems of
Cardamine grown under water develop no mechanic tissue. The
length of the vascular bundles is less in plants grown in moist places
over plants which transpire strongly. Stahl found in his study of the
leaves of Lactuca scariola, that the mesophyll consists of palisade cells
throughout in the vertical leaves and in horizontal leaves lighted from
above of palisade cells only on the upper side of the leaf. If we call
upon homoplasia to explain the formation of shade leaves (Fig. 142), as
the unavoidable product of some arresting factor, then the structure of

shade leaves and those from alpine habitats, as well, as those placed under
water and which have a shade leaf structure, lose their remarkable char-
acter. Taking into consideration all of the experiments which have been
performed, it may be stated in concluding this chapter, that all of the
described hypoplasias may ' be traced back to scanty nourishment.
We are probably correct in assuming that there is poor nutrition in
plants grown an atmosphere deprived
in distilled water, in the dark, in
of its carbon dioxide in moist places, or under water. Insufficient
nourishment leads to an arrestment of differentiation and this becomes
evident in a number of ways.
Metaplasia
Metaplasia has been defined as the progressive change of any cell,

which is not connected with cell division and cell growth. The empha-
sis in this definition is upon the word progressive in contradistinction to
the word regressive. Metaplasia is less important in the histology of
plants than it is in animal histology. Changes of a metaplastic kind
are produced in the cells of plants, especially in the production of new
cell contents, or of the cell wall by increase in thickness.
Cell Contents. —
Frequently, it happens with tubers, bulbs, rhizomes
and roots of many plants that they develop a green color in place of
their normal chlorophylless character. Potato tubers kept in a damp,
warm, sunny place sometimes develop a green color and become
poisonous through the formation of metaplastic solanin. Bonnier
found that the tissues of his experimental plants exposed to strong arc
lights turned green even to the pith. Likewise red pigment dissolved
in cell sap may appear as a metaplastic change. For example, the nor-
mally green pitchers of Sarracenia purpurea become purplish green when
the plant is grown in intense sunlight. Such is also true in the heather,
Calluna vulgaris, Azolla, many succulents as Opuntia and Sedum. In-
jury to plant parts may be followed by the development of a red color.
The normal color of the leaves of Saxifraga Ugulata are green, but if leaves
are cut through the midrib, a red coloration developed along the edges
of the wound. Parasitic fungi may cause a local reddening of the cells
affected as in certain fruit and leaves spot diseases. The metaplastic

formation of coloring matters appears in the so-called graft hybrids.
The excessive formation of starch in the leaves of such plants as the

buckwheat, Polygonum fagopyrum, when insufficiently supplied with
chlorine is a case in point, as also the unfavorable nutrition occasioned
by potassium salts, while Schimper succeeded in getting the same ac-
cumulation of starch in unusual amounts in the leaves of Tradescantia
selloi by cultivation in nutrient solutions free from calcium.
Cell Membranes. —The metaplastic modifications of cell walls may
be considered under two heads. The first condition is found where
bordered pits are formed, as in such orchids as Cymbidium ensifolium,
and Epidendrum ciliare, whose leaves have been scarred.
LcBlia anceps
The second modification is seen where the cell walls have been thick-
ened abnormally by cellulose knobs, or thickenings. Such cellulose
deposits occur about calcium oxalate crystals, oil drops, as in Piper-
ace^, Laurace^ and about the hyphae of fungi which penetrate cells,
the hyphae along with certain cytoplasmic inclusions being surrounded
by the cellulose sheath bridging the space of the cell. Wortmann has
found heavy wall thickenings in the epidermis and bark of beans and
other twining plants, if they are prevented from carrying out their
reaction curvatures, while Kiister noticed the lignification of the cell
walls in the leaves of Juglans under the influence of certain plant lice.
CHAPTER XXX
PATHOLOGIC PLANT ANATOMY (CONTINUED)
HYPERTROPHY
The plant pathologist applies the word hypertrophy to an abnormal
process of growth in which the individual cells are larger than the nor-
mal, or when whole tissues become enlarged, or distended. Cell
division is left out of account as a means of the formation of hyper-
trophied cells, or tissues. The cells which are enlarged may be derived
from the meristematic elements, which have continued their growth to
the enlarged size, or cells continue their growth longer and more in-
tensively, or cells of permanent tissue are concerned, which take up
anew the process of growth in size. The cell may enlarge in all of its
dimensions, so that the original shape of the cell is maintained, or it may
enlarge in one or two directions, when the original shape is no longer
kept. If the enlargement is in two directions the cell will be distorted, if
in one direction it will grow abnormally long. The extent of the en-
largement and its direction will be determined by the character of the
surrounding cells, or their absence. An hypertrophied cell may be
surrounded by cells incapable of distention, hence its enlargement will
be limited to the size of the available free space. Kiister distinguished

two kinds of hypertrophy, cataplastic and prosoplastic. Cataplastic
hypertrophy is an abnormal increase in the volume of cells associated
with degenerative atrophy of their living contents, for the functional
decline of the cell has been termed by Beneke, cataplasia. Prosoplastic
hypertrophy involves new anatomic characteristics and functional
activities, for the cells store up fats, proteins and starches, or develop
chlorophyll, or red coloring matter. The involution forms of Bacillus
radicicola,which forms the leguminous root tubercles, and those of
the crown-gall organism, Pseudomonas turn efac tens, are examples of
simple hypertrophied cells (Fig. 143). With these preliminary remarks
it is important to illustrate the different kinds of hypertrophy which
have been described by plant pathologists. The most simple cases are
those in which the meristematic cells capable of division have grown to
364
an abnormal size by the omission of cell division. Under the influence

of a fungous parasite, Chytridium sphac ell arum, the apical cells of the
lateral branches of an alga, Cladostephus spongiosus, stop dividing and
enlarge into club-shaped swellings at their upper end. If specimens of
Padina pavonia, a siphonaceous alga, be inverted and are exposed to

Etiolated plants afford interesting examples of hypertrophy, for

in the absence of light the internodes of the stems and the petioles
of the leaves become inordinately long. If this follows cell divisions,
then it is a hyperplastic phenomena, but where it is due to the abnormal
lengthening of existing cells, it is a simple case of hypertrophy. Kiister
found in the etiolated peduncles of Tulipa Gesneriana, that the cells
were from a third to a half longer than the normal ones. Longer cells
than usual are produced in plants grown experimentally in moist air.
Hyperhydric abnormal and are formed by an excess of

tissues are
water within the plant. They
constitute a homogeneous group from
a causative (etiologic) point of view. As examples may be cited
the spongy white masses of cells which appear in the lenticels of the
twigs of alder, poplar, willow when such twigs are placed in water.
The individual cells of this porous tissue are chlorophylless, have a thin
layer of cytoplasmand a clear abundant cell sap. Such water lenticels
were compared by Schenck with typic aerenchyma found on numerous
water plants. Such lenticel excrescences arise from normal lenticels
by the enlargement of the phelloderm cells and in some cases the bark
cells lying under the lenticel hypertrophy. Von Tubeuf and Devaux
give extensive lists of the plants which produce hypertrophied lenticels.^
Bark excrescences form another kind of hypertrophied tissue.
They have been produced experimentally on the bark of the red currant,
Rihes aureum (Fig. 144). In such boss-like excrescences the paren-
chyma cells of the bark grow out into long sac-like cells of different
form and size by growth in a radial direction. Not only the cells of
the outermost bark layers take part, but all the elements down to the
wood take part in the abnormal growth and have become completely
or nearly colorless. The firm connection between bark cells is lost and
they are separated from each other by large intercellular spaces.
Sorauer kept cuttings of shoots of Ribes aureum several years old in a
vessel of water and in moist air. At the end of four weeks extensive
excrescences were formed.
Intumescences are small pustules, which are formed only in limited
areas, and their formation follows the same processes of growth as in
the case of bark excrescences. They are known in the branches of
Acacia pendula, Eucalyptus rostratus, Lavatera trimestris and Malope
^ KtJSTER, Dr. Ernst: Pathological Plant Anatomy, authorized translation by
Frances Dorrance, 1913: 74-75.
grandiflora. They are formed on the side of the branches exposed to

the sun and the bark cells are elongated in a radial direction, finally
breaking through the epidermis as spongy masses of cells. Leaves
also produce intumescences. Originating in the mesophyll cells, they
—
Fig. 144. Cross-section of a part of a strongly hypertrophied bark of Ribes
aureuni. K, Cork; P, periderm; H, abnormally elongated bark cells. {Kiisler,
Pathologische Pflanzenanatomie, 1903: 80.)
appear as greenish or whitish pustules of varying size and beneath the

cells lose their chlorophyll content. Cataplastic hypertrophy explains
the origin of some intumescences. For example, the lower cells of the
several-layered epidermis of Ficus elastica are pressed together by the
growth of the mesophyll cells and the space originally occupied by the
former is finally filled with the cells of the mesophyll. Excess of water
is one of the contributing causes in the formation of intumescences,
as also treatment of plants with poisons, especially copper salts.
Abnormal succulence, as an hypertrophy, is such where plants with
normally thin leaves, develop thick ones in their place. Salt solutions,
if used experimentally upon certain plants, may induce succulency.
LeSage produced artificial succulence in the leaves of Lepidium sativum
by abundant doses of common salt, NaCl. The mesophyll cells were
elongated greatly.
Fig. 145. — Cross-section through the wounded border of a cabbage leaf. The
hypertrophied mesophyll cells are enlarged into vesicular swellings. {Kilster, Palh-
ologische PJlanzenanatomie, 1903: 94.)
Callous hypertrophy arises after an injury when the living cells of an

organ enlarge without division, especially at the edge of the wound,
where they may enlarge to many times their normal volume (Fig. 145).
As it frequently happens that cell divisions follow an injury, it is not
always easy to distinguish between callous hypertrophies and callous
hyperplasias. We find callus hypertrophies among the thallophytes,
as in Padina pavonia, and in the higher plants where the bark, wood
parenchyma, leaves are affected. Kiister produced callous hyper-
trophies near the upper surface of the cut by keeping one end of the
cutting under water, the other extending into moist air. The bark
cells were enlarged greatly, producing ball-Hke or weakly lobed forms.
Only single cells in the bud hypertrophied and they grew out into large
colorless vesicles. Miehe has found Tradescantia virginica a suitable
object to produce callous hypertrophies experimentally. The destruc-
tion of cells, or cell groups, of the epidermis causes the formation of
empty places which are filled by the neighboring cells which close the
Fig. 146. —Pitted vessel of black locust, Fig. 147. —

Cross-section through
Robinia pseudacacia, filled with enlarged old wood Mespilodaphne sassafras.
of
parenchyma cells or tyloses. At a the con- The lower vessels contain .stone
nection between tyloses and original cell is tyloses, the upper besides stone
seen. (Kiister, Pathologische Pflanzenanal- tyloses, contain thin-walled tyloses.
omie, 1903: 100. {After Molisch in Kiister, Pathologische
Pflanzenanatomie, 1903: 100.)
opening. Haberlandt in his culture of isolated tissue elements obtained

abnormally large cells which should be classed among callous hyper-
trophies. He kept alive isolated mesophyll cells from the leaves of the
purple dead nettle, Lamiuni purpureiim, for weeks in Knop's solution,
or in nutrient sugars, and these cells grew perceptibly at the same time
that a thickening of their membranes took place. The exact causative
influence in the development of callous hypertrophies is still an open
question.
24
Tyloses^ are more or less closely packed, bladder-shaped intrusions

derived from the parenchyma cells adjoining the cavities of water-
conducting elements into which they project, often completely blocking
the cavities (Fig. 146). They were first investigated by Hermine von
Reichenbach, who noticed that the swelling is not cutofiF from the
parent cell by a septum. They arise frequently in association with one-

sided bordered pits, the limiting membranes of which undergo active
surface growth and thus push their way into the cavities of the vessels
(Fig. 147). Several tyloses may
from a single epidermal cell. They
arise
occur beneath branch scars that have been formed by a branch breaking
off and also at the wounded end of cuttings being formed in such
numbers, that they become flattened by mutual pressure. The cavities

of vessels are thus filled and they probably serve, as Boehm first sug-
gested, to plug up the cavities of the water-conducting tubes that have
suffered mechanic injury. This explanation suffices for such special
cases of injury, but tyloses are formed in uninjured vessels where they
obviously do not serve to close up a wound. Haberlandt believes that
tyloses of this last-mentioned type take some part in the process of
conduction, by increasing the surface of contact between the vessels
and the neighboring parenchyma cells. Kiister in his "Pathological
Plant Anatomy" gives a detailed account of the different kinds of
tyloses and their method which need hardly be discussed
of formation,
in a text-book for student use. Molisch gives a
list of plants in which
tyloses have been found. Sometimes tyloses fill the air chambers of
the stomata partially or almost entirely, where the epidermal cells
adjacent to the guard cells grow out into large unicellular bags, as in
Tradescantia viridis.
Gall hypertrophies are those which are produced by the effect of a

poison formed by an attacking animal, or plant. The tissue products
are the most diverse and a sharp distinction cannot be drawn between
hypertrophic and hyperplastic gall tissues. Gall hypertrophies usually
occur in the epidermal and the fundamental tissues of various plants.
The galls of the fungi belonging to the family Chytridiace^, namely,
those occasioned by species of Synchytrium, are very simple, for the
entire life history of the fungous parasite is passed in a single cell of the
1Gerry, Eloise: Tyloses: Their Occurrence and Practical Significance in Some

American Woods. Journal of Agricultural Research, i: 445-470, with 8 plates,
March 25, 1914.
1
host. The zoospores of the species of Synchytrium penetrate the epi-

dermal cells and growth causing their enlarge-
incite these cells to active
ment, as in the cells attacked by Synchytrium
drabcB. Sometimes the
infected cell grows inordinately and pushes the mesophyll cells lying
below apart, until it projects into the underlying cells as a spheric
pouch. If the neighboring epidermal cells are stimulated warts are
formed.
The second group of gall hypertrophies are certain hair-like develop-
ments of epidermal cells due to the irritation of certain mites of the
genus Phytoptus, which produce felt-galls, or Erineum. These erineum
structures arise in clusters on the surface of leaves of such trees as
maples, alders, birches, beeches, oaks, willows, limes and on herba-
ceous plants belonging to the genera Geranium, Mentha, Salvia, etc.
These outgrowths so resemble fungi, that Persoon was deceived into
so believing. They are usually pale, or even white at first, and they
turn brown aS the hair-like outgrowths die and lose their sap, but
since the latter may be colored yellow, red or purple, the outgrowths are
conspicuous objects on smooth leaves. The botanist Malpighi in
1675-1679 was the first to call attention to these galls. One-celled
erinea are the rule, but multicellular abnormal hairs are formed by the
hypertrophies of the normal trichomes as Frank reports on Quercus
(Bgilops.
Gall hypertrophies, where the ground tissues of plants participate in

their formation, are known. The roots of the Cycadace^ develop
sacs out of their parenchyma cells, so that large intercellular spaces are
formed which a blue-green alga, Anabcena cycadearum, the causal
in
organism, Galls produced by flies and belonging to the group of
lives.
zoocecidia may be taken as illustrations of gall hypertrophies. One is

known as the window gall of the maple, and the other is a reddish-brown,
bladder gall occurring on the leaves of Viburnum lantanum.
Multinuclear giant cells may be formed in plants, if the nuclei divide
regularly, but for some reason the formation of cross-walls becomes
impossible. The cells are stimulated to abnormal growth forming the
so-called giant cells. Such hypertrophies are associated with an in-
crease of the cytoplasmic contents of the cells. Such giant cells are
those produced by certain Nematode worms of the genus Heterodera on
such host plants as Beta, Coleus, Daucus, Plant ago and Saccharum (Fig.
148). Prilleux produced multinuclear giant cells in seedlings which
were cultivated at an abnormally high temperature. The number of

nuclei rarely exceeded three.
Multinucleate cells occur in crown gall which are perhaps compar-
able to the giant tells of the animal histologist. Cancer specialists have
divided these into two groups, viz., foreign-body giant cells in which the
Fig. 148.-— Cross-section of a part of a root gall of Circaa luteliana in old stage,
numerous giant cells are seen, the nuclei of which have begun to degenerate; b, irreg-
ularly branched nuclei out of the giant cells dividing by amitosia within anuceoli;
C, a single multinucleate giant cell. {After Tischler in Kuster, Pathologische Pflanzen-
anatomie, 1903: 128.)
stimulus is some introduced foreign substance, and genuine ones in
which no foreign bodies are visible. There is probably no real distinc-

tion other than that those occupied by parasites are malignant and those
induced by non-Hving granules are harmless. The cells in question in
crown gall are not very large, but they contain several nuclei (Fig.
149). Four nuclei in one cell is the most we have seen, but it is prob-
able that larger numbers occur. It would seem from the studies of
Erwin F. Smith, which, however, are incomplete, that most of the cell
divisions in crown gall are by mitosis. Frequently, however, there
have been found nuclei variously lobed and in process of amitotic
division, and this is probably the way in which several nuclei are
formed in one cell (Fig. 149).
—
Fig. 149. Nuclear division in crown gall; 1-16, cells showing amitotic (direct)
division; 17, mitotic division in which more chromosomes have passed to one pole
than to the other. (After Smith, Brown, McCulloch, Bull. 255, U. S. Bureau of Planl
Industry, 1912.)
HYPERPLASIA
Virchow in his " Cellularpathologie " (1858: 58) defined hyper-
plasia as allabnormal quantitative increase, produced by cell division,
and that definition will be adopted here. It is very difficult in practice
to distinguish without a careful study between hypertrophy and hyper-
plasia, but in the latter abnormalities are produced by cell division,
while in hy})crtiophy they are not. A number of well-defined groups of

vegetative hyperplasias may be distinguished by their etiology. Chemic
stimulation may be the cause of some, injury the cause of others. The
normal currents of foodstuffs may be clogged, the food may be irregu-
larly distributed and these interferences with normal processes may
result in proliferations and other abnormalities. Special stimuli may
also bring about abnormal supplies of food with consequent hyperplas-
tic tissue formation. The study of the abnormality to determine its
kind must be based on histologic analysis. If in our histologic examina-
tion, we discover that the abnormal tissues resemble the corresponding

normal plant parts, we are dealing with homooplasia; if they differ from
the normal, that is are composed of cells different from the correspond-
ing normal ones, then we have a case of heteroplasia.
Heteroplastic excrescences are of great interest histologically. The
difference between normal and abnormal states is sometimes greatly
diverse. This difference may be one of size, of tissue differentiation, of
constitution, and it is important in our pathologic study to determine
the nature of the differences between normal and abnormal conditions.
Thus, when we find a less differentiated tissue produced by abnormal
cell division without regard to the increase in the numbers of cells, we can
speak of the degeneration of tissue formation combined with an increase

of volume. This is known as cataplasy, and the products of the cata-
plastic processes as cataplasms and the kind of hyperplasia illustrated
in these abnormal changes as cataplastic hyperplasia. When, on the
other hand, we find new histologic characteristics and functional activi-
tiesassociated with hyperplasia, we speak of prosoplasy, of prosoplasms,
and of prosoplastic hyperplasia.
—
Homooplasia. This term may be defined as abnormal tissue forma-
tion produced by an increase of the normal elements; it has a limited
use to abnormalities, not to increase in size of normal organs by a mere
increase in the number of cells. We would not use the word homo-
oplasia for the unusually large leaves which of normal form and texture
appear on the shoots which arise from tree stumps and which have been
studied by the writer in a number of our American forest trees, such as
the tulip tree, Liriodendron tulipijera. Homooplasia is opposed to the
phenomena of giant growth here mentioned.
Localized tissue excrescences composed of the same histologic ele-
ments and of homooplastic character are not common. Occasionally
sugar beets continue their growth to abnormal thickness by the forma-

tion of ridge-like tissue excrescences composed of normal layers of tis-
sues which extend longitudinally. De Vries investigated a case where

new cariibial rings were formed outside of the latest ones of the first year
coincident with an arrestment of activity. Hottas incased roots of
Viciafaba in plaster casts pierced by holes. He found that by correla-
tivegrowth homooplastic excrescences filled the holes.
Some kinds of homooplasias are characterized by the fact, that only
single tissue forms of an organ are developed unusually without the for-
mation by which means the histology of the organ
of local excrescences
is altered. demand upon a tissue may result in the formation
Increased
of abnormally abundant tissue and to this the name of activity homo-
oplasias has been given. Various experiments have been conducted in
the attempt to form mechanic tissue by putting an increased mechanic
demand upon plant tissues. The experiments of Kiister with sunflower
stems were negative, as also those of Wiedersheim with branches of
beech and ash, for he found no strengthening of the hard bast in his
experiments. He proved, however, an increase of stereids in the
strained branches of Corylns avellana. Vochting has shown that hori-
zontal stalks of the Savoy cabbage strained at the extremity by hanging
weights developed thickenings on the upper side of the branch. De
Vries has described an abnormal potato tuber in which through the need
of conduction of plastic substances the bundles of the tuber had devel-
oped to an extent unusual to the normal plant. The wood and bast
portions were both increased. Vochting's experiments with potato
tubers supplement those of de Vries; for he succeeded in interpolating
the potato tuber as an element in the potato plants grown from it and
succeeded in getting hyperplastically developed vascular bundles.
Correlation homooplasias result when there is a local arrestment of
growth, and growth is started elsewhere with homooplastic changes in
the tissues. The experiments of Boirivant and Braun have proved this
in a number of plants. Only one case of callus homooplasia has been
reported and it is described by Schilberszy, who succeeded in stimulat-
ing an increase of vascular tissue in the stalks of Phaseohis mtdtiflorus
through injury. No positive cases are known where homooplasias
occur in the formation of galls.
Heteroplasias. —This term of pathologic anatomy is used when

there is a quantitative increase of an organ in which by abnormal di-
vision of the cells there are produced tissues, the single elements of which
have no resemblance to normal ones. Size of cells is of relatively little
interest in the study of these abnormalities. More important are cata-
plasmic and prosoplasmic tissues, which are formed in heteroplasia.
Cataplasmic tissues are those which are more simply constructed than
the corresponding normal tissues, while prosoplasmic tissues are those in
which we can see processes of differentiation in the formation of their
single cells and in the distribution of their different elements, which are
not manifest in the formation of the corresponding normal tissue.
The material illustrating the various kinds of heteroplasia may be
treated of under the following heads:
1. Correlation-heteroplasms 1
2. Calluses \ Cataplasms
Heteroplasias 3. Wound- wood J
4. Wound-cork
(a) Cataplasms
Galls
(b) Prosoplasms
I . Correlation-heteroplasms
This term is applied to cases where the normal growth of any plant
is arrested at its vegetative points by any causative factors whatsoever,
and where under the stimulus of the unused nutritive materials some part
of the plant develops abnormal growth and tissues. Vochting has
studied this subject in all of its details. He found that decapitation
of sunflower plants resulted in the production of tuber-like swellings
on the roots and that in the aerial runners of Oxalis crassicaulis filled
with reserve materials that removal of the apical cells and all axillary
bud cells resulted in the formation of swellings on the leaves and

internodes. According to Vochting, the parenchyma participates, also
the vascular bundles, which have fewer ducts than the normal ones.
The sieve tubes, however, are richly developed and extensive funda-
mental tissue outgrowths are found between bast and wood. The first
experimentally produced correlation-heteroplasms were made by Sachs.
He cut off all the vegetative points of pumpkin plants. He found, as a
result, that theembryonic root cells present in the stem at the right
and left of each petiole grow out into short-stalked tubers, as large
as marbles, in which the root cap and vegetative point are absent and
the axillary fibrovascular cord is resolved into a circle of isolated bundles

separated by chlorophyll-containing cells.
2. Callus
Callus may be defined in the widest sense of the word as all cell and
tissue forms produced subsequent to and as a result of injury. In
many plants and plant organs, only a metaplastic change of the cells
was incited by the injury (callus-metaplasia); in others, the cells laid
bare showed an abnormal growth and were changed into voluminous
vesicles and sacs (callus-hypertrophy), or an
increase of the normal tissue may result from
wound stimuli (callus-homooplasia). The
cells may be abundant after an injury owing
to active cell division and heteroplastic tissue
arises (callus-heteroplasia). When excres-
cences arise, which are composed of cells very
little differentiated and of the simplest form,
they are called cataplasms. If produced after
injury, they are found to differ greatly. The
tissues produced after an injury, if resembling
cork, are termed wound-cork, if similar to those
of wood, they are called wound-wood and
where we have the healing tissue composed of
nearly homogeneous parenchyma, it is called Fig. 150. — Longitudinal
section of a callused end of
simply callus. a cutting. C, C, Callus de-
Callous tissue may be formed as wound veloped from cambium; H,
wood; R, bark. (After
tissue in very different plant groups. It has Kiisler, p. 159.)
been found and vascular
in the algal fungi
cryptogams. The woody seed plants have been studied carefully as
to the formation of callus, because of its economic importance in forestry
and horticulture. Rose, poplar, or willow cuttings kept in moist air
and at a proper temperature after a few days form a ring-like tissue
excrescence from the cambium of the cut surface. This spreads out
rapidly and finally closes over the wound. Such rolls of tissue have
been called callus (callus, hard skin).
Callus at least in its first stages appears in the form of a ring, some-
times it is irregular in its formation, often being lacking in some places
and sometimes due to limitations of space relations. Sometimes

this is
the callus is most luxuriant, as in Cuttings of Populus pyramidalis
(Figs. 150 and 151) and Lamium orvala (Fig. 152), which produces the
largest callous rolls among herbaceous plants. All organs of the plant
Fig. 151. — Cross-section of a calloused end of a Fig. 153. — Stem of

poplar cutting. G, Vessel; M, pith ray. {After Lamium orvala with strong cal-
Krister, p. 159-) lous growth^ {After Kuster.)
are capable of producing callus, such as roots, stems and leaves, yet
allparts of all plants do not have the capacity of forming it. Such
growth seems to reside in the living elements of exposed tissue and the
productive power of different kinds of tissues varies greatly. Cam-
bium is the most active layer in the production of callus and next to
the cambium the primary and secondary bark tissues. The epidermis
plays an unimportant role. Pith also can develop callus.
The investigations of R. Hoffman, Kiister and Stoll go to show that
the cambial when division takes place after
cells injury are not re-
stricted to the mode of normal division but can grow in every direction.
It is certain, therefore, that the conditions of changed pressure are of
importance and significance, and yet this fact alone is hardly sufl&cient
to explain the phenomena of growth subsequent to an injury. The
cell divisions are very regular and rapid in those woody plants which
form callus.
Cuttings of woody plants, such as Populus pyramidalis (Fig. 150), if
placed in water and covered with a bell glass, so that the upper end
extends above the water into the moist air, shows early division of the
cambial cells near the upper wounded surface. We find these cells
are divided by walls perpendicular to their long axis, and in a lively
manner, by forming tangential walls, causing an abnormally intensive
growth in thickness of the cutting near the injured place. A strong
callus has been formed by abundant division of the cambial cells and the
cutting assumes a club-shaped form at its upper end. The wedge,
which is formed in this way between the wood and bast, has been termed
by Th. Hartig the "Lohdenwedge," which might be termed more ap-
propriately in English the healing wedge. In the formation of this
wedge, the cambial cells have divided just as under normal conditions,
but the relief of pressure has caused some of the outer cells to protrude
to form the enlarged part of the wedge with the outer cells bent
strongly. Primary bark as in Salix easily forms callus, and petioles and
leaves often form abundant callus.
Histologically the tissues of callus are distinguished by the slight
differentiation of their cells. The cushions of callus in many kinds of
cuttings are made up of the same kinds of cells and in a homogeneous
fashion. The cells are typically nascent ones with thin cell wall, pro-
toplasmic contents and a colorless cell sap. growth is slow, the
If the
callous cells are small and closely fitted together, but with rapid growth
the cells are large and loosely placed with conspicuous intercellular
spaces. Tracheids are absent from the upper cells of the cushion of
callus, but wedge some of the cells
in the lower part of the healing
assume the tracheal character. The formation of a tegumentary layer
is next to the development of tracheids the most interesting process of
differentation in the callus. The callus of poplar cuttings is favorable

for a study of its formation. The outer cells of the wedge of healing
are long and pouch-like, and their outer walls give the cork reaction,
since they take up Sudan III with avidity, and at the same time are
colored with hydrochloric acid and phloroglucin. Sooner or later, a
cork cambium is produced in the outer cell layers of most callous for-
mations. Massart, who first studied the nuclear phenomena in callous
tissue, rarely found that the cells contained more than one nucleus.
He found that direct nuclear division took place after wounding in
Cucurbita, Ricinus and Tradescantia, while Nathansohn found mitosis
in the callus of the divided roots of Vicia faba and both mitosis and
amitosis in that of poplar cuttings.
Conditions of Callous Formation
The behavior of cuttings from different plants varies within rather

wide limits. Some cuttings develop callus quickly, others slowly, and
the quality of the callous tissues differs as greatly. The poplar develops
a large amount of callus, while cuttings of elm, willow and oak form
only a low callus ring. Organs rich in foodstuffs form callus more
quickly than those poor in food materials. For example, the cotyle-
dons of Phaseolus and Vicia, rich in proteins and starch, develop callus
to an extraordinary degree. Moisture is an important factor in the
formation of callus, for it is formed in water, but better in moist air,
and not at all in dry air. Cuttings of poplar with both cut ends in
moist air develop callus at both extremities, but usually there is a
polarity shown. Cut-off petioles of the poplar form a more prolific
callus at the basal end of the petiole than at the end nearer the leaf
blade With stem cuttings, the callus is best developed at the basal
end in preference to the apical. Pieces of dandelion roots, 3 cm. long,
kept in a moist place, show most abundant callus on the upper stem
ends and not at all, or only slowly at the apex, but in alfalfa a power-
produced at the root end and feebly at the sprout
ful tuber-like callus is
end. So that having varied the external conditions of their formation,
it becomes evident that internal conditions are active and these prob-
ably depend upon inequalities in the nutritive condition of the cut

parts and also on the direction of established sap flow.
Loosely connected with pathologic anatomy ^re the regenerative
1
processes ivhich result in the formation of the vegetative points of roots

and shoots following an injury. Following an injury in very many
woody plants, there is a formation of adventitious roots and adventi-
tious shoots which grow from vegetative points developed directly
from the permanent tissue of the wounded plant organs, but this opera-
tion is necessarily preceded by formation of callus and in some cases
the new vegetative points are developed directly from the callus.
Upon these functional operations depend the success of the horticultural
operations of the making and establishment of cuttings of roots, stems,
and leaves. A very large number of plants may be raised by means of
cuttings. Soft-wooded, or herbaceous cuttings having leaves are used
in many cases, the shoots being in a half-ripened condition, that is
neither too young nor too old, dry and woody. Such cuttings are
usually inserted in sandy or gritty soil, and most of the leaves are
stripped off to check transpiration of moisture. Several leaves are
retained, so that a certain amount of assimilation can be carried on to
induce callus formation.
WOUND- WOOD.
The wood, whichis formed on the surface of the exposed wood of
the stem and on the inner surface of the detached bast, is distinguished
from ordinary wood by its abnormal structure, and especially by the
shortness of its cells and the absence, or scarcity of vessels. Hugo de
Vries,^ who was the first to direct attention to this abnormality, called
such wood, wound-wood. Such abnormal wood is distinguished from
the normal xylem by its simple histologic character, and is to be added
to the list of cataplasms.
The difference between wound-wood and normal wood depends
upon whether its formation has been brought about by cross cuts into
the cambium, or by longitudinal wounds. In the latter, the wound-
wood is distinguished by a wide-celled structure and by more numerous
ducts than in normal wood, but the libriform fibers are less in evidence.
Hugo de Vries studied Caragana arborescens and proved that the
wound stimulus caused the formation of wound tissue 7 cm. from the
wound itself. The nearer the cells of the cambium are to the wound
the more cross walls are formed, so that the short-celled zone of the
1 DE Vries, Hugo: Ueber Wundholz. Flora, 1876: 2.
wound-wood is produced near the place of injury, the transitional forms

at a greater distance and then the long-celled zone, which is formed
from undivided cells of the cambium. The daughter cells of the cam-
bium of the short-celled zone form near the edges of the injured part,
a wound-wood composed of polyhedric fundamental tissue cells re-
sembling the medullary ray cells of normal wood, only a few of such ele-
ments develop into parenchymatous tracheids. The cells of the long-

celled zone retain the character of wood parenchyma, but between them
narrow vascular cells united into strand-like groups are formed, while
wood fibers and broad ducts are absent. Such formed elements have
been termed primary wound-wood by de Vries, and later, there occurs
the production of a secondary wound-wood in which the cells gradually
assume a normal form. Abnormal resin ducts are formed in wound-
wood and these ducts are often more numerous in abnormal wood than
in the normal.
Sometimes the wound-wood does not form definite stratified tissues.
Occasionally tracheid-like cells are found in the callus which become
united into ball-like groups separated from the normal wood. Wood
fibers, which have an irregular course, have formed the gnarled wood.
The pith may take part in the formation of wound-wood, for it is

highly capable of producing callus, and also from the ground tissue of
injured leaves. No definite outer form is characteristic of wound-wood.
Frost action may kill the cambium in places, and if the dead places are
surrounded by cushions of wound-wood, then we speak of frost canker.
Frost cracks are filled with wound-wood, which close up the wound
followed by the formation of a frost ridge. Such canker tissue may be
destroyed during a frosty spell and a new attempt to form Callus results
in the addition of new wound-wood to the old and frost cankers are
formed.
Sometimes without an injury, tissues resembling wound-wood are
formed by the activity of the normal cambium, or from a newly formed
independent cambium. Under some conditions, the parenchyma of
the medullary rays increases at the expense of the formed elements of
the wood, so that broadened medullary rays are formed. Fasciated
branches frequently show such broadened medullary rays. Tuber-like
gnarls are formed in fruit trees that have stone fruits, and also in
beech bark and the structure of gnarls has been investigated by
Sorauer, and the bark tubers of beech by Krick.
Wound-cork
Injury to different plant organs such as roots, tubers, rhizomes,

stems, leaves and inflorescences is followed by the formation of cells
in rows and adjacent to the place of injury. The walls of these new
cells react to sulphuric acid, chlor-iodide of zinc and Sudan III and the
application of such reagents demonstrates the formation of cork,
which has been termed wound-cork. It is developed generally on all
parts of the wound, and at its edges connects directly with the normal
membranes, thus closing the wound. The walls of wound-cork cells
are always thin and are often folded, and the cork cells thus formed are
larger than those of the phelloderm. A stem wounded by a knife cut
soon heals up unless disturbed. The cut cells die, while those next
below grow out as a result of the decreased pressure, giving rise to cork
cells. As the opposing cushions of callus close together, this cork is
squeezed between them and finally a shearing of the cork cells results
as the tips of callus close together and unite. The only external sign
of the wound is a slight ridge-like elevation beneath which are traces
of the dead cells and the cork trapped here and there beneath the ridge.
Normally, wound-cork closes over the broken surface of the scars
formed in the autumn by the fall ofthe leaf, which is actually occasioned
by the formation of a cork layer, which cuts off the leaf from the stem.
CHAPTER XXI
GALLS
Galls may be defined as all abnormal tissues produced by the action
of animal, or vegetal parasites. The great majority of galls arise either
through the growth of cells alone (gall hypertrophy), or by cell division
(gall hyperplasia). The number of galls constructed heteroplastically is
very large, exceeding the diverse gall hypertrophies. Galls of heteroplastic

origin occur in the most diverse kinds of plants and on all organs of these
plants. The term gall, or cecidium (cecidia), is applied to those varia-
tions in form which are caused by foreign organisms. In the forma-
tion of the cecidium, an active participation of the host plant is neces-
sary and the biologic connection between the host plant and the gall-
producing organism must be considered. Only those cases fall within
our purview in which abnormal tissues are produced.
Considered biologically and etiologically galls form a well-defined
group without, however, any one feature common to all. Even when
considering only gall hyperplasias, we will find no common characteristics
except that a production of heteroplastic tissue is involved in all. This
is either extraordinarily simple histologically, showing little or no dififer-
entiation, or there are specific differentiationswhich produce structures

entirely distinct from those of normal tissues. The first kind are cata-
plasmic galls, and the second kind prosoplasmic. Galls may be clas-
sified as to their morphologic characteristics, as well as by their histolo-
gic. They may be found on every part of plants, roots, stem, branches,
leaves, flowers and fruits and plants capable of producing galls belong-
ing to all groups of the plant kingdom.
The following descriptive terms for galls will serve as a rough clas-
sification of their morphologic forms. Connold^ gives an example of
each kind.
As to morphologic character, galls are: axillary, coalescent, con-
glomerate, cymbiform, elongated, globose, glossy, gregarious, hirsute
1 CoNNOLD, Edward T.: British Vegetable Galls, 1901: 24-25.
384
GALLS 385
imbricate, pedunculate, pilose, pubescent, pustulate, rugose, rosaceous,

and thickening of the
scabious, separate, sessile, solitary, spiny, rolling
leaf,upon the upper surface of the leaf, upon the under surface of the leaf,
upon the margins of the leaf. Some cecidologists would classify galls
by the causal animal or fungus, by the natural families of the host
plants, according to the situation of the galls upon the plant, according
to their modes of growth, etc. Anton Ke.rner in his "Natural History
of Plants" (translated from the German by F. W. Oliver) divides galls
into simple, where one plant organ is involved, and compound, where
several plant organs are concerned in their formation. The simple
galls he divides into (a) felt galls, (b) mantle galls and (c) sohd, or
tubicular galls.
Cataplasmic galls are often produced by the action of parasitic
fungi,which invade the interior of the plant after an infection by ani-
mals, which by their wanderings over the surface of the plant may en-
large the field of their stimulation. Domiciled organisms are the cause
of prosoplasms, where the extent of the field of stimulation remains the
same under all circumstances, and is effective only in certain phases of
the development of the host plants.
The etiology of galls is of great interest. Malpighi in his " Anatome
Plantarum" published 1675-79 attributes the formation of insect
in
galls to the action of a poison excreted by the gall insect. Darwin and
Hofmeister explained galls, as the action of different kinds of poisons.
The stimuh, which cause the formation of galls, is undoubtedly chemic,
some unknown substances excreted by the causal parasite, excite
the cells of the host plant to growth and cell division and to different
kinds of differentiation. We know nothing definite about the chemic
substance, nor have the attempts to produce artificial galls been suc-
cessful. Traumatic stimuh, too, must come into play, for injury to the
plant goes hand in hand with infection, for the first stage of the develop-
ment of galls resembles callous tissue. The galls produced may be due
to plants, phytocecidia, or to animals, zoocecidia. The fungiand a few
flowering plants are largely responsible, while dipterous and hymenop-
terous insectsand mites are gall-producing animals.
(a) —
Cataplasms. Cataplasmic galls are those which are distin-
guished from the normal tissue of the corresponding organs by the small
amount of their tissue differentiation. The cell elements may often be
abnormally large, and the union of these elements usually forms a thin-
Fig 153. — Tubercles bean produced by inoculation. {After Moore, Geo.

of velvet T.,
Yearbook. Depi. Agric, 1902, pi. xxxvii.)
GALLS 387
walled often homogeneous parenchyma, while in other cases the cata-

plasms are marked by the absence of any definite form, or size. Almost"
all phytocecidia, or plant-induced galls, are cataplasmic. The swell-
ings on the roots of various members of the CRUCiFERiE caused by the
slime mould Plasmodiophora hrassiccB are of this nature. It is known
as Hanbury, clubroot, finger- and- toes by the practical grower of

plants. Root nodules, or tubercles, are produced on the roots of legu-
minous plants by bacteria (Figs. 153, 154, 155, 156), which can utilize
Fig. 154. — Cross-section of root tubercle of Lupinus angiislifolius containing bac-

teria, X 46. {After Moore, Geo. T., Yearbook Dept. Agric. pi. xxxviii, 1902.)
free atmospheric nitrogen and by their activity the leguminous plants

secure large amounts of nitrogen. A species of Actinomyces, or ray
fungus, is probably the cause of the mycodomatia of Myrica. Bacteria
also cause tumors on the Pinus halepensis and Oka europcea, on the latter
in the nature of a crown gall suggested to be somewhat like animal
cancer (Figs. 157, 158, 159). Recently Erwin F. Smith in relation to
the abnormal multiplication of the tissues which result in a crown-
gall tumor, or hyperplasia, concludes that the removal of growth inhibi-
tions is brought about by the physical action of substances liberated
within the tumor cells as the result of the metabolism of the im-
prisoned bacteria iPseudomonas tumefaciens) .Growth of the tumor
comes about not by the direct application of stimuh, but indirectly
by the removal of various inhibitions. Under normal conditions the
physiologic brakes are on at all times, more or less, in both plants
and animals, and only when they are entirely or largely removed in
GALLS 389
form cataplastic plant galls. Galls are due to species of Synchytrium,

to the aecidial stage of the rusts on violets, barberries, nettles and buck-
FiG. 156. —Longitudinal section through red clover rootlet, showing formation
of tubercle, a, Rootjhairs; 5, normal root parenchyma; c, vascular tissue; </, infected
areas showing infection strands; e, growing cells of tubercle. (Fig. 44, page 95,
Schneider, Pharmaceutical Bacteriology, 1912.)
%
GALLS 391
thorns. Branches of Vaccinmm enlarged by Calyptospora

vitis-idcBa are
Goeppertiana and those of Juniperus and Chanicecyparis by rusts of the

genus Gymnosporangiiim. Various species of the genus Exobasidium
produce soft, edible galls. All such galls are mycocecidia (Fig. 84).
Various algae, such as Cystoseira opuntioides, C. ericoides, and Ecto-
carpus Valiantei, live parasitically and cause tissue excrescences, while
the higher plants, especially of the family Loranthace^, produce large
gallsand the so-called wood roses on their host plants. These wood
roses areformed by the woody tissues of the host forming a ridge-like
growth about the clasping part of the parasite.
The animal-produced galls known as
zoocecidia aresome of them of cattiplastic
nature and are caused by nematode worms,
insects and mites. The most important
nematode worm responsible for the forma-
tion of galls is Heterodera radicicola, which
attacks many cultivated plants both in the
greenhouse and in the open. The mite
galls include the fleshy (hyperplastic) curl-
ings of the leaf edges, shoot tips of various
woody plants. Erineum galls, consisting of
multicellular cones and ridges, are to be

placed here. Dipterous insects produce
galls with a prosoplasmatic structure, while
the cataplasms produced by them have the
form of fleshy curlings of the edges of the
leaves of the host plants. Galls are produced Fig. 158. — Crown gall on
also by the attack of bugs, aphids, or plant raspberry. {After Conn, a gri-
,, 1
cultural Bacteriology,
,
^06.) -p.
lice, leaf wasps and gall wasps. They are
found on roots, stems, leaves, inflorescences and fruits. Such are
those on the roots of the grape due to Phylloxera vastatrix, etc.
—
Histology of Cataplasms. Usually aside from the slight tissue
differentiation cataplasms are composed of abnormally large cells with
an abundant protoplasmic content and sometimes with red cell sap,
also a large starch content. The primary and secondary tissues are
both involved in the formation of the galls.
—
Primary Tissues. ^Leaves, which are infected by fungi on which are
formed mycocecidia, show an arrestment of the tissue differentiation.
For example, the distinction between the pahsade and spongy paren-
chyma is often lost, because the palisade layer is not formed as such
and sometimes the spongy parenchyma undergoes a rich proliferation
Fig. 159. Section uf tobaccu. Margin of infected needle wound. Tumor in

middle part of back parenchyma; sieve tubes at x. (After Smith, Broiun, McCulloch,
Bull. 255, U. S. Bureau of Plant Industry, 1912, pi. cl.)
and red pigment sometimes appears. The same failure to form the regu-
by the zoocecidia. The vascular and mechanic
lar tissues is displayed
tissues may also undergo the same reductions in cataplasm, as do the

assimilatory tissues, so that the vascular bundles in infected parts are
GALLS 393
often only moderately developed. Wakker describes the disappear-

ance of the coUenchyma in the stalks of Vaccinium vitis-idcBa infected
with Exohasidium. Hyperplastic excrescences may be found by the

pith as in branches of Clematis attacked by jEcidiuni Englerianum.
Secondary Tissues. — Under this head will be considered the products
of the cambium. The formation of galls may be due to the division of
the living derivatives of the already-formed annual ring, or as in wound-
wood, itsown cells are used in the production of the cataplasmatic tis-
sue. The wood and bark swellings formed by the attack of animals and
fungi may be clustered or knob-like and resemble the frost-induced
cankers or even the witches' brooms.
Abnormal wood found in many woody galls is induced by many
fungi belonging to the genera Dasyscypha, Gymnosporangium and Peri-
dermium, by insects, and by parasitic flowering plants. A character-
istic feature of such galls is the abnormal increase in the parenchyma,
produced by the division of the cambial derivatives, which give rise to
group of parenchymatous cells. Sometimes the cambial rays are
broadened, so that extensive continuous masses of parenchymatous
wood are produced. The same kind of tissue formation is seen in an
examination of mycocecidia and zoocecidia. The mycocecidia may be
illustrated by a brief consideration of the spindle-like, or ball-like,
woody induced on different species of Juniperus by forms of the
galls
genus of rust fungi, Gymnosporangium. In the diseased wood, the

difference between the spring and autumn wood is scarcely recognizable,
and the parenchyma occupies a relatively broad space. The cambial
rays in the parts of the branches infected by Gymnosporangium clavar-
iceforme, instead of being only two to ten cells deep, are often ten to sixty
cells deep and at least three cells broad. The woody gall of Gymnospor-
angium juniperi-virginiancB shows still broader cambial rays, when
viewed in tangential longitudinal section. According to the investi-
gations of Reed and CrabilV the cedar apple gall is a modification of
the leaf of the red cedar (Fig. i6o). The cedar leaf parenchyma makes
up the greater portion of the cedar apple with the fungous hyphae in
the intercellular spaces of the parenchyma cells.
The fibro vascular system of the gall is a modified continuation of the
1 Reed, Howard S. and Crabill, C. H.: The Cedar Rust Disease of Apples
Caused by Gymnosporangium Junipcri-Virginiana. Technical Bull. 9, Va. Agric.
Exper. Sta., May, 1915.
fibrovascular system of the cedar leaf (Fig. i6i). From, or near the
base of the cedar apple, where the vascular elements are much con-
torted, arise many branches, which extend radially almost to the cortex.
Harshberger^ has investigated the galls produced by two species of
Gymnosporangium on the coastal white cedar, ChamcBcyparis thyoides,
and Stewart^ has published an
account of the anatomy of
Gymnosporangium galls and
Peridermium galls.
There may be an over-pro-
duction of the wood paren-
chyma and the parenchymatous
elements may divide without
abnormal widening of the
annual ring. The production
of abnormal resin canals which
are always surrounded with
parenchyma illustrate this
point. Hartig produced an in-
crease of resin ducts in the dis-
eased areas of coniferous trees
infected by Ar miliaria mellea.
—
Abnormal Bark. Many gall
formations exist where exten-
sive bark excrescences are pro-
duced whereby there is an ab-
normal formation of paren-
Fig. 160. —
Unopened cedar apples on red
chyma. An examination of
cedar, Juniperus virginiana. (After Jones the galls due to species of

and Bartholomew, Bull. 257, Agric. Exper. Stat.
Gymnosporangium shows that
Univ. Wise, July, 1915.)
the bark and wood form excres-
cences simultaneously. Wornle found that in weak branches of
Juniperus communis a rust fungus, Gymnosporangium davaricBforme,
incited the bark to' form woody parenchyma.
1Harshberger, John W.: Two Fungous Diseases of the White Cedar. Proc.
Acad. Nat. Sci., Phila., 1902: 461-504, with 2 plates.
2 Stewart, Alban: An Anatomical Study of Gymnosporangium Galls. Amer.

Journ. Bot, ii: 402-417, October, 1915; Notes on the Anatomy of Peridermium
Galls, do, iii: 12-22, January, 1916.
GALLS 395
Witches' —
Brooms and Stag-head. The branches of the silver fir, the
and portions of stem of various species of plants are trans-
flowers, fruits
formed into witches' brooms, or stag-head by the action of fungi of the
genus Exoascus and in the silver fir by Mcidium elatinum. The shoots
—
Fig. i6i. Diagram of a longitudinal section of a cedar twig bearing a small
cedar apple in June, a, Epidermis of cedar leaf; b, sclerenchymatous layer; c, fibro-
vascular bundle; d, resin gland; e, parenchyma; /, parenchyma of cedar apple; g,
fibro-vascular system of cedar apple; h, cortex. (After Reed, H. S., and Crabill,
are annual instead of perennial and are always and branch out
sterile
into broom-like, or antler-like forms called thunder bushes by some.

{b) Prosoplasms.^ —
Those galls are included, as prosoplasms, which
do not have arrested tissue dififerentiations, nor in which callus tissues
are found, but in which new kinds of tissues are formed differing from
the normal and in which definite proportions of form and size normal for
the species are repeated in them. Therefore, prosoplasms display in
their external form, something independent and well defined from the
organs of the normal plant both internally and externally. Hyper-
plastic tissues of this sort have been found until now only in the excres-
cences caused by parasites and almost entirely those of the animal
world, which produce zoocecidia. Six different orders of insects are the
principal producers of galls and various fungi. They are as follows:
The Acarina, or Mites of diminutive size, produce galls of simple form
and structure.
The Diptera, or Flies, cause many prosoplasms. The galls produced
by the gall gnats, or gall midges, are very different in character and often
very complicated.
The Hemiptera, which include the aphides commonly known as
green fly and plant lice, also produce numerous usually simple proso-
plasms.
The Hymenoptera, or gall-wasps, produce striking galls on account
of their size, diversityand complexity of form external and internal.
The Coleoptera and Lepidoptera (Heterocera) are responsible for
relatively few galls, and if formed their structure is relatively simple.
There are several plant-produced galls, or mycocecidia, in which
there is a regular arrangement of certain elements such as the cells in
which anthocyanin is formed. Ustilago Treuhii causes the production
of canker-like excrescences on the stems of Polygonum chinense, which
consist of spongy, parenchymatous, wood tissue. The excrescences,
which develop from the canker swelling, are fleshy, succulent, easily
breakable, irregularly bent, cyhndric and often longitudinally furrowed
broadened at the top Hke the head of a snail. The fruit galls, which
represent the part which produces the spores of the fungus, are repre-
sented by this part of the gall.
Histology of Galls
Three types of abnormal cell divisions, connected with the formation
of galls, may be distinguished, according to the direction that the di-
vision takes. In the first type, the regular orientation of the trans-
verse partitions cannot be recognized in young galls. In the second
type, the cells divide usually in a plane perpendicular to the upper sur-
GALLS 397
face of the affected organ. The third type is where no definite direction
of cell division may be found.
The tissue material used in the formation of galls may be considered
from several viewpoints. Thomas asserts that only those tissues are
able to form galls which are attacked during development, or in other
words permanent tissue cannot form galls and this is certainly true of
prosoplasmatically formed galls, but with cataplasms there seem to be
exceptions, where callus has been formed from bark parenchyma several
years old. Definite experimental proof of the contested points cannot
be obtained, because all attempts with experimentally producing cecidia
have failed. It is certain, however, that many galls are produced from
completely undifferentiated tissue, that is,from the primary meristem
of the tips of shoots, or from callus tissue, but not from cells and tissues
with lignified walls. It has been proved that all living cells belonging
to the epidermis, the ground tissue, or the vascular bundle tissue, can
under certain circumstances participate in the formation of galls. The
fundamental tissue, or parenchyma, produces the largest mass of the
galls, and it should be remarked in passing that the pith, bark and
mesophyll cells often proliferate with astonishing luxuriance. If in
leaf galls, for becomes ten or twelve

example, the infected part of the leaf
times the thickness of the normal leaf, it is in nearly all case's the meso-
phyll which has been active, for in nearly all galls the tendency to form
parenchyma is striking. The epidermis is concerned only occasionally
in the formation of galls and the chlorophyll content of galls is scanty.
The comparison of galls with animal tumors has been made but in-
advisedly because with the exception of a diseased new formation of
tissue being involved and in the absorption of appreciable amounts of
foodstuffs from the fundamental tissue galls and tumors have little in
common. Galls in contrast to tumors are developed by a typic infection
growth. Mixed swellings occur in galls where epidermis, bark, meso-
phyll and other tissues unite to form an homogeneous whole while no
tumor is known, which consists of characteristic tissue zones of such
diversity as those of the galls of the dipterous insects.
CECIDIAL TISSUE FORMS

We are next concerned with a study of the different kinds of tissue
forms in galls and in their consideration we will treat first the two most
important, namely, the protective and nutritive tissues.
Protective Tissues. —The protective tissues of galls consist of the

epidermal, or covering tissues, and the stone cells which form part of
the mechanic tissue. The epidermal tissue will be considered as a pro-
tective tissue irrespective of its origin whether from the epidermis of the
host, or as a new formation. The outer epidermis of sac and walled
galls consists of relatively large, ofteti flat cells which have a cuticle of
moderate development. Occasionally this epidermis may consist of
more than one layer. A gall found on a Calif ornian oak Quercus Wisli-
zeni, has the outer walls of its epidermal cells and the upper part of the
side walls thickened so that the cell cavity becomes conic in shape (Fig.
162). Cork, as a covering for galls, is extremely rare. Wound-cork
is found occasionally in these galls, while bark is even rarer in a few
apterous galls.
Hair structures, or trichomes, are not unusual in galls. The

majority of prosoplasmatic galls are naked or only slightly pubescent
and some galls are entirely without any covering tissue.
—
Mechanic Tissue.- These consist of stereids (sclerotic, or stone cells)
or sclerenchyma fibers almost entirely and they surround the larval
chambers so that their occupants are protected from outside pressure,
or sudden blows. Lacaze-Duthiers called the stone cell tissues in galls
"couche protectrice." The arrangement of the stone cells, their
structure and their position in the gall tissues are of the greatest diver-
sity. In the majority of cases, the stereids are round, in other galls
they are angular, while in others, they are stretched like palisade cells
and stand perpendicular to the upper surface of the gall body similar to
those in many fruit and seed shells. Sometimes the sclereid cell is thick-
ened only on one side, the delicately walled part being outside as in the
galls of Andricus quadrilineatus and sometimes they are inside as in an
elliptic gall of the oak, etc. The walls of the sclereids may be pitted,
and, therefore, porous, while in other cases the pitting may be very
scanty and other peculiarities have been described by pathologists
who are intimately acquainted with the structure of galls.
Nutritive Tissues.- —The tissues of galls which are eaten by the animal
occupants of the different galls, or the contents of which are beneficial
to the larvaehave been termed by cecidologists nutritive tissues. The

position of these nutritive tissues in the galls and their contents must
be considered next. No gall is entirely without nutritive tissues and
these not infrequently form the largest part of the gall and in those
GALLS 399
formed by dipterous insects the nutritive layers are often sharply sepa-
rated from the mechanical tissue adjoining. The epidermis of the gall
may represent the nutritive tissue when it develops as an inner hairy
lining to the larval chamber. Albuminous substances are found in such
and small grains of starch, so
papilla, or hairs, as well as drops of fat
by abundant supplies of a rich pabulum.
that the larvae are surrounded
Nutritive parenchyma may be formed within the mechanic mantel and
here it is available to the larval occupant of the cell (Fig. 162). In
Fig. —
Cross-section of an un-
162. Fig. 163. —
Insect gall on scrub oak,
known on Quercus Wlslizeni. Ep,
gall Quercus nana, due to gall insect, Amphi-
pidermis; Mi, outer mechanic mantle; bolips ilicifolia with interior of gall.
St, starch-filled outer nutritive layer; Pine Barrens near Chatsworth, N. J.,
Ms, inner mechanic mantle. (After May 27, 1916.
Kiister, p. 252.)
Other cases, the food materials are stored outside the mechanic mantel,
and they become available only by the larvae breaking through the
stereid layer. The cells of the nutritive parenchyma are usually iso-
diametric, elongated and sac-like forms, or as delicate cell threads. In
the highly organized galls of the Cynipidae, the cells of the innermost
layers on which the larvae feed contain a cloudy dense cytoplasm in
which small fat globules are seen and this layer may be termed appro-
priately the protein layer. A starch layer lies outside of the protein
layer. Here the cells contain starch. Besides the nutritive bodies
just mentioned occur tannic substances and lignin bodies. The latter
are produced at corners where several cells come together as local
thickenings of the walls. It is improbable that this lignin is nutritive
in function.
Tissues of Assimilation.- — Almost all galls are characterized by the
almost entire absence of chlorophyll. In a few galls, if present, the
chloroplasts are small, twisted and feebly colored besides being extremely
scanty.
Vascular Tissues. —The tissue of galls is intimately associated with
the vascular bundles of the host plants on which the galls occur and
some are actually formed from the tissue of the vascular bundles. In-
side the galls the vascular strands are usually delicate cords both in
cataplasms and prosoplasms. Where they occur inside galls, we find
that their individual elements resemble those of the normal bundles.
In a few exceptional cases, as in the galls of Andricus albopundatus,
these are concentric bundles. The arrangement of the gall bundles
varies greatly for we find them in a circle, or they pass through the bark
of the gall as a delicate network.
Tissues of Aeration. —The structure of many galls is an open porous
one (Fig. 163). The gall parenchyma cells in some cases are star-
shaped, fitting together by their projections, so that large intercellular

spaces are formed. Stomata and lenticels constituting pneumathodes
are found in galls. The stomata, however, have lost their ability to
close and remain, therefore, permanently open. Lenticels are present

in some cases. The stomata and parts of the epidermis disintegrate and
large roundish lenticels develop beneath them. Perhaps this aerating
tissue enables the larva to get sufficient oxygen for its metabolism.
Anthocyanin is present in the cells of many galls, as their red cheeks
abundantly testify.
—
Secretions and Secretory Reservoirs. -The elements concerned with
secretion in the normal epidermis are present in galls in unchanged form,
or they are increased, richly furnishing the secretions which are asso-
ciated with gall formations. Less frequently new forms of secreting
cells and tissues are found in galls. Crystals of calcium oxalate are not
found usually in galls, but yet their entire absence is a rare feature.
In some cases, the crystals when present are associated with the stereids.
The presence of tannic bodies has been noted previously, and it
seems that the tannin is found in the cells of certain gall tissues. The
GALLS 401
outer cell layers in some of the galls produced by Cynipid.e is rich in
tannin, so that these galls have been used from time immemorial in
the tanning of leather and in the production of ink. Tannin balls occur
in the nutritive parenchyma of many galls and are devoured by the
larvie of the same.
BIBLIOGRAPHY OF GALLS
Adler, Hermann, Transl. b}- Straton, Charlks R.: Alternating Generations.

A Biological Study of Oak Galls and Gall Flies. Oxford, at the Clarendon
Press, 1894, pages 198.
AsHMEAD, W. H.: Proc. Ent. Spc. Am., new ser., 1881: ix-xx,
Galls of Florida.
xxiv-xxviii, 1885and x-xix. Trans. Am. Ent. Soc, xiv: 125—128.
BEUTENMtJLLER, WiLLiAM: Catalogue of Gall-producing Insects Found Within
Fifty Miles of New York City, with Descriptions of Their Galls, and of Some
New Specjes. Bull. American Museum Natural History, iv: 245-278, with
8 plates.
Beutenmuller, William: The Insect Galls of the Vicinity of New York City,
Guide Leaflet No. 16, American Museum of Natural History. Reprinted from
American Museum Journal, iv. No. 4.
CoNNOLD, Edward T.: British Vegetable Galls: an Introduction to Their Study,
1902, pages 312, with 130 plates.

Cook, Mel T.: Some Problems in Cecidology. Botanical Gazette, 52: 386-390,
November, 191 1; A Common Error concerning Cecidia. Science, new ser., 34:
683-684, Nov. 17, 19 II.
CosENS, A.: A Contribution to the Morphology and Biology of Insect Galls. Trans.
Canadian Institute, ix: 297-387, 1912, with 13 plates.
Darboux, G. and Hovard, C: Hilfsbuch fii.r das Sammeln der Zoocecidien, mit
Beriicksichtigung der Nahrpflanzen Europas und des Mittelmeergebietes.
Felt, Efhraim Porter: A Study of Gall Midges II. 29th Report of the New
York State Entomologist, 1913: 79-213, with 16 plates, Albany, 1915.
Howard, C: Les Zoocecidies des Plantes d' Europe etdu Bassin dela Mediterranee.
Description des Galles. Illustration. Bibliographic detaillee. Repartition
geographique.Index bibliographique, 2 tomes, Paris, 1908.
Kerner, Anton: Natural History of Plants, transl. by F. W'. Olh-er, ii: 518-554,
1895.
KtJSTER, Dr. Ernst: Die Gallen der Pfianzen. cin Lehrbuch fiir Botaniker und
Entomologen, mit 158 Abbildungen.
KtJSTER, E.: Pathologische Pflanzenanatomie, Gustav Fischer in Jena, 1903; Zweite
Auflage, 1916.
KtJSTER, E.: Pathological Plant Anatomy, authorized translation by Fr.xnces
Dorrance, 19 13-19 1 5.
L.\caze-Dltthiers, H.: Recherches pour servir a I'historie des Galles. Ann. Sc.
Nat. Eot., xii: 353, 1849; xiv: 17, 1850; xLx: 273, 332, 1853.
26
Magnus, Prof. Dr. Werner: Die Entstehung der Pflanzengallen verursacht durch
Hymenopteren, Jena, 1914.
Mayr, Dr. GustavL.: Die Mittel-Europaischen Eichen Gallen in Wort und Bild,
Berlin, 1907.
Osten-Sacken, C. R. von: On the Cynipidae of the United States and Their Galls.
Proc. Ent. Soc. Phil., I: 47, 62 (1861); IV: 380 (1865).
Ross, Dr. H.: Die Pflanzengallen (Cecidien) Mittel-und Nordeuropas ihre Erreger
und Biologie und Bestimmungs tabellen, 191 1.
RtJBSAAMEN, Ew. H.: Die Zoocecidien durch Tiere erzengte Pflanzengallen Deutsch-
lands und ihre Bewohner, Leipzig, 191 1.
Thompson, Millett Taylor: An Illustrated Catalogue of American Insect Galls,
published and distributed by Rhode Island Hospital Trust Co., executor in
accordance wdth the provisions of the will of S. Millett Thompson, edited by
E. Felt, 1915, pages 66, with 21 plates.
CHAPTER XXXII
MECHANIC DEVELOPMENT OF PATHOLOGIC TISSUES
Our study of plant pathology would not be complete without a brief
reference to the reactions which influence the genesis of the abnormal
tissues of diseased plants. The investigation of these questions is a
matter of recent development ever since prominence has been given to
the experimental methods of studying plant diseases and abnormalities.
Kiister gives considerable prominence to these problems in the second
edition of his " Pathologische Pflanzenanatomie" (pages 328-398),
where we have the last and most authoritative treatment of the subject.
As an important factor he mentions the reaction ability of the living
cells, both in normal cell division and with inequalities in cell division, for
it is recognized that unequal division of the dividing cells plays an im-
portant part in pathologic plant anatomy. The polarity of cells is

another important element to be considered by the pathologic anatomist,
for if by unequal division, there is produced a change in the polarity of
the cells concerned in such division, the tissues which arise from such
cells will show a different kind of differentiation.
Miehe has demonstrated the physiologic polarity of cells by plas-

molyzing the cells of a marine species of Cladophora. He found, after
the destruction of the continuity of the protoplasm from cell to cell by
plasmolysis, and the transference of the plant into a solution of deter-
mined concentration, that elongated filaments developed, and that
rhizoids developed from the basal pole of each of the cells. The epi-
dermal cells of the leaves of linden, Tilia platyphylla, when attacked by
Eriophyes tilicB develop long cylindric trichomes from the same pole
of each cell.
The reaction capabilities of the cells of different tissues are both

quantitative and qualitative. The cells of the epidermis, parenchyma,
sap bundles react differently and this is expressed in the formation of
intumescences, wound-cork and wound-wood out of them.
callus
The change is also a noteworthy feature in the
in the reaction of cells
study of abnormal plant structure. There is a difference between young
403
organs, tissues and cells, as expressed in the growth, plasticity and

processes of differentiation under the influence of the exciting cause, as
is evidenced in the formation and nutrition of galls comprehended
under the general head of cecidogenesis.
The recent study of the developmental mechanics of pathologic

tissues calls for an investigation of stimuli, and the reaction to stimuli
where every reaction presupposes a capacity for reaction and where the
cells of different tissues vary in this respect and no cell remains always
the same, but changes without any influence of the external world with
the age of the cell, as well as the fact that every reaction presupposes
previous conditions which permit the reaction to take place. Such
considerations as these introduce the student to the investigation and
terminology of Roux, as set forth in his " Terminologie der Entwick-
lungs Mechanik der Tiere und Pfianzen," 191 2, and to the work of
Vochting, Kiister, Klebs, Haberlandt, N^mec and others along experi-

mental lines. Correlation, Neoevolution, Neoepigenesis are terms with
which the pathologic student must become acquainted. He learns that
Osmomorphosis comprehends all osmotic and turgor influences which
determine the form and differentiation of cells and tissues; that mechano-
morphosis is where plant cells and tissues have been modified in develop-
ment by mechanic pressure and pull; that chemomorphosis is where
chemic influences are the determining factors in molding the form and
controlhng the differentiation process; that trophomorphosis is where
abnormal nutrition is influential locally in the transformation of plants.
The consideration of chemomorphosis shows us that we may deal
with known chemic bodies the action of which can be studied experimen-
tally, or we may be concerned with unknown chemic substances, as the
poisons injected into the tissues of a plant by the gall forms which pro-
foundly influence the formation of the gall tissues.
Trophic correlation, or trophomorphosis, exists between the parts of

a cell, as well as between the organs of a plant, or the tissues of the
organs. The action within the cell may be between the nucleus and the
cytoplasm, and its importance in pathologic plant anatomy has been
experimentally studied by Gerassimoff and N^mec. Gerassimoff's
research dealt with the influence of the size of the nucleus on the cyto-
plasm, while N^mec discovered that in chloralized roots of Viciafaba the
cellswith normal diploid chromosome content had didiploid and tetra-
diploid chromosome-rich nuclei, and that the greater the content of the
MECHANIC DEVELOPMENT OF PATHOLOGIC TISSUES 405
cell in nuclear material the greater becomes its volume. Equally re-
markable discoveries were made in an investigation of the action of tis-
sues and organs upon one another. Vochting has produced a bending
growth in the root of the kohlrabi by removal of the leaves of one
side of the plant, so that the development of the normal side was
markedly greater than that of the other. The same effect was secured
in the petiole of a compound leaf of Pklea mollis by removal of a lateral
leaflet and the result of this experiment is displayed in the accompany-
ing figure. Narcotics and the vitiation of the atmosphere by poisonous

gases inhibit growth in length. Mathuse figures the effect of removal
of the growing point of a plant in the promotion of superficial leaf
growth and other anatomic changes in the leaves of Achyranthes
VerschafeUii. Other experiments of a somewhat similar nature are
equally illustrative. Hardly a more important and inviting field of
research has been opened than that which has been revealed by the
investigation of the experimental plant morphologists, or the experi-
mental pathologic plant anatomists.
BIBLIOGRAPHY OF DEVELOPMENTAL MECHANICS

OF PATHOLOGIC TISSUES
BoRDNEE, J. S.: The Influence of Traction on the Formation of Mechanical Tissue
in Stems. Botanical Gazette, 48: 251, 1909.
BucHER, H.: Anatomische Veranderungen bei gewaltsamer Kriimmung und geo-
tropischer Induktion. Jahrbucher ftir wissenschaftliche Eotanik, 43: 271, 1906.
CowLES, H. C: A Text-book of Botany for Colleges and Universities, vol. ii, Ecol-
ogy, 1911.
Daniel, W.: Zur Kenntnis der Riesen- und Zwergblatter, Dissertation, Gottingen,
1913-
EwART, A. J. and Mason- Jones, A. J.: The Formation of Red Wood in Conifers.
Annals of Botany, 20: 201, 1906.
GoEBEL, K.: Organography of Plants (English edition), i: 206, 1900.
H.^BERLANDT, G.: Verglcichende Anatomic des assimiherenden Gewebesystems der
Pflanzen. Jahrbucher fiir wissenschaftliche Botanik, 13: 74, 1882.
Haberlandt, G.: Zur Physiologic der Zellteilung. Sitzungsber. Akad. Wiss., Ber-
lin, 1913, Nr. xvi.
Haberlandt, G. transl. by Drummond, Montagu: Physiological Plant Anatomy,
Macmillan and Co., London, 1914.

Hoffmann, R.: Untersuchungen iiber die Wirkung mechanischen Krafte auf die
Teilung, Anordnung und Ausbildung der Zellen beim Aufbau des Stammes der
Laub und Nadelholer. Dissertation, Berlin, 1885.
Hartig, R.: Das Rotholz der Fichte. Forstl. naturwiss. Zeitschr., 5: 96, 1896.
HiBBARD, R. P.: The Influence of Tension on the Formation of Mechanical Tissue

in Plants. Botanical Gazette, 43: 361, 1907.
Keller, H.: Ueber den Einfluss von Belastung und Lagelauf die Ausbildung der
Gewebe in Fruchtstielen. Dissertation, Kiel, 1904.
Kny, L.: Ueber den Einfluss von Zug und Druck auf die Richtung der Scheidewande
in sich teilenden Pflanzenzellen. Jahrbiicher fiir wissenschaftliche Botanik,
37: 55, 94, 1901-

KiJSTER, Ernst: Histologische und experimentelle Untersuchungen liber Intumes-
zenzen. Flora, 96: 527, 534, 1906.
KtJSTER, Ernst-. Aufgaben und Ergebnisse der entwickelungsmechanischen Pflan-
zenanatomie Progressus Rei Botanicae, 2: 455, 1908.
KiJsTER, Ernst: Gallen der Pflanzen, Leipzig, 191 1.
MiEHE, H.: Wachstum, Regeneration und Polaritat isoliertcr Zellen. Berichte der
Deutschen botanische Gesellschaft, 23: 257, 1905.
N£mec, B.: Studien iiber die Regeneration, 1905.
Newcombe, F. C.: The Regulatory Formation of Mechanical Tissue. Botanical
Gazette, 20: 441, 1895.
Nordhausen, Max: Ueber Richtung und Wachstum des Seitenwurzeln unter dem
Einfluss ausserer und innerer Faktoren. Jahrbiicher fiir wissenschaftliche
Botanik, 44: 557, 1907.
Pieters, A. J.: The Influence of Fruit-bearing on the
Development of Mechanical
Tissue in Some Fruit Trees. Annals of Botany, 10: 511, 1896.
Prein, R.: Ueber den Einfluss mechanischer Hemmung auf die histologische Ent-
wicklung der Wurzeln. Dissertation, Bonn, 1908.
Roux, Wilhelm: Der Kampf der Telle im Organisms, Leipzig, 1881.
Roux, Wilhelm; Terminologie des Entwicklungs mechanik der Tiere und Pflanzen
Leipzig, 19 1 2.
Schulte, W.: Ueber die Wirkung der Ringelung auf Blattem. Dissertation,
Gottingen, 191 2.
Simon, S.: Experimentelle Untersuchungen iiber die Entstebung von Gefassver-
bindungen. Berichte der Deutschen botanische Gesellschaft, 26: 364, 393,
1908.
Smith, L. M.: Beobachtungen iiber Regeneration und Wachstum aus isolierten
TeLlen von Pflanzen embryonen. Dissertation, Hallea S., 1907.

Snow, L. M.: The Development of Root Hairs. Botanical Gazette, 40: 12, 1905.
Strasburger, E.: Ueber die Wirkungssphare der Kerne und die Zellgrosse Histo-
logische Beitr age, 1893: 5.
Strasburger, E.: Die Ontogenie des Zelle seit 1875. Progressus Rei Botanicae,
i: I, 90, 1907.
Vochting, Hermann: Ueber die Bildung der Knollen. Bibliotheca Botanica, 4: 11,
1887.
Vochting, Hermann: Untersuchungen zur experimentellen Anatomie und Patholo-
gic des Pflanzenkorpers, Tubingen, 1908.
voN Schrenk, H.: Intumescences Formed as a Result of Chemical Stimulation.
Report Mo. Bot. Gard., 1905: 125.
WoRGiTzKY, G.: Vergleichende Anatomie der Ranken, Flora, 70: 2-25 etseq., 1887.
^
WoRTMANN, J.: Zur Kenntnis der Reizbewegungen, Botanische Zeitung, 45: 819,
1887.
Suggestions to Teachers and Students
The investigation of plant diseases in general is most important and

it should be approached from a number of standpoints.^ The teacher is
interested in it, because he desires to arrange the subject matter, so that

it may be presented in the laboratory and lecture course. The experi-
ence of the writer along these lines may be of service to other teachers,
and it is given, therefore, with some detail. Living plants should be
kept for experimentation along pathologic lines. The best plants for
this purpose will be determined by the locality, by their availability, by
the ease of their cultivation and by their successful growth in the green-
house during the short days of winter. The experiments outlined in the
lessons of partIV can be tried upon these plants, such as the influence
of chemicals upon growth, the action of illuminating gas on the health of
the plant, and the extremely minute, or excessive action of amounts of
chemic reagents, for some experiments conducted by Free at Johns Hop-
kins University indicate that various plants react in a specific way to
extreme dilution of poisonous substances.
The plants can be wounded in various ways and on different organs.
The repair tissue can be studied by sectioning the healed part and stain-
ing with appropriate stains. Various infection experiments can be tried
with fungi and the lesions produced can be fixed and imbedded in paraf-
fin for sectioning, mounting, and for study later under the microscope.
The stock of such material for study can be increased materially by
collecting galls, insect depredations on plants, examples of callus for-
mation from street trees, which have been injured by horses biting off
the bark, orby abrasion with wagon wheels. This material, collected
from the streets and highways, from the woods and fields, should be
fixed and hardened and finally embedded in paraffin for sectioning and
microscopic study. These sections should be furnished along with
alcoholic, or dried material of the abnormal plant, so that the student
^
Cf. Shear, C. L.: Mycology in Relation to Phytopathology. Science, new,
ser., xli: 479-484, April 2, 1915.
Smith, E. F.: Plant Pathology. Retrospect and Prospect. Science, new ser.
xv: 601-612, April 18, 1902.

^Free, E. E.: Symptoms of Poisoning by Certain Elements, in Pelargonium
and other Plants. Contributions to Plant Physiology, The Johns Hopkins Uni-
versity, March 191 7; 195-198.
becomes familiar with the gross anatomy, as well as the microscopic.

Photomicrographs can be made readily by the use of the Edinger appara-
tus which has been used successfully at the University of Pennsylvania
in class work. It adds materially to the interest of the work to take
photographs of the sections studied and make permanent prints of the
diseased structures. After a few years, the alcoholic stock material
will have increased to such an extent that all phases of pathologic
plant anatomy can be demonstrated, not only by actual specimens,
but also by sections. The sections, if made directly by the sliding
microtome, can be kept in large numbers in small bottles in 50 per
cent, alcohol, where they are available for class use at any time. The
paraffin mounts can be kept in block form ready for use when required
by the sequence of laboratory exercises and lectures. If alcohol is not
available on account of its high price, other materials may be used in its
place.
The sections and alcoholic material having been prepared for use
can be studied for hypertrophy, for metaplasia, hypoplasia and other
pathologic conditions. Such an investigation presupposes a thorough
grounding in the technique of plant anatomy and histology, so that no
time may be wasted in unnecessary explanations. From the stand-
point of curriculum, such a course in mycology and pathologic plant
anatomy should be given in the junior, or senior years, or deferred until
the post-graduate years because of the special nature of the work.

Written reports should be required of all students based upon the
experiments with the inoculation and infection of various cultivated
plants and their reaction to various fungi. Similarly, where pathologic
anatomy and histology of plant organs and tissues are concerned photo-
graphic prints may take the place of microscopic drawings. Each
topic considered in the lecture course should receive attention in the
laboratory and in the field and indoor experiments, because this work
is designed to prepare future plant doctors, teachers and investigators,
who are interested in the science of phytopathology and who are
anxious to be proficient in the study of plant diseases.
Stock material should be kept of all the more common insect and
fungous diseases of cultivated and wild plants not only for such patho-
logic study, but also for a systematic and morphologic work with insect
and fungous parasites. The mycologic student should be able to
identify not only the more common insects and fungi after such a
course, but should be able also to diagnose the more common diseases
and suggest remedies in the form of insecticides, or fungicides, or other
remedial measures from a knowledge of the physiology of pathologic
plants. A change in the soil, or a change in the temperature and
exposure may be all that is needed to keep a plant in a perfect state
of health.
The problems which may be assigned to the post-graduate student
for experimental investigation are unlimited in America, where the
nation is confronted by serious pests introduced from various lands.
The anatomic and histologic characters and the development of cecidia
have been the subject of extensive studies in Europe, but American
botanists have done very little in the study of American galls along these
lines of investigation. The character of the poisons which cause
the stimulation of the plant to produce the galls is a matter well worth
the attention of botanists experimentally inclined. The equipment of
the laboratory and the facilities for experimentation should be con-
sidered before the problem is assigned to the post-graduate student.
The previous training and bias of the individual should be weighed
carefully for the research work may be of a cytologic nature. It may be
a histologic study pure and simple with pathologic tissues, or the prob-
lem may deal with prophylaxis, or preventive measures. It may be
that the student is better prepared to investigate the etiology of disease,
or the composition of sprays and their effects on the plant tissues.
Some advanced students would find keener zest in the systematic or
biologic study ofsome fungus, or group of fungi, or the bias may be
toward detailed experimentation with insects, or other forms of animal
life. The teacher should weigh carefully all of these details and act
accordingly. Problems with an economic bearing would be more suit-
able for the students of agricultural colleges and experiment stations,
while matters of pure science might be properly relegated to the
endowed colleges and universities, where investigation with a practical
trend would not be absolutely essential. The laboratory work should
be combined with field work in the study of inorganic and organic dis-
eases. The character of the field work will be determined by the
nature of the investigation and by the season and by the climatic con-
ditions. The work in the field at first would consist in the observation
of diseases, the taking of notes from the living trees and the collection of
material for more detailed study. The extent of the injury should be
4IO GENERAL PLANT PATHOLOGY
determined. Extension and the work of prevention can be carried on.

Cooperative work with the progressive farmers and horticulturists can
be inaugurated with profit to the farmer and the investigator. The
etiology of diseases can be investigated by properly directed field experi-
ments. Inoculations can be made on plants growing in the field, or in
the laboratory or greenhouse.^ Such original investigation presup-
poses the accumulation of apparatus and a suitable working library.
With the limited appropriation available for the purchase of apparatus
and books, such an equipment seems beyond the ordinary school and
college, but it will be surprising to those who have not tried the plan how
many books, diagrams, etc., can be accumulated, and how much
apparatus can be secured by spreading the purchase of such needful
things over a series of years. If the books and apparatus are cared for,
little deterioration need be suffered and at the end of twenty or twenty-
five years, a respectable stock of these desiderata will be on hand for use
in the class room, laboratory, research rooms and greenhouses.
The growth of the study of plant pathology as a distinct branch
of science has been by leaps and bounds. It is now on a more satisfac-
tory basis than ever before, and a larger number of men and women are
directing their attention to phytopathology as a life work. The men
who enter this field from now on must have a better and an all-sided
training. This presupposes an acquaintance with the literature of the

subject in his own and several foreign languages. There should also be a
training in chemistry and physics. He should know something about
zoology and should be conversant with the physiology and histology of
plants and other phases of botanic inquiry. To meet this demand our
American colleges and universities have introduced subjects which will
be of direct benefit to the future plant pathologist. The curricula^
have been arranged to introduce the study, hot only of plant pathology,
but also cognate subjects some of which may not have a direct bearing,
but which make the man a well-trained and a competent "plant
doctor."
iC/. Heald, F. D.: Field Work in Plant Pathology. The Plant World, lo:
104-109, May, 1907.
2 Fink, Bruce: A College Course in Plant Pathology. Phytopathology, II:
150-152, August, 1912. Consult Stevens, F. L.: Problems of Plant Pathology,

The Botanical Gazette, Ixiii: 297-306, Apr., 191 7; also Harshberger, John
W. The Need
: of Competent Plant Doctors, Education, 1895, 140-144.
PART III
SPECIAL PLANT PATHOLOGY

CHAPTER XXXIII
LIST OF SPECIFIC DISEASES OF PLANTS
The remarkable growth of the work of the United States Depart-
ment of Agriculture, and that of the agricultural experiment stations of
the different states, has been along the most diverse lines. Mycology
has been given prominence and the number of trained workers in this
field has increased to such an extent, that a separate organization,
known American Phytopathological Society, has been found
as the
necessary. The meetings
of this society have been largely attended
and the papers read have been of the greatest value and interest. The
organ of the society, "Phytopathology," has published already a con-
siderable number of important papers, and it has set a high standard for
the future work along mycologic and pathologic lines. One of the
specific problems, which it has attempted to do through special com-
mittes appointed for the purposes, has been to suggest the use of com-
mon names of fungous diseases based on recognized rules of procedure
and to prepare a list of the common and important diseases of economic
plants in the United States and Canada. The preliminary report of
the committee on common names has been made, but considerable
time must elapse before the list of common and important diseases is
completed.
As this book will be printed and issued before the preliminary list of
the American Phytopathological Society of fungous diseases appears,
it has been deemed advisable to compile a list from various sources of
information for the common host plants in the United States and
Canada, using the "Literature of Plant Diseases" given by W. C.
Sturgis in the Report of the Connecticut Agricultural Experiment
Station for the year ending Oct. 31, 1900, part III, pages 255-293, as
the basis of such a list.
411
+
412 SPECIAL PLANT PATHOLOGY
That the list might be made as complete as possible and repre-

sentative of the plant diseases of the United States and the tropic
countries to the southward, the following publications have been
used in its compilation.
Atkinson, Geo. F.: Studies of Some Shade Tree and Timber-destroying Fungi.
Cornell Univ. Agric. Exper. Sta., Bull. 193, June, iqoi.
CoiT, J. E.: Citrus Fruits, 1915: 364-402, The Macmillan Co.
Cook, Melville T.: The Diseases of Tropical Plants, 1913, The Macmillan Co.
DuGGAR, Benj. M.: Fungous Diseases of Plants, 1909, Ginn and Co.
Freeman, E. M.: Minnesota Plant Diseases, 1905.
Graves, Arthur H.: Notes on Diseases of Trees in the Southern Appalachians.
Phytopathology, III (1913) and IV (1914).
Heald, Fredk. D. and Wolf, Fredk. A.: A Plant-disease Survey in the Vicinity
of San Antonio, Texas. U. S. Bureau of Plant Industry, Bull. 226, 191 2.
Hesler, Lex R. and Whetzel, Herbert H.: Manual of First Diseases, xx
146 pages, 126 figs., 191 7, The Macmillan Co.
HuME,H. Harold: Citrus Fruits and Their Culture, 191 1: 466-492, Orange Judd Co.
Jackson, H. S.: Some Important Plant Diseases of Oregon in Biennial Crop Pest
and Horticultural Report, 1911-1912, Oregon Agric. E.xper. Sta., 177-308.
Longyear, B. O.: Fungous Diseases of Fruits in Michigan. Michigan State Agric.
Coll. Exper. Sta., Special Bull. No. 25, March, 1904.
Meinecke, E. p.: Forest Tree Diseases Common in California and Nevada. U. S.
Forest Service, A Manual for Field Use, 1914.
Reed, Howard S. and Cooley, J. S.: Plant Diseases in Virginia in the Years 1909
and 1910.
RoBBiNS, W. W. and Reinking, Otto A.: Fungous Diseases of Colorado Crop
Plants. Agric. Exper. Sta. Colo. Agric. Coll., Bull. 212, October, 191 5.
Selby, a. D.: a Brief Handbook of the Diseases of Cultivated Plants in Ohio,
Bull. 214, Ohio Agric. Exper. Sta., 1910.
Shear, C. L. and Wood, Anna K.: Studies of Fungous Parasites Belonging to the
Genus Glomerella. U. S. Bureau of Plant Industry, Bull. 252, 1913.
Smith, Ralph E. and Smith, Elizabeth H.: California Plant Diseases. Coll. of
Agric, Agric. Exper. Sta., Bull. 218, June, 1911.
Stevens, F. L. and Hall, J. G.: Diseases of Economic Plants, 1910, The Mac-
millan Co.
von Schrenk, Hermann: Some Diseases of New England Conifers. U. S. Div. Veg.
and Pathol., BuU. 25.
Physiol,
von Schrenk, Hermann: Sap-rot and Other Diseases of the Red Gum. U. S.
Bureau of Plant Industry, Bull. 114, 1907.
von Schrenk, Hermann and Spaulding, Perley: Diseases of Deciduous Forest
Trees. U. S. Department of Plant Industry, Bull. 149, 1909.
Whetzel, H. H. and Rosenbaum, J.: The Diseases of Ginseng and Their Control.
U. S. Bureau of Plant Industry, Bull. 250, 191 2.
This list will serve as an index of the diseases which will be described
LIST OF SPECIFIC DISEASES OF PLANTS 413
in full in the remainder of part III. As it will be impossible to describe

in detail all of the diseases of the list, number will be chosen,
a selected
which and which, if mastered by the student,
will illustrate the subject
will lay the foundation for a more thorough acquaintance with the
diseases, which are prevalent in the United States, and which the
student, the teacher, the horticulturist, the forester, the agriculturist,
and the practical mycologist are likely to meet in their plant-growing
experience. It is recommended that for each of the diseases described
in the following pages the outline for the use of students given in
Lesson 29 be used to facilitate an investigation of the disease in the
laboratory, greenhouse, or in the open field. This is a method of
study approved by the best teachers of the United States. ^ The author
wishes to state emphatically that he has designedly kept down the
number of diseases described in the following pages because the
thorough mastery of a limited number is better than a superficial study
of a larger list.
The general list precedes the descriptive pages of part III dealing
with a series of specific plant diseases, especially chosen because of the
author's familiarity with them, or because, they stand out prominently
as some of the more important diseases, which concern the American
plant-grower.
These specific diseases are divided into two groups. One group
includes the parasitic diseases due to fungi as the causal organisms. The
other group includes the non-parasitic, or so-called physiologic diseases
of plants. These have been treated in general in part II of this book,
but certain of the non-parasitic diseases have become of such general
interest that they merit a more detailed treatment. The literature of
these diseases is very much scattered, the only general account being
one published by Sorauer, Lindau and Reh in their "Handbuch der
Pfianzenkrankheiten" (3d Edition of Sorauer), 1908. This work is be-
ing translated by Frances Dorrance. Four parts of Vol. I have been
printed and the other parts will appear as fast as translated and printed.
The English edition beginning 1914 is entitled "Manual of Plant Dis-
eases." To this work the student of plant pathology is referred for
many details.
1 Whetzel, H. H. and Collaborators: Laboratory Exercises in Plant

Pathology, Ithaca, N. Y., 1916.
Parasitic Diseases of Plants
A LIST OF THE COMMON AND IMPORTANT DISEASES OF ECONOMIC

PLANTS IN THE UNITED STATES AND CANADA
Alfalfa
{Medicago saliva, L.)
Anthracnose {CoUetotrichum trifolii, Bain).

Journ. MycoL, Vol. XII, p. 192 (1906).
Bacterial Blight {Psendomonas mcdicaginis Sacket).
Bull. 212, Colo. Agr. Exp. Sta. (October, 1915).
Downy Mildew (Peronospora trifoliorum, de By.).
N. Y. Agr. E.xp. Sta., Bull. 305, p. 394 (1908).
Leaf -blotch {Pyrenopeziza medicaglnis, Fckl.).
Phytopathology 6, Abstracts of Columbus Meeting.
Leaf -spot {Pseudopesiza medicaginis (Lib.), Sacc).
Ibid., p. 384.
Root-gall {Urophlyciis alfalfa; (v. Lagerh.), Magn.).
Duggar, Fungous Diseases of Plants, p. 140 (1909).
Texas Root-rot {Ozonium omnivorum, Shear.)
Tex. Agr. Exp. Sta., Bull. 22 (1892).
Rust {Uromyces slriatus Schrot).
Bull. 218, Calif. Exp. Sta. (June, 191 1).
Iowa Bull. 131, p. 209 (April, 1912).
Violet Root-rot {Rhizodonia crocorum (Pers.) DC.).-
Phytopathology i, p. 103 (1911).
Winter Injury.
N. Y. (Cornell) Agr. Exp. Sta., Bull. 221, p. 6 (1904).
Almond
{Prunus amygdalus, Baill.)
Armillaria Root- Rot {Annillaria mellea, Vahl.).

Cal. Agr. Exp. Sta., Bull. 218, p. 1084 (191 1).
Crown-gall {P seudomonas tiimefaciens, E. F. Sm. & Towns).

Ariz. Agr. Exp. Sta., Bull. t,t, (1900).
Die-back [N on- par.).
Cal. Agr. E.xp. Sta., Bull. 218, p. 1086 (1911).
Rust {Puccinia pruni-spinosm Pers.).
Shot-hole (Cercospora circumcissa, Sacc).
Journ. MycoL, Vol. VII, p. 66 (1892).
Ampelopsis
Leaf-spot {Phyllosticla ampelopsidls, Ell. & Mart, Laestadia BidweUii (Ell.) V. & R.
and SphcEropsis hedericola (Speg.).
N. J. Exp. Sta., Rep. (1914).

Die-back {Cladosporium sp.).
N. J. Exp. Sta. Rep. (1914).

Apple
{Pirus mains, L.)
Anthracnose {Gleosporium malicorticis, Cordley; Ascigerous stage said to be Neofab.

rcea malicorticis (Cordley) Jackson, see Phytopathology 2: 94, 1912).
Oregon Sta., Biennial Rep., pp. 178-197 (1911-12).
Arsenical Poisoning.
Cal. Agr. Exp. Sta., Bull. 131 (1908).
Bark-canker (Myxosporium corticolum, Edg.).
Ann Myc, Vol. VI, p. 48 (1908).
Bitter-rot (Glomerella riifomaculans (Berk.) Spauld. & v. Schr.).'
Bitter-rot canker, U. S. Dept. Agr. Bur. Plant Industry, Bull. 44 (1903).
Black-rot {Sphceropsis malornm, Berk.).
Black-rot Canker.
N. Y. State Agr. Exp. Sta., Bull. 163 (1899).
Black-rot Leaf-spot.
U. S. Dept. Agr. Bur. Plant Industry, Bull. 121, p. 47 (1908).
Blight (Bacillus amylovorus (Burr.), Trev.).
Blight-canker, N. Y. (Cornell) Agr. Exp. Sta., Bull. 236 (1906).
Blossom- blight. Phytopathology, Vol. IV, p. 27 (1914).
Collar-blight, Penn. Agr. Exp. Sta., Bull. 136, p. 7 (1915).
Fruit-blight, Ibid., p. 20.
Twig-blight, N. Y. (Cornell) Agr. Exp. Sta., Bull. 329, p. 322 (1913).
Blister-canker {Nummularia discreta (Schw.) Tul.).
111. Agr. Exp. Sta., Bull. 70 (1902).
Blotch {Phylloslicta solitaria, Ell. & Ev.).
Blotch canker, U. S. Dept. Agr. Bur. Plant Industry, Bull. 144, p. 10 (1909).
Blotch leaf-spot. Ibid., p. 11.
Fruit-blotch, Ibid., p. 9.
Blossom End Rot (Alternaria sp.).

N. J. Exp. Sta. Rep., p. 471 (1914).
Blue Mold Rot (Penicillium spp.).
Stevens Diseases of Economic Plants, p. 94 (1913).
Brown-rot^ {Sclerotinia frucligena (Pers.) Schrot.).
Stevens and Hall; Diseases of Economic Plants, p. 92 (1913).
Canker (Pacific Coast) {Macrophoma curvispora, Pk.).
Stevens & Economic Plants, p. 83 (1913).
Hall, Diseases of
Common Rust {Gymnosporangium juniperi-virginlance, Schw.).
U. S. Dept. Agr., Rep., 1888, p. 376 (1889).
1 Consult Shear, C. L. and Wood, Anna K:
Studies of Fungous Parasites Belong-
ing to the Genus Glomerella. U. S. Bureau of Plant Industry, Bull. 252, 1913.
^For apple rots consult Phytopathology 4, p. 403, December, 1914, and Manual
of Fruit Diseases by Hesler and Whetzel.
6
Crown-gall (Pscitdomonas tumefaciens, E. F. Sm. & Towns).

U. S. Dept. Agr. Bur. Plant Industry, Bull. i86, p. 13 (1910).
Hairy-root, Ibid., p. 14.
Fly-speck {Leptothyrium pomi (Mont. & Fr.), Sacc).
Ohio x\gr. Exp. Sta., Bull. 79, p. 133 (1897).
Frost-blister (N on- par.).
N. Y. Agr. Exp. Sta., Bull. 220 (1902).
Frog-eye Spot {PhyUosticta pirina Sacc.)
Va. Rep., pp. 95-115, figs. 16 (1911-12).
Fruit-pit {Non-par.).
Bull., Torr. Bot. Club, Vol. XXXV, p. 430 (1908).
i Phyllachora pomigena (Sch-w.), SsLCC.
Fruit-spot {Phonia pomi, Pass. (Phytopath., Vol. II, pp. 63-72).
[Sphceropsis malorum, Pk.
Bull. 121, U. S. Bureau PI. Indst.
Leaf -spot (Frog-eye) {PhyUosticta pirina, Sacc).
R. I. Agr. Exp. Sta., Rep. 7, pp. 188-192 (1895).
Pink-rot {Cephalothecium roseum, Cda.).
N. Y. Agr. E.xp. Sta., Bull. 227 (1902).
Powdery Mildew {Podosphcera leucotricha (Ell. & Ev.) Salm. and P. oxyacantlicp {DC.)
deBy.).
U. S. Dept. Agr., Bull. 120 (1914).
Ripe-rot {Glcosporium fructigenum, Berk).
Journ. Mycol., Vol. VI, pp. 164, 172 (1891).
Root-rot {Armillaria mcllea Vahl).
Oregon State Biennial Report, pp. 226-233 (1911-12).
Scab {Venturia inaequalis (Cke.), Wint.).
N. Y. (CorneU) Agr. Exp. Sta., Bull. 335 (1913).
Mont. Bull. 96, pp. 65-102, pi. I, figs. 3 (February, 1914).
Scab {Fusicladimn dendriticiim (Wallr.) Fckl.)
Wash. Bull. 64, pp. 24, pis. 2, figs. 5 (1904).
Rusts {GymtiosporangimnJHniperi.virginiancB Schw. {Rmstelia pi rata (Schw.), Thaxt.);
G. globosum, Farl. {Ra'stclia laccrata, y, z. Thaxt.).
Scald {Non-par.).
Vt. Agr. Exp. Sta., Rep. 10, p. 55 (1897).
Scurf {PhyUosticta prunicola (Opiz), Sacc).
Stevens & Hall, Disease of Economic Plants, p. 78 (191 1).
Silver-leaf {Stercum purpureum, Pers.).

Sooty-blotch {Leptothyrium pomi (Mont. & Fr.), Sacc).
Spongy Dry-rot {VolutcUa friicti, Stev. & Hall).
Spray Injury {N on- par.).
Bordeaux injury, N. Y. Agr. Exp. Sta., Bull, 287 (1907).
Lime-sulphur injury, Ore. Agr. Exp. Sta., Research Bull. 2 (1913).
Stem-blight {Pseiidomonas mcdicaginis, Sackett.)

Col. Bull. 158, April, 1910, pp. 3-32; Bull. 159, pp. 3-15 (April, 1910).
Stem-rot {Schizopkylliim commune Fr.)
Bull. 218, Calif. Agr. Exp. Sta. (June, 191 1).
Volutella Rot {Voluidla fritcti, Stev. & Hall).
N. C. Agr. Exp. Sta., Bull. 196, pp. 41-48 (1907).
Water-core (Non-par.).
Phytopathology 3, p. 121 (1913).
Winter Injury (Non-par.).
Winter bark-splitting, Canada Exp. Farm. Rep., 1908, p. 112 (190S).
Winter black heart. Ibid., p. 113.
Winter bud-injury, Mont. Agr. Exp. Sta., Bull. 91 (191 2).
Winter crotch-injury. Me. Agr. E.xp. Sta., Bull. 164, p. 17 (1909).
Winter crown-rot, N. Y. Agr. Exp. Sta., Techn. Bull. 12, p. 370 (1909)
Winter die-back, Canada Exp. Farms, Rep., 1904, p. 108; 1908, p. 113.
Winter root-injurj^ Iowa Agr. Exp. Sta., Bull. 44, p. 180 (1899).
Winter sunscald, Canada Exp. Farm Rep., p. 112 (1908).
Apricot
(Primus armeniaca, L.)
Bacteriosis (Pseudomonas pritni, E. F. Sm.).
N. Y. (Corn.) Agr. Exp. Sta., Mem. 8 (1915).
Black-knot (Plowrightia morbosa (Schw.), Sacc).
Bull. 212, Colo. Exp. Sta. (October, 1915).
Blight (Bacillus amylovorus (Burr.), Trev.)
Colo. Agr. Exp. Sta., Bull. 84 (1903).
Blossom-rot (Scleroiinia lihcrliana, Fckl.).
Cal. Agr. Exp. Sta., Bull. 218, p. 1097 (191 1).
Brown-rot (Sckrotinia fructigena (Pers.), Schrt.).
Ibid.
California Blight (Coryneum Beijerinckit, Oud.).
Cal. Agr. E.xp. Sta., Bull. 203, p. 33 (1909).
Die-back (Valsa leucostoma (P.), Fr.)
Heald & Wolf, Plant Disease Survey, San Antonio, Tex. (1912).
Gummosis (Various causes).
Amer. Card., Vol. XIX, p. 606 (1898).
Shot-hole (Cylindrosporium padi, Karst).
Heald and Wolf, Plant Disease Survey, San Antonio, Tex. (191 2).
Arbor-vit.e
(Thuja occidental is, L.)

Die-back (Peslalozzia sp.)
N. J.Agr. E.xp. Sta., Rep., p. 517 (1912).
Root- rot (Polyporus Schiweinitzii, Fr.).
U. S. Dept. Agr. Div. Veg., Phys. & Path., Bull. 25, p. 23 (1900).
27
8
Ash
{Fraxinus sp.)
Decay, or Brown-rot {Polyporus sulphureus (Bull.), Fr.).

Heart-rot {Fomes fraxinophilus (Pk,), Sacc).
U. S. Dept. Agr. Bur. Plant Industry, Bull. 32 (1903).
von Schrenk, H., Diseases of Deciduous Forest Trees, U. S. Bur. of Plant
Industry, Bull. 149 (1909).
Leaf-spot {Cercospora fraxinites, Ell. & Ev.; Cylindrosporium viridis, Ell. & Ev., and
Septoria submaculata, Wint.).
Rust (yEcidium fraxini, Schw.).
Rep., Conn. Exp. Sta., p. 304 (1903).
Asparagus
(Asparagus offi-cinalis, L.)
Blight {Cercospora asparagi, Sacc).

Heald & Wolf, Plant Diseases Survey in Texas (191 2).
Rust {Puccinia asparagi, DC).
N. J. Agr. E.xp. Sta., Bull. 129 (1898).
Calif. Bull. 165, pp. 1-7, 18-9S, 98, 99, figs. 32 (Jan., 1905).
Aster, China
{Callistephus chinensis, Nees)
Rust [Coleosporium solidaginis (Schw.), Thiim).

Wilt {Fusarium sp.).
Mass. (Hatch) Agr. Exp. Sta., Bull. 79, p. 5 (1902).
Yellow (undetermined).
Ibid., p. I r.
Azalea
Rust {Piicciniastrmn minimum (Schw.), Arth.).
Conn. Exp. Sta., Rep., p. 854 (1907-1908).
Bamboo
{Phyllostachys henonis, Mitf. and P. quilioi, Riv.)
Smut (Ustilago Shiraiana, Henn.).

Patterson, Flora W. and Charles, Vera K., The Occurrence of Bamboo Smut
in America. Phytopath. 6, pp. 351-356 (1916).
Banana
(Musa spp.)
Trinidad Bud-rot {Bacillus miisce, Rorer.).

Phytopath. i, pp. 43-49 (191 1).
Ripe Fruit-rot {Gleosporium musorum, Cke. and Mass.).

Root Disease {Marasmius semiustus, Bri. & Cav.).
See Cook, Diseases of Tropical Plants, 1889, pp. 133-136 (1913).
Barley
{Hordeum sativum, Jess.)
Anthracnose {CoUetotrkhum cereale, Manns) = graminicola (Ces.) Wilson Phyto-

path, 4:110.
Ohio Agr. E.xp. Sta., Bull. 203, pp. 187-212 (1909).
Covered-smut {Ustilago hordei (Pers.), K. & S.).
Kan. Agr. Exp. Sta., Rep. 2, p. 269 (1890).
Nebr. Rep., pp. 45-53) figs- 4 (iQoT)-
Ergot (Claviceps purpurea (Fr.), Tul.).
So. Dak. Agr. Exp. Sta., Bull. 33, p. 38 (1893).
Leaf-rust {Puccmia simplex (Korn), Erikss. & Henn.).
Loose-smut {Ustilago nuda (Jens.), K. & S.).
U. S. Dept. Agr. Bur. Plant Industry, Bull. 152, p. 7 (1909).
Powdery Mildew {Erysiphe graminis, DC).
Bull., Ill State Lab. Nat. Hist, Vol. II, pp. 387-432 (1887).
Scab (Gibberella saubinetii (Mont.), Sacc).
Ohio Agr. Exp. Sta., Bull. 203, pp. 212-232 (1909).
Stem-rust {Puccinia graminis, Pers.).
U. S. Dept. Agr. Bur. Plant Industry, Bull 216 (1911).
Stripe Disease (Blade-blight) {H dminthos poritim gramineiim, Rabenh.).
Iowa Agr. E.xp. Sta., Bull. 116, p. 179 (1910).
Bull. 218, Calif. Agr. Exp. Sta. (June, 1911).
Bean
{Phaseolus vulgaris, L.)
Anthracnose {Colletotrichum lindemuthianum (Sacc. & Magn.), Bri. & Cav.).

N. Y. Agr. Exp. Sta., Bull. 48, p. 310 (1892).
Cornell Bull. 255, pp. 431-447, figs. 6 (May, 1908).
Mich. Spec. Bull. 68, pp. 12 (March, 19 14).
Bacterial-blight {Bacterium phaseoli, E. F. Sm.).
La. Bull. 139, pp. 43, pis. 6 (January, 19 13).
Leaf-spot {Cercospora canescens, Ell. & Mart.).
Heald and Wolf, Plant Disease Survey in Texas. (1912.)
Damping-off {Fungi spp.).
Pod-blight {Phoma subcircinata, Ell. & Ev.).
N, J. Exp. Sta., Rep., p.
472 (1914).
Rhizoctoniose {Corticium vagum, Bri. & Cav. var. solani Burt).
Rhizoctonia damping off, N. Y. (Cornell) Agr. Exp. Sta., Bull. 94, p. 266 (1895).
Rhizoctonia pod-spot, Science, new ser., Vol. XIX, p. 268 (1904).
Rhizoctonia stem rot. Science, Vol. XXXI,
new ser., p. 796 (1910).
Rust {Uromyces appendiculatus (Pers.), Lev.).
Southern Blight {Sderoiium Roljsii, Sacc).
Fla. Agr. Exp. Sta., Bull. 21, p. 27 (1893).
Bean (Lima)
(Phaseohis liinalus, L.j
Bacterial Blight {Pseudomonas phaseoU, E. F. Sm.).

Downy Mildew {Phylophthora phascoli, Thax.).
Conn. Agr. Exp. Sta., Rep. 167 (1889).
Beech
(Fagiis gnindifolia, Ehrh.)
Sap-rot {Polysticlus pergameniis, Fr.).

von Schrenk, H., Disease of Desiduous Trees, U. S. Bureau of Plant Industry,
Bull. 149 (1909).
White Heart-rot (Pomes igniarlus (L.), Gill.).
Beet
{Biia vulgaris, L.)
Bacterial Leaf-spot {Bacterium aptatum, Brown & Jamieson).

Journ. Agr. Research 190 (19 13).
i, p.
Cercospora Leaf-spot {Cercospora belicola, Sacc).
Nebr. Bull. 73 (1902).
W. and McKay, M. B., Relation of Stomatal Movement to Infection
Poole, V.
by Cercospora beticola, Journ. Agr. Research 5, pp. 1011-1038 (1916).
Crown-gall (Pseudomonas tumefaciens, E. F. Sm. & Towns).
Curly-top (undetermined).
U. S. Dept. Agr. Bur. Plant Industry, Bull. 122 (igcS).
Smith, Ralph E, and Boncquet, P, A.: Connection of a Bacterial Organism
with Curly Leaf of the Sugar Beet Phytopath. 5 pp. 335-342 (1915).
Damping-off (Fungi spp.).
Downy Mildew (Peronospora Schachtii, Fckl.).
1
Leaf-scorch (Non-par.).
N. Y. Agr. Exp. Sta. Bull. 162, p. 167 (1899).
Mosaic (undetermined).
Science, new ser. Vol. XLII, p. 220 (1915).
Phoma Crown-rot (Phoma beta: (Oud.) Fr.).
Frank Die Krankheiten der Pflanzen Zweitl. Aufl. 2, p. 399 (1896).
Journ. Agr. Research 4, pp. 135-168, pis. 11, pp. 169-177 (1915).
Phoma Leaf-spot (Phoma beta; (Oud.), Fr.).
Journ. Agr. Research 4, p. 169 (1915).
Phoma Root-rot (Phoma beta; (Oud.), Fr.).
Frank Die Krankheiten der Pflanzen Zweite. Aufl. 2, p. 399.
Puccinia Rust (Puccinia siibnilens, Diet.).
Rhizoctonia Root-rot (Cortkimn vagiim, Bri. & Cav. var. Solani Burt).
N. Y. (Corn.) Agr. Exp. Sta., Bull. 163, p. 34 (1899).
Rust (Uromyces beta;.).
Scab (Actinomyces chromogenes).
N. Dak. Agr. Exp. Sta., Bull. 4, p. 15 (1891).
Soft-rot (Bac'erium leutlium, Mete).
Nebr. Agr. Exp. Sta., Rep. 17, p. 69 (1904).
Tuberculosis (Bacterium beticolmn, E. F. Sm.).
Ct. Dept. Agr. Bur. Plant Industry, Bull. 213, p. 194 (1911).
Uromyces Rust (Uromyces beta; (Pers.), Lev.).
U. S. Dept Agr. Rep., 1887, p. 350 (1888).
Bermuda Grass
(Capriola daclylon (L.), Kuntze)
Leaf-spot (Helminthos poriiim giganleum, Heald & Wolf).

Heald and Wolf, Plant Disease Survey in Texas (191 2).
Birch
(Bclula spp.)
Decay (Fomcs fomenlarius (L.), Fr.).

Red Heart-rot (Fomes ftdvus, Fr.).
von Schrenk, H., Diseases of Deciduous Forest Trees, U. S. Bureau Plant
Industry, Bull. 149 (1909).
Sapwood Decay (Polyporus bclulini(s (Bull.), Fr.).
von Schrenk, H., p. 57.

White Heart-rot (Fomes iguiariits (L.), Gill).
N. Y. (Corn.) Agr. Exp. Sta., Bull. 193, p. 214 (1901).
Blackberry
{Riibus spp.)
Anthracnose {Gleosporium venetum, Speg.)-

U. S. Dept. Agr., Rep., 1887, p. 357 (1888).
Wash. "Bull. 97, pp. 3-18 (1910).
Cane-blight (Coniothyrium Fuckelii, Sacc).
Crown-gall {Bacterium tumefaciens, E. F. Sm. & Towns).
U. Dept. Agr. Bur. Plant Industry, Bull. 213 (1911).
S.
Double-blossom (Fiisarium ruhi, Wint.).

Del. Agr. Exp. Sta., Bull. 93 (191 1).
Gall {Pycnochytrium globosum, Schrot).
Late-rust (Kuehneola albida (Kiihn), Magn.).
Mass. (Hatch) Agr. Exp. Sta., Rep. 9, p. 74 (1897).
Leaf-spot (Septoria ruhi, Westd.).
Conn. Agr. Exp. Sta., Rep. 27, p. 309 (1904).
Orange-rust {Gymnoconia Peckiana (Howe), Tranz).
111. Agr. Exp. Sta., Bull. 29, pp. 273-300 (1893).
Box Elder
{Acer negiindo californicum (T. & G.), Sarg.).
Leaf-spot {Glwosporlumncgundinis, Ell. & Ev.).

Leaf-tip Blight {Septoria marginata, Heald & Wolf).
Leaf-blight Buckeye {Aiscnlus octandra, Marsh). {Phylloslicla cesculi, Ell. & Mart.)
Boxwood
{Buxus sp.)
Leaf-blight {Macrophoma Candollei (B. & Br.), Berl. and Vogl.).

Leaf and Stem Disease {Volutella buxi (Cda.), Berk.).
Buckwheat
{Fagopyrum esculenlum, Moench)
Leaf-blight {Ramularia rufomaculans, Pk.).

Descr., Conn. Agr. Exp. Sta., Rep. 14, 1890, p. 98 (1891).
Butternut
{Juglans cinerea, L.)
Leaf-spot {Gnomonia leptostyla (Fr.), Ces. & de Not.).

Mass. (Hatch) Agr. Exp. Sta., Rep. 10, p. 69 (1898).
BUTTONBUSH
{Ccphalanthiis occidentaUs, L.)
Leaf-blight {Cercospora perniciosa, Heald and Wolf).

Leaf-spot {Ramularia cephalanthi (Ell. & Kell.), Heald).
Cabbage
{Brasska oleracca, L.)
Bacterial Leaf-spot {Bacterium maculicoliim, McCul.).

U. S. Dept. Agr. Bur. Plant Industry, Bull. 225 (191 1).
Black-leg {Phoma lingam (Tode), Desm.).

Black-mold {Allcrnaria hrassicce (Berk.), Sacc).
U. S. Dept. Agr., Farmers' Bull. 488, p. 31 (191 2).
Black leaf-spot, Ibid.
Black-mold storage-rot, Ibid.
Black-rot {Pseudonionas campeslris (Pam.), E. F. Sm.).
Wis. Agr. Exp. Sta., Bull. 65 (1898).
Black-spot {Macros port urn brassiccB, Berk.). {Allcrnaria brassica, (B.), Sacc).
Va. Agr. Exp. Sta., Rep. (1909-1910).
Club-root {Plasmodio phora brassica, Wor.).
Journ. Mycol., Vol. VII, p. 79 (1892).
Va. Bull. 191, pp. 12, figs. 5 (Apr., 1911).
Vt. Bull. 175, pp. 1-27, pis. 4, figs. 6 (Oct., 1913)-
Damping-off {Fungi spp.).
U. S. Dept. Agr., Farmer's Bull. 488, p. 31 (191 2).
Downy Mildew {Peronos pora parasitica (Pers.), deBy.).
Ibid., p. 29.
Drop {Sclerotinia libertiana, Fckl.).
Mo. Bot. Card. Rep. 16, p. 149 (1905).
Leaf-spot {Cercospora Bloxami, B. & Br. (?)).
Heald and Wolf, Plant Disease Survey in Texas (191 2).
Root-rot {Corticium vagtini, Bri. & Cav., var. Sclani, Burt.).
Soft-rot {Bacillus carotovoruss, Jone.)
Journ. Science, new ser.. Vol. XVI, p. 314 (1902).
Yellows {Fusarium conglutinans Wollenw.).
,
Ohio Agr. Exp. Sta., Bull. 228, p. 263 (191 1).
Cacao
{Theobroma cacao, L.)
Bark Disease {Corticium javanicum, Ziram. = C. Zimmermanni, Sacc. & Syd.)

Diseases of Tropical Plants, pp. 180-191 (1913).
Black-rot {Phylophlhora Faheri, MaubL).

Exovin-rot {Thyridaria tarda, HsLncxoit).
Canker (Nectria theohromcB, Mass., and Caloneclrla jlavida, Massee)
Pink Disease {Corticium lilacofiiscum, Berk, and Curt.).
Root Disease {Macrophoma veslila, Prill & Del.).
Scabby-pod {Lasidoplodia theohromce (Pat.) Griff. & Maubl).
Seedling Disease {Ramularia nccator, Mass.).
Diseases of Tropical Plants, pp. 1 80-1 91 (1913).
Thread-blight (Marasmius equicrinus, Mull.).
Calla
{Richard la cthlopica, Spreng.)
Soft-rot {Bacillus aroidecd, Towns.).

Leaf-spot {Phyllosticta Richardia, Hals.).
Black-edge {Cercospora RichardlcBcola, Atk.).
Carnation
{Dianthus caryophyllus, L.)
Alternariose {Alternaria dianthi, Stev. & Hall).

A.nihxdiCno?,e {Volulella dlanthi, Xikin'i),
Descr., N. J. Agr. Exp. Sta., Rep. 14, 1893, pp. 385-386 (1894).
Bud-rot {Sporotrichum anthrophilmn, Pk.).
Nebr. Bull. 103, pp. 3-24 (Jan., 1908).
Leaf-mold or Fairy-ring {Helerosporium cchinulatmn (Berk.), Cke.).
Descr., N. J. Agr. Exp. Sta., Rep. 14, 1893, p. 386 (1894).
Die-back {Fusarium sp.).
Descr. Illus., N. Y. Agr. Exp. Sta., Bull. 164, pp. 219-220 (1899).
Leaf-spot {Septoria dianthi, Desm. and Heterosporlum cchinulatmn).
Bull. 218, Calif. Agr. E.xp. Sta. (June, 1911).
Descr., N. J. Agr. Exp. Sta., Rep. 14, 1893, pp. 384-385 (1894)-
Rust {Uromyces caryophyllmus (Schrank), Wint.j.
Descr. Illus., Gar. and For., Vol. V, pp. 18-19 (1892).
Treat., N. Y. Agr. Exp. Sta., Bull. 100, pp. 50-68 (1896).
Cf. N. Y. Agr. E.xp. Sta., Bull. 175 (1900).
Wilt {Fusarium sp.?).
Descr., Conn. Agr. Exp. Sta., Rep. 21, 1897, pp. 175-181 (1898).
.
Carrot
(Daucus carota, L.)
Root-rot {Corlicium vagum, Bfi. & Cav., var. Solani, Burt.)-

Rot {Phoma sanguinolenla, Grove).
Soft-rot {Bacillus carotovoriis, Jones).
Catalpa
{Catalpa hignonloides, Walt.)
Leaf-blight {Macros porium catalpce, Ell. & Mart.).

Descr. Illus., U. S. Dep. Agr., Rep. for 1887, pp. 364-365 (r888).
Treat, (rec), U. S. Dep. Agr., Rep. for 1887, p. 366 (1888).
Leaf-spot {Phyllosticta catalpce, Ell. & Mart.).
Descr. Illus., U. S. Dep. Agr., Rep. for 1887, pp. 364-365 (1888).
Soft Heart-rot {Polystictus versicolor (L.), Fr.).
Stevens, Neil, Mycologia IV, p. 263 (September, 1912).
Cedar
{Libocedriis; Thuya; Juniperus)
Leaf-pit {Keithia thujina, Durand).

Phytopath 6, pp. 360-363, 1916, on T. plicata.
Red-rot or "Pecky" Disease {Pomes carneus, Nees).
Descr. Illus., U. S. Dep. Agr., Div. Veg. Phys. & Path., Bull. 21, pp. 16-20 (1900).
{Gymnosporangium globosum, Farl).
{Gymnos porangiimt junipcri-virginiance, Sch w. )
Rust 1 Nebr. Rep. i, pp. 103-127, pis. 13, map i (1908).
{Gymnos porangi urn sabinm, Plowr).
Duggar, Fungous Diseases of Plants, pp. 425-426.
White-rot {Polyporusjunipcrinus, v. Schr.).
Descr. Illus., U. S. Dep. Agr., Div. Veg. Phys. & Path., Bull. 21, pp. 7-16
(1900).
\\'hitening {Cyanospora albiccdra:, Heald & Wolf).
Celery
{Apium graveolcns, L.)
Bacteriosis {Bacterium apii, Brizi).

Descr. Illus., N. J. Agr. Exp. Sta., Rep. 12, 1891, pp. 257-258 (1892).
Cf. U. S. Dep. Agr., E.xp. Sta. Rec, IX-9, p. 850 (1898).
Late-blight (Septoria pdroselini, Desm, var. apii, Br. & Cav.).

Oregon Sta. Biennial Rep., p. 273 (1911-12).
Calif. Bull. 208, pp. 83-115, pi. I, figs. 18 (Jan., 1911).
Leaf-blight {Cercospora apii, Fres.).
Descr. Illus., U. S. Dep. Agr., Rep. for 1886, pp. 11 7-1 20 (1887).
Treat, (pos.). Conn. Agr. Exp. Sta., Rep. 21, 1897, pp. 167-171 (1898).
Leaf-spot {Phyllosticia apii, Hals.).
Descr. Illus., N. J. Agr. Exp. Sta., Rep. 12, 1891, p. 253 (1892).
Leaf-spot {Septoria petrosclini, Desm., var. apii, Bi. & Cav.).
Descr. N. Y. Agr. Exp. Sta., Bull. 51, pp. 137-138 (1893).
Illus.,
N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 132, pp. 206-215 (1897).
Treat, (rec), N. Y. Agr. Exp. Sta., Bull. 51, pp. 139-141 (1893).
Rust {Puccinia bullata (Pers.), Wint.).
Century Plant
{Agave amcricana, L.)
Blight {Stagonospora gigantea, Heald & Wolf).

Plant Disease Survey in Texas (1912).
Cherry
{Priinus cerasus, L.)
Black-knot {Ploivrightia morbosa (Schw.), Sacc).

Descr. Mass. Agr. Exp. Sta., Rep. 8, 1890, pp. 200-210 (1891).
Illus.,
N. J. Agr. Exp. Sta.. Bui. 78. pp. 2-10 (1891).

Cf. N. Y. Agr. E.xp. Sta., Rep. 12, 1893, pp. 686-688 (1894).
Treat, (pos.), N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 81, pp. 646-653 (1894).
Fruit-mold {Sclerotinia cinerea (Bon.), Schrot.).
Ky. Agr. Exp. Sta., Rep. 2, 1889, pp. 31-34 (1890).
Mass. Agr. Exp. Sta., Rep. 8, 1890, p. 213 (1891).
Treat (pos.), N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 98, p. 409 (1895).
Leaf-curl {Exoascus cerasi (Fckl.), Sadeb.).
Descr., N. Y. Agr. Exp. Sta., Rep. 14, 1895, pp. 532-533 (1896).
Leaf-spot {Cylindrosporium padi, Karst., = Septoria cerasina, Pk.).
Descr. Illus., Scribner, Fung. Dis., p. 119 (1890).
Iowa Agr. Exp. Sta., Bull. 13, pp. 61-65 (1891).
Treat, (pos.), Iowa Agr. Exp. Sta., Bull. 30, pp. 291-294 (1895).
Leaf-spot {Cercospora cerasella, (Aderh.); Sacc).
Powdery Mildew {PodosphcEra oxycanth(B (DC), deBy.).
Descr. Illus., U. Dep. Agr., Rep. for 1888, pp. 352-356 (1889).
S.
Treat, (pos.), Iowa Agr. Exp. Sta., BulL 17, pp. 421-433 (1892).
Rust {Puccinia prtmi-spinoscB, Pers.).

Descr. U. S. Dep. Agr., Rep. for 1887, pp. 353-354 (1888).
Illus.,
Scab (Cladosporium carpophilntn, Thiim).

{Sclerotinia fructigena (Pers.), Schrot.).
Twig-blight
(Sderotinia cinerea (Bon.), Schrot.).
Chestnut
{Castanea dentata (Marsh.), Borkh.).
[
(Cylindrosporium castanicolum (Desm.), Berl.).
Anthracnose \
{Cryptosporiiim epiphyllum, C. & E.). { = Marssonia ochrolenca
[
(B. & C), Humph.).
Treat, (pos.), Amer. Gardening, Vol. XX, p. 559 (1899).
Blight {Endothia parasitica (Murrill), Anders. Hall).
Diseases of Economic Plants, p. 436 (igio).
Conn. Rep., pt. 5, pp. 359-453, pls- 8 (1912).
Leaf-spot (Marssonia ochrolenca, (Bri. & Cav.), Humph.).
Descr., Mass. Agr. Exp. Sta., Rep. 10, 1897, p. 69 (1898).
Sap-rot {Polysticlus versicolor (L.) Fr.).
Chrysanthemum
{Chrysanthemum- sinense, Sabine & C. indicum, L.)
Leaf-blight {Cylindrosporium chrysanthemi, Ell. & Dearn.).

Treat, (rec), N. J. Agr. Exp. Sta., Rep. 15, 1894, p. 369 (1895).
Ray-blight {Ascochyta chrysanthemi, Stev.).
Leaf-spot {Phylloslicta chrysanthemi, Ell. & Dearn.).
Occ, N. Agr. Exp. Sta., Rep. 15, 1894, p. 368 (1895).
J.
Leaf-spot {Septoria chrysanthemi, Cav.). {=S. chrysanthemella (Cav.), Sacc.)
Treat, (pos.), N. Y. Agr. Exp. Sta., Rep. 11, 1892, pp. 557-560 (1893).
Ray-blight {Ascochyta chrysanthemi, Stev.).
Rust {Puccinia chrysanthemi, Roze).
Occ, N. J. Agr. Exp. Sta., Circ, Nov. 15 (1899).
Descr., Treat, (rec), Ind. Agr. Exp. Sta., Bull. 85 (1900).
Cf. Gardening, Vol. VI, p. 277, '98.
Chives
{Allium schosnoprasum, L.)
Rust {Puccinia porri (Sow.), Wint.).

Conn. Exp. Sta., Rep., 1909-1910, p. 726.
Clematis
(Clematis spp.)
Anthracnose (GlcBosponumdcmatldis, Sor.).

Leaf-spot (Ascochyta clcmatidina, Thiim).
Journ. Agr. Research 4, pp. 331-342 (igiS)-
Root-rot (Phoma sp.)
Descr., N. Y. Agr. Exp. Sta., Rep. 3, 1884, pp. 383-384 (1885).
Clover
{Trifolium spp.)
Anthracnose (Collctolrichum Irifolii, Bain).

Damping-off (Pythiiim de Baryanum, Hesse).
Leaf-spot (Pscudopcziza IrifoHi (Pers.), Fckl.).
Leaf-spot {Phyllachora trifoln (P.), Fckl.).
Rust (UromycesTrifolii (Hedw. f.),Lev. and U. fallens (Desm.), Kern).
Descr. lUus., N. Y. (Corn. Univ.) x\gr. Exp. Sta., Bull. 24 (1890).
Iowa Agr. Exp. Sta., Bull. 13, pp. 51-55 (1891).
Phytopath. i, pp. 3-6 (February, 191 1).
Treat, (rec), N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 24, p. 139 (1890).
Sooty spot {Polythrincium Irifolii, Kze.).
Stem-rot {Sclerotinia trifoliorum, Eriks.).
Descr. lUus., Del. Agr. Exp. Sta., Rep. 3, 1890, pp. 84-88 (1891).
N. J. Agr. Exp. Sta., Rep. 18, 1897, pp. 314-318 (189S).
Treat, (rec), Del. Agr. Exp. Sta., Rep. 6, 1893, p. no (1894).
COCKLEBUR
{Xanthimn spp.)
Rust {Piiccinia xantkii, Schw.).

Coconut
(Cocos niicifcra)
Bud-rot {Bacillus coli, (Esch.) Mig.).

Johnston, John R., The History and Cause of the Coconut Bud Rot, U. S.
Bureau of Plant Industry, Bull. 228 (191 2).

Godaveri Disease {Pylkiiim palmivorum, Butler).
Cook, Diseases of Tropical Plants, pp. 197-206 (1913).
Leaf Disease {Peslalozzia palmarum, Cooke.)
Stem-bleeding {Thiclaviopsis ethacelicus, Went.).
Coffee
{Coffea arabica.)
Foot Disease {EiiryachoraUberica, Oud.)-

Porto Rico Bull. 17, pp. 29 (Feb., 1915).
Leaf-rot {Pellicularia koleroga, Cke).
Porto Rico Bull. 17, pp. 29 (Feb., 1915)-
Leaf -spot {Cercospora coffeicola, Bri. & Cav.).
Mancha de Hierro {Sphccroslilbc flavida, Massee).
Root Disease {Irpcx flaviis, Klotsch).
Rust {Hemileia vastatrix, Berk. & Broome).
Porto Rico Bull. 17, pp. (Feb. 29, 1915).
Stem Disease {Nccator dccretus, Mass.).
Corn
{Zca mays, L.)
Downy Mildew {Scleras pora macros pora, Sacc).

Leaf-blight {Ilelminthosporiiim inconspicuum, C. & E.).
Descr. lUus., N. Y. Agr. Exp. Sta., Rep. 15, 1896, p. 452 (1897).
Dry-rot (Diplodia zece (Schvv.), Lev. = D. maydis (Berk. Sacc).
Stevens & Hall, Diseases of Economic Plants, p. 335 (1910).
111. Bull. 133, pp. 73-85, 92-100, pi. I, figs.20 (Feb., 1909).
Rust {Puccinia sorglii, Schw.= P. maydis Bereng.)
Descr. Illus., U. S. Dep. Agr., Rep. for 1887, p. 390 (1888).
Cf. U. S. Dep. Agr., Div. Veg. Phys. & Path., Bull. 16, p. 65 (1899).
Smut {Ustilago zecB (Beckm.), Unger) and (U. Reiliana, Kiihn).
Descr. Illus., Kans. Agr. Exp. Sta., Bull. 62, pp. 179-189 & 198-201 (1896)
Ind. Agr. Exp. Sta., Rep. 12, pp. 99-112 (1900).
Treat, (rec), 111. Agr. Exp. Sta., Bull. 57, p. 335 (1900).
Wilt (Pseudomonas Stewarti, E. F. Sm.).
Treat, (rec), N. Y. Agr. Exp. Sta., Bull. 130, pp. 438-439 (1897)
Cosmos
(Cosmos hipinnahis, Cav.)

Stem-spot {Phlydana sp.).
Descr. lUus., N. J. Agr. Exp. Sta., Rep. 15, 1894, pp. 371-372 (1895).
Cotton
{Gossypium spp.)
Angular Leaf- spot (Bacterium malvacearum, E. F. Sm.).

Anthracnose (Colletotrichum gossypii, South worth);
Descr. Illus., Ala. Agr. Exp. Sta., Bull. 41, pp. 40-49 (1892).
U. S. Dep. Agr., Office Exp. Sta's, Bull. 33, pp. 293-299 (1896).
Ala. Bull. 153, pp. 27-33 (Feb., 191 1).
Boll-rot (Bacillus gossypina, Stedm.).
Descr. Illus., Ala. Agr. Exp. Sta., Bull. 55 (1894).
Treat, (rec), Ala. Agr. Exp. Sta., Bull. 107, p. 313 (1900).
Damping-off (Corlicium vagum, B. & C, var. Solani, Burt.).
Descr., Ala. Agr. Exp. Sta., Bull. 41, pp. 30-39 (1892).
Cf. Ala. Agr. Exp. Sta., Bull. 107, pp. 295-296 (1900).
Ala. Agr. E.\p. Sta., Bull. 41, pp. 58-61 (1892).
Leaf-mold (Ramularia areola, Atk.).
Descr. Illus., Ala. Agr. Exp. Sta., Bull. 41, pp. 55-58 (1892).
Root-rot (Ozonium omnivormn, Shear).
Descr. Illus., 2, 1889, pp. 67-76 (1890).
Tex. Agr. Exp. Sta., Rep.
Dep. Agr., Office Exp. Sta's, Bull. 2,3, P- 300 (1896).
U. S.
Treat, (rec), U. S. Dep. Agr., Office Exp. Sta's, Bull. 2,2,, p. 304 (1896).
Rust (Uredo gossypii, Lagerh.) and (Mcidiiim gossypii. Ell. & Ev.).
Descr., Journ. Mycol., Vol. VII, pp. 47-48 (1891).
Texas Root-rot (Ozonium omnivorum, Shear).
Smut (Doassansia gossypii, Lagerh.).
Wilt (Neocosmospora vasinfecta (Atk.), Smith).
Descr. Illus., Ala. Agr. E.xp. Sta., Bull. 41, pp. 19-29 (1892).
U. S. Dep. Agr., Div. Veg. Phys. & Path., Bull. 17 (1899).
Cow Pea
(Vigna catjang)
Angular Leaf-spot (Cercospora cruenta, Sacc).

Stevens and Hall, Diseases of Economic Plants, p. 395 (1910).
Leaf-spot (Amerosporium cecotunnicum, Ell. & Tracy).
Stevens & Hall, p. 394 (1910).
1
Rust (Uromyces appendiculatus (P.), Lk.).
Wilt (Neocosmospora vasinfecta (Atk.), E. F. Sm.)-
Cranberry
(Vaccinium oxycoccus, L.)
Anthracnose {Glomerella rujomaculans (Berk.) Sp. & v. Schr. var. vaccinii, Shear).
Gall [Synchytrium Vaccinii, Thomas).

Descr. lUus., N. J. Agr. Exp. Sta., Bull. 64, pp. 4-9 (1889).
Treat, (rec), N. J. Agr. Exp. Sta., Rep. 11, 1890, p. 2,2>3 (1891).
Hypertrophy (Exobasidium oxy cocci, Rost = Ex. vaccinii (Fckl.) Wor.).
Rot (Acanlhorhynchus vaccinii. Shear).
Shear, C. L., Bull. 10, U. S. Bur. Plant Industry.
"Scald" {Guignardia vaccinii, Shear).
Descr. lUus., N. J. Agr. Exp. Sta., Bull. 64, pp. 30-34 (1889).
Treat, (rec), N. J. Agr. Exp. Sta., Bull. 64, pp. 39-40 (1889).
Sclerotial Disease {Sclerotinia oxycocci, Wor.).
Spot {Pestalozzia Guepini, Desm., var. vaccinii. Shear).
Cucumber
{Cucitmis sativus, L.)
Anthracnose {Colletolrichum lagenarium (Pass.), Ell. & Hals.).

Descr. lUus., Ohio Agr. Exp. Sta., BuU. 89, pp. 109-110 (1897).
Treat, (pos.), N. J. Agr. Exp. Sta., Rep. 17, 1896, pp. 340-343 (1897).
W. Va. Bull. 94, pp. 127-138, pis. 5 (Dec. 2, 1904).
Bacteriosis or Wilt {Bacillus tracheiphilns, E. F. Sm.).
"Damping-off" or Seedling-Mildew {Pythium de Baryanum, Hesse).
Descr. lUus., Mass. Agr. E.xp. Sta., Rep. 8, 1890, p. 220 (1891).
Treat, (rec), Mass. Agr. Exp. Sta., Rep. 8, 1890, p. 221 (1891).
Downy Mildew {Plasmopara cubensis (Bri. & Cav.), Humphrey).
Descr. lUus., N. Y. Agr. Exp. Sta., Bull. 119, pp. 158-165 (1897).
Cf. Ohio Agr. Exp. Sta., Bull. 105, pp. 219-220 (1899).
Treat, (pos.), Ohio Agr. Exp. Sta., BuU. 105, pp. 223-229 (1899).
Leaf-glaze {Acremonium sp.).
Illus., Mass. Agr. Exp. Sta., Rep. 10, 1892, p. 230 (1893).
Leaf-spot {Phyllosticta cuciirbitacearum, Sacc).

Occ, Ohio Agr. Exp. Sta., Bull. 105, p. 222 (1899).
Powdery Mildew {Erysiphe polygoni, DC).
Descr. Illus., Mass. Agr. Exp. Sta., Rep. 10, 1892, p. 225 (1893).
Treat, (pos.), N. Y. (Corn. Univ.) Agr. Exp. Sta., BuU. 31, p. 138 (1891).
Scab {Cladosporium cncunierinum, Ell. & Arth.).

Descr. lUus., Ind. Agr. Exp. Sta., Bull. 19, pp. 8-10 (1889).
Mass. Agr. Exp. Sta., Rep. 10, 1892, pp. 227-229 (1893).
Stem-rot {Sclerolinia libertiana, Fckl.).
Descr. lUus., Mass. Agr. Exp. Sta., Rep. 10, 1892, pp. 212-224 (1893).
Currant
{Rlbes, spp.)
Anthracnose {Gloeosporium rihis (Lib.), Mont. & Desm.).

Descr. lUus., N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 15, p. 196 (1889).
Cane- wilt {Dothiorella).
Descr., N. Y. Agr. Exp. Sta., Bull. 167, pp. 292-294 (1899).
Cane-blight {Neclria cinnabarina (Tode), Fr.).
Descr. Illus., N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 125 (1897).
Knot {Pleonectria beroUnensis, Sacc).
Cornell Agr. Exp. Sta., Bull. 125 (February, 1897).
Leaf-spot {Septoria rihis, Desm., and Cercospora angulata, Wint.).
Descr. Iowa Agr. Exp. Sta., Bull. 13, pp. 68-69 (1891).
Illus.,
Treat, (pos.), Iowa Agr. Exp. Sta., Bull. 30, pp. 289-291 (1895).
Powdery Mildew {Sphcerotheca mors-itvce (Schw.), Bri. & Cav.).
Rust {Puccinia Ribis, DC).
See U. S. Dep. Agr., Exp. Sta. Rec, X-6, p. 559 (1899).
Wilt {BotryosphcBria ribis, Gross. & Dug.).
N. Y. Techn. Bull. 18, pp. 1 13-190, pis. 2, fig. i (July, 1911).
{Cronartium ribicola, Diet.), representing the uredo- and teleuto-stages of the
white pine blister rust, Peridermium slrobi, Kleb, a serious disease of
white pines against which a strict quarantine is maintained.
N. Y. State Techn. Bull. 2, pp. 61-74, pls. 3 (1906).
Cyclamen
Dark Leaf -spot (Phoma cydamena, Halst.).

Watery Leaf-spot (Glomerclla rufomaculans (Berk.), Spauld. & v. Schr., var.
cyclaminis, Patt. & Ch.).
Cypress
{Taxodium distichiim (L.), Rich.)
Leaf -blight {Pestalozzia funerea, Desm.).

"Pecky" Disease {Fungus indet.).
Dandelion
{Taraxacum officinale, Web.)
Leaf-spot {Ramtilaria taraxaci Karst.)-

Conn. Agr. Exp. Sta., Rep., p. 862 (1907-08).
Egg-plant
{Solanum meJongena, L.)
Anthracnose {Ghvosporiitm melongcnce. Ell. & Hals.).

Occ, N. J. Agr. E.xp. Sta., Rep. 12, 1891, p. 281 (1892).
Cf. N. J. Agr. Exp. Sta., Rep. 13, pp. 330-333 (1892).
Blight {Pseudomonas solanacearum, E. F. Sm.).
Descr. Illus., U. S. Dep. Agr., Div. Veg. Phys. & Path., Bull. 12 (1896).
Treat, (rec), U. S. Dep. Agr., Div. Veg. Phys. & Path., Bull. 12, pp. 23-24
(1896).
"Damping-off," or "Seedling-mildew" {Pylhium de Baryannm, Hesse).
Fruit-mold, Gray Mold {Botrylis fascicularis (Cord.), Sacc).
N. J. Agr. Exp.
Descr., Sta., Rep. 11, 1890, p. 357 (1891).
Leaf-spot Phomopsis vexans (Sacc. & Wint.), Hart. = Ascochyla hortorum (Speg.),
C. O. Smith, a fruit rot).
Journ. Agr. Research H, pp. 331-338, pis. 5 (1914).
Descr. Illus., N. J. Agr. Exp. Sta., Rep. 11, 189c, pp. 355-357 (1891).
Del. Bull. 70, pp. 10-15, pi. I, figs. 2 (March, 1905).
Treat, (pos.), N. J. Agr. Exp. Sta., Rep. 17, 1896, pp. 337-340 (1897).
Rot {Penicilliiim sp.).
Seedling-rot {Phomopsis vexans, Sacc. & Syd., Hart.).
Stem-rot {Neciria ipomcecE, Hals.).
Descr. Illus., N. J. Agr. Exp. Sta., Rep. 12, 1891, pp. 281-283 (1892),
Elder
{SambucHS canadensis, L.)
Rust {Aecidiiim samhiici, Sacc).
Leaf- spot {Ccrcospora catenas pora, Atk.).
Elm
(Ulmus spp.)
Black-spot {Dolhidella idmi (Duv.), Wint.) and {Gnomonia uhnea (Sacc), Thiim )
Blister-canker {Nmnmularia discreta, (Schw.) TuL).

Duggar, p. 282 (1909).
28
.
'
Leaf-scab (Gnomonia nlmea (Sacc), Thiim.).

White-rot {Polyporus squamosus (Huds.), Fr.)-
Duggar, p. 453 (1909).
English Ivy
{Hedera helix, L.)
Anthracnose {CoUetolrichum glceosporioides, Penz, var. hedera, Pass.

Leaf-blight {Phyllosticla concentrica, Sacc).
Leaf-spot {Ramidaria hedericola, Heald & Wolf).
Evening Primrose
{Oenothera biennis, L.).
Gall, or Chytridiose {Synchytriiim fulgens, Schrot.).

Duggar, p. 139 (1909).
.
Fig
{Ficiis carica, L.)
Anthracnose {Glomerella riifomaculans (Berk.) Spauld & von Schr. = G. Jrucligena

(Clint.), Sacc.)
Canker {Tubercularia fici, Edgerton).
Cook, Diseases of Tropical Plants, p. 139 (19 13).
Phytopath. i, pp. 12-17 (February, 191 1).
Die-back {Diplodia sycina, Mont., var. syconophila, Sacc).
Fruit-rot {Glomerella riifomaculans (Berk. Spauld. & von Schr.).
Leaf-blight {Cercospora Bolleana (Thiim.), Sacc).
Occ, U. S. Dep. Agr., Div. Pomol., Bull. 5, pp. 27-28 (1897)
Leaf-spot {Cercospora fici, H. & W.).
Limb-blight {Corticium latum, Karst.).

Rust {Uredofici, Cast. = Physopella fici (Cast.), Arth.).
Occ, N. C. Agr. Exp. Sta., Bull. 92, p. 117 (1893).
Scab {Fusarium roseum, Lk.)
Soft-rot {Rhizopus nigricans Ehrenb.).
La. Agr. Exp. Sta., Bull. 126 (March, 1911).
Filbert
{Corylus avellana, L. and C amerlcana, Walt.)
Black-knot {Cryptosporella anomala (Pk.), Sacc).

Descr., N. J. Agr.Exp. Sta., Rep. 13, 1892, pp. 287-289 (1893).
Fir
{Abies balsamea (L.), Miller)
Dry-rot {Trametes pini (Brot.), Fr.)

Descr. Illus., U. S. Dept. Agr., Div. Veg.
Root-rot {Polyporus Schweinitzii, Fr.)
Phys. & Path., Bull. 25 (1900).
Wet-rot {Polyporus subacidus, Pk.?)
Rust {Aecidium elatinum, Alb. & Schw.)
Flax
{Liniim spp.)
Rust {Melampsora lini (DC), Tul.).

Occ, Journ. Mycol., Vol. V, p. 215 (1889).
Wilt {Fusariiim lini, BoUey).
N. Dak. Bull. 50, December, 1901, pp. 27-60, figs. 18.
Geranium
{Pelargonium spp.)
Leaf-spot {Bacteria'^).
Descr., Mass. Agr. Exp. Sta., Rep. le, 1899, p. 57 (1900).
Kot {Bacillus i^.).
Descr. Illus., Journ. Mycol., Vol. VI, pp. 114-115 (1891).
Ginseng
{Panax quinquefolium, L.).^
Anthracnose {Vermicular ia dematium (Pers.), Fr.).

Blight {Alternaria panax, Whetz).
Leaf Anthracnose {Pestolozzia funerea, Desm.).
Wilt {Neocosmopara lasinfectum (Atk.) E. F. Sm. var nivea (Atk.) E. F. Sm.).
Mo. Bull. 69 (October, 1905).
Gladiolus
Hard-rot {Septoria gladioli, Passer).

Phytopathology 6 (Columbus Meeting Abstracts).
GOLDENROD
{Solidago spp.)
Red-rust {Coleosporium solidaginis (Schw.), Thum).

Rust Uromyces solidaginis (Somm.), Niessl.
1 See Whetzel, H. H.: The Diseases of Ginseng and Their Control, U. S. Bur.
of Plant Industry, Bull. 250 (191 2).
Gooseberry
{Ribes grossularia, L.)
Leaf-spot {Septoria ribis, Desm., and Cercospora angulata, Wint.)-

Leaf-spot {Sphmrella grossularia (Fr.), Awd.?).
Occ. Illus., Iowa x\gr. Exp. Sta., Bull. 13, p. 70 (1891).
Powdery Mildew {Spkarotheca mors-uvcR (Schw.), Bri. & Cav.).
Descr. U. S. Dep. Agr., Rep. for 1887, pp. 373-378 (1888).
Illus.,

Treat, (pos.), N. Y. Agr. Exp. Sta., Bull. 161 (1899).
Root-rot {Dcmatophora sp.?).
Descr., N. Y. Agr. Exp. Sta., Bull. 167, pp. 295-296 (1899).
Rust {Aecidium grossularicB, Schum.).
Grape
{Vilis spp.)
Anthracnose {S phaceloma ampdinitm, deBy. = GJocosporium ampdophagmn (Pass.)

Sacc).
Descr. Illus., Tenn. Agr. Exp. Sta., Bull. IV-4, pp. 111-112 (1891).
Descr., U. S. Dep. Agr., Div. Veg. Path., Bull. 2, pp. 170-172 (1892).
Shear, C. L. Grape Anthracnose in America. Rep. Int. Congr. Viticulture,
San Francisco, July 11-13, 191 5: 111-117.
Hawkins, LonA. Circ. 105, Bureau PI. Industry, 1913.
Bacteriosis {Bacillus sp.).
See U. S. Dep. Agr., Exp. Sta. Rec, VI-3, pp. 231-232.
Bitter-rot{Melanconium fuligincmn (Scrib. & Viala.), Cav.).

Scribner, Fung. Dis., pp. 37-40 (1890).
Cf. N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 61, pp. 302-305.
Black-rot {Guignardia {Lacsiadia) BidweUii (Ell.), Viola. & Rav. and G. ba'cca (Cav.),
Jacq.).
Del. Agr. Exp. Sta., Bull. 6, pp. 18-27 (1889).
Tenn. Agr. Exp. Sta., Bull. IV-4, pp. 97-102 (1891).
Tex. Agr. Exp. Sta., Bull. 23, pp. 219-228 (1892).
Penna. Bull. 66, pp. 1-16, pis. 2, map. i (Jan., 1904).
N. Y. Cornell Bull. 293, pp. 289-364, pis. 5 (March, 191 1).
Treat, (pos.), Conn. Agr. Exp. Sta., Rep. 14, 1890, pp. loo-ioi (1891).
U. S. Dep. Agr., Farm. Bull. 4, pp. 8-9 (1891).
Tex. Agr. Exp. Sta., Bull. 23, pp. 228-231 (1892).
Chytridiose {Cladochylrium vilicolum, Prunet.).

See U. S. Dep. Agr., Exp. Sta. Rec, VI-7, pp. 642-644 (1895).
Dead-arm {Cryplosporella viticola, Shear.).
Circ. 55, N. J. Agr. Exp. Sta.
N. Y. State Bull. 389, pp. 463-490 (Julj', 1914).
Phytopath. i, pp. 116-119 (1911).
Uowny Mildew (Plasmopara viticola (B. & C), Berl. & De Ton.).
Descr. IIlus., U. S. Dep. Agr., Rep. for 1886, pp. 96-99 (1887).
Tenn. Agr. E.\p. Sta., Bull. IV-4, p. 108 (1891).
Mich. Agr. E.xp. Sta., Bull 83, pp. 9-12 (1892).
Phytopath. 2, pp. 235-249 (1912).
Treat, (pos.), U. S. Dep. Agr., Farm. Bull. 4, p. 8 (1891).
Fruit- mold (Bolrytis sp.).
Leaf-blight Isariopsis claviipora (B. & C.) Socc.
Descr. Illus., Scribner, Fung. Dis., pp. 60-62 (1890).
N. Y. Agr. Exp. Sta., Rep. 9, 1890, p. 324 (1891).
Cornell Bull. 76, November, 1894.
Treat, (rec), N. C. Agr. Exp. Sta., Bull. 92, p. 122 (1893).
Leaf-mold {Leptosporium hderosporum, Ell. & Gall.).
Leaf-spot {Isariopsis davispora, Sacc).
N. J. Exp. Sta., Rep., p. 474 (1914).
Powdery Mildew {Uncimda necator (Schw.), Burr.).
Descr. Illus., U. Dep. Agr., Rep. for 1886, pp. 105-108 (1887).
S.
N. Y. Agr. Exp. Sta., Rep. 9, 1890, pp. 322-323 (1891).

U. S. Dep. Agr., Div. Veg. Path., Bull. 2, pp. 166-170 (1892).
Treat, (pos.), U. S. Dep. Agr., Farm. BulL 4, p. 8 (1891).
N. C. Agr. E.xp. Sta., Bull. 92, pp. 120-121 (1893).

Ripe- rot or Anthracnose {Glceosporium fructigemim, Berk.).
Descr. Illus., U. S. Dep. Agr., Rep. for 1890. p. 408 (1891).
Journ. Mycol., Vol. VI, pp. 164-171 (1891).
Root- rot {Demalophora necatrix, Hartig).
Descr. Illus., Scribner, Fung. Dis., pp. 64-69 (1890).
S. Dep. Agr., Div. Veg. Path., Bull. 2, pp. 153-159 (1892),
U.
Treat. N. C. Agr. Exp. Sta., Bull. 92, p. 122 (1893).
Root-rot {Annillaria mcllca, Vahl.).
Scab (Cladosporium vilicolum, Ces. = Cercospora viticola (Ces.) Sacc.)
Descr., U. S. Dep. Agr., Div. Veg. Path., Bull. 2, pp. 173-174 (1892).
Scald {Aureobasidium litis, Viala & Boyer).
See U. S. Dep. Agr., E.xp. Sta. Rec, VI-3, pp. 230-231 (1894).
Twig-blight {Bolrytis cinerea, Pers.).
White-rot {Charrinia diphdiella, Viala & Rav.; Syn. Coniolhyriiim diplodidla
(Speg.) Sacc).
Scribner, Fung. Dis., pp. 41-44 (1890).
Treat, (pos.), U. S. Dep. Agr., Sec. Veg. Path., Bull. 11, p. 69 (1890).
. . .
GUAVA
{Psidium guajava, L.)
Ripe-rot {Glonierclla psidii (G. Del.) Sheldon).

W. Va. Bull. 104, pp. 299-315 (April, 1906).
Hackberry
{Cellis spp.)
Leaf-spot {Cylindros poriuni defoliaium, Heald and Wolf and {Ramularia ccUidis,
EU. & Kell.).
Powdery Mildew {Uncinula polychccta, Bri. and Cav.).
Hazel
{Corylus spp.)
Black-knot {Cryplosporella anomala (Pk.), Sacc).

Descr. lUus., Mass. Agr. Exp. Sta., Rep. 10, 1892, p. 242 (1893).
Hemlock
{Tsuga canadensis (L.), Carr.)
Dry-rot {Trametcs pini (Brot.), Fr.).

Heart-rot {Polyporus borealis (Wahl.), Fr.).
Bull. 193 Corn. Univ. Agr. Exp. Sta. (June, 1901).
Timber Rot {Fames pinicola, Fr.)
Graves, A. H., Phytopath. 4, p. 69 (April, 1914).
Wet-rot {Polyporus siihacidus, Pk. ?)
Rust {Peridermium Peckii, Thiim.)
Phytopath. i, pp. 94-96 (191 1).
Hemp
{Cannabis saliva, L.)
Leaf- wilt {Bolryosphceria Marconii (Cav.), Charles & Jenkins).

Journ. of Agr. Research 3, pp. 81-84 (Oct. 15, 1914).
Hickory
{Carya spp.)
Leaf-spot {Marsonia juglandis (Lib.), Sacc).

Hollyhock
{AlthcEa rosea, Cav.)
Anthracnose {Colletolrichum malvarum (Braun. & Casp.), Southworth).

Descr. Journ. Mycol., Vol. VI, pp. 46-48 (1890).
Illus.,
Treat, (pos.), Journ. Mycol., Vol. VI, p. 50 (1890).

N. J. Agr. Exp. Sta., Rep. 11, 1890, p. 362 (1891).
Leaf-blight [Cercospora althaina, Sacc).
Descr., N. j. Agr. Exp. Sta., Rep. 11, 1890, p. 361 (1891).
Treat, (pos.), N. J. Agr. Exp. Sta., Rep. 11, 1890, p. 361 (1891).
Leaf-Spot {Phyllosticta althmina, Sacc.).^
Rust {Puccinia malvacearum, Mont.).
Descr. Illus., N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 25, p. 154 (1890).
Phytopath. i, pp. 53-62 (191 1).
Rust {Puccinia helerogenea, Lagerh.)
Descr. Illus., Journ. Mycol., Vol. VII, pp. 44-47 (1891).
Hop
{Humuliis japonicus, Sieb & Zucc.)
Powdery Mildew {Spharotheca hiimtdi (DC), Burr.).

N. Y. Corn. Bull. 328, pp. 281-310, figs. 19 (March, 1913).
N. Y. State Bull. 395, pp. 29-80, pis. 2, figs. 2 (February, 1915).
Horse-chestnut
{jEschIus hippocastaniim, L.)
Leaf-blotch {Gitignardia csscidi (Pk.) Stewart). Phytopath. 6, 5-19, 1916.

Leaf-spot {Phyllosticta pavice, Desm.).
Descr., N. Y. Agr. Exp. Sta., Rep. 15, 1896, p. 456 (1897).
Tr. (pos.), Journ. Mycol. Vol. VII, p. 353; Phytopathology 4, 399 (December,
1914)
Horseradish
{Cochlearia armoracia, L.)
Leaf-blight {Ramularia armoracia, Fckl.).

Occ, N. J. Agr. Exp. Sta., Rep. 11, 1890, p. 360 (1891).
Leaf- mold {M acres porium herculeum. Ell. & Mart.).
Occ, N. Y. Agr. Exp. Sta., Rep. 15, 1896, p. 452 (1897).
' The be found described in The North Ameri-
dififerent species of Phyllosticta will
can Phyllostictas with Descriptions of the Species, published up to August, 1900 by

J. B. Ellis and B. M. Everhart, Vineland, N. J., December, 1900.
Leaf-spot {Scploria armoracice, Sacc.)-

Descr., N. J. Agr. Exp. Sta., Rep. ii, 1890, p. 360 (1891).
Huckleberry
(Gaylussacia sp.)
Gall {Exohasidiitm vaccinii (Fckl.) Wor.j.
Hyacinth
{Hyacintlius oricnlalis, L.)
Yellow Disease {Pscudomonas hyacinihi (Wakk.) E. F. Sm.).
Hydrangea
{Hydrangea horlensia, Siebold)
Leaf-spot {Pliyllostkta hydrangea, Ell. & Ev.).

Rust {Melampsora Hydrangea = Thecopsora hydrangea B. & C.) Magn.
Incense Cedar
{Libocedrus decurrens, Torr.)
Dry-rot {Poly poms amarus, Hedgcock).

Rust {Gymnosporanglum Blasdaleanum (Diet. & Holw.) Kern).
Meinecke, E. A., Forest Tree Diseases Common in California and Nevada, 1914.
Iris
{Iris spp.)
Bulb-spot {Mystrosporium adustum, Mass.).

Leaf-blight {Botrytis galanthina, (B. & Br.) Sacc).
Johnson Grass
{Andropogon halepensis (L.), Brot.).
Leaf-blight {Helminthosporiuni turcicum Pass, and Septoria pcrlitsa Heald & Wolf).
Leaf-spot {Cercospora sorghi (Ell. & Ev.) and Colletolrichiim lineola Cda var. hale-
pense, Heald & Wolf).

Rust {Puccinia purpurea, Cke.).
Kaffir Corn
{Sorghum vulgare, Pers.)
Grain Smut {Sphacetolheca sorghi Lk.) Clint.

Clint Conn., Exp. Sta. Rep., p. 351 (1912).
Larch
(Larix lan'cina (DR.) Koch)
Canker (Dasyscypha Willkommii, Hartig).

Dry-rot (Tramdes pint (Brot.) Fr.).
(1900).
Laurel
{Kalmla laiijolia, L.)
Leaf-spot {Seploria kalniicola (Schw.) Bri. & Cav.).
Lemon
{Citrus mcdica, L. var. Union, L.)
Black pit {Bacillus citripuleale spp.)

Coit, Citrus Fruits, p. 401, 1915; Phytopath. 3, pp. 277-281 (1913).
Brown-rot {Pythiacystis dtrophthora, R. E. Smith).
Calif. Bull. 190, pp. 1-72, pi. I, figs. 30 (July, 1907).
Foot-rot {Fusisporium limonis, Bri.).
Cotton-rot {Sclerotinia libertiana, Fuckl.j. Phytopath. 6, pp. 268-278 (1916).
Fruit-spot {Trichoseploria alpci, Cav.).
Leaf-spot {Ccrcospora aurantia.^ Heald & Wolf).
Melanose {Fungus indel?).
Canker {Pscudomonas citri, Hassej. Journ. Agr., Res. 4: 97-150 (1915).
Scab {Cladosporium, sp.).
Descr. Illus., U. S. Dept. Agr., Div. Veg. Phys. & Path., Bull. 8, pp. 20-23
(1896).
Treat, (pos.), U. S. Dept. Agr., Div. Veg. Phys. & Path., Bull. 8, pp. 23-24
(1896).
Sooty- mold {Mcliola Penzigi, Sacc. and M. Camdlice (Catt.), Sacc.j.
Twig- blight {Diplodia aurantii, Catt and Sphceropsis maloruin, Berk.).
White-rot {Sclerotinia libertiana Fckl.) Bull. 218, Calif. Agr. Exp. Sta. (June, 191 1).
Wither-tip {Colletotrichum glceosporioidcs Penz.
Plant Disease, Survey San Antonio Texas (191 2).
Lettuce
{Lactuca sativa, L.)
Anthracnose {Marssonia perforans, Ell & Ev-.).

Descr. Illus., Ohio Agr. Exp. Sta., Bull. 73, pp. 222-223 (1897).
Treat, (rec), Ohio Agr. Exp. Sta., Bull. 73, pp. 225-226 (1897J.
.
Downy Mildew (Breniia lacluca, Regel).

Descr. N. Y. Agr. Exp. Sta., Rep. 4, 1885, p. 253 (1886).
Illus.,
Treat, (pos.), Ohio Agr. Exp. Sta., Bull. 73, p. 226 (1897).
Drop {Schrotinia libertiana, Fckl.).
Descr. Illus., Mass. Agr. Exp. Sta., Bull. 69, pp. 12-15 (1900).
N. C. Bull. 217, 1-21, figs. 8 (July, 1911).
Treat, (pos.), Mass. Agr. Exp. Sta., Bull. 69, pp. 17-35 (19°°)
Leaf-mold, Gray Mold or Rot {Botrytis cinerea, Pers.;.
Descr. Illus., Mass. Agr. Exp. Sta., Bull. 69, pp. 7-12 (1900).
Leaf-rot {Rhizoclonia sp.).
Descr. Illus., Mass. Agr. Exp. Sta., Bull. 69, pp. 16-17 (1900)
Treat, (pos.j, Mass. Agr. Exp. Sta., Bull. 69, pp. 39-40 (1900).
Leaf-spot {Septoria conslmilis, Ell. & Mart.).
Descr. Illus., Ohio Agr. Exp. Sta., Bull. 44, pp. 145-146 (1892)
Stem-rot (Bacterial).
Descr., Vt. Agr. Exp. Sta., Rep. 6, 1892, p. 87 (1893).
Treat, (rec.) Vt. Agr. Exp. Sta., Rep. 6, 1892, p. 88 (1893).
Lilac
{Syringa vulgaris, L.)
Leaf-spot {Phyllosticta Halstedii, Ell. & Ev.).

Powdery Mildew {Microsphcera aim (Wallr.) Wint.).
Leaf-blight {Cercospora macromaculans. Heald & Wolf).
Lily
(Liliiim spp.)
Bermuda Disease.
See U. S. Dept. Agr., Div. Veg. Phys. & Path., Bull. 14 (1897)
Bulb-rot {Rhizopus necans, Massee).
Mold or Ward's Disease {Sclerotinia Fuckcliana deBy.).
Treat, (pos.). Gar. and For., IX-414, p. 44 (1896).
See N. J. Agr. Exp. Sta., Rep. 14, 1893, pp. 392-394 (1894)
Linden
{Tilia spp.)
Leaf-blight {Cercospora microsora, Sacc).

Occ, N. Y. Agr. Exp. Sta., Rep. 15, 1896, p. 454 (1897).
Stem-rot {Botrytis cinerea, Pers.).
Locust
{Rohinia pseiidacacia, L.)
Leaf-spot {Cylindrosporium solitarinm, Heald & Wolf).

Heart-rot {Trametes robiniophila, Murr. and Fames rimosus Berk.)
Diseases of Deciduous Trees (1909).
LIST OF SPECIFIC DISEASES OF PLANTS 44^3,
LOQUAT
(Eriobotrya japonica, Lindl.)
Scab {Fitsicladium dendrUicum (Wallr.), Fckl. var. EriobolrycB, Scalia.
Lupine
{Lupinus, spp.)
Blight (Fcstalozzia litpini, Sor.).
Magnolia
{Magnolia grandijlora, L.)
Leaf-spot {PhyUoslicla magnolia Sacc. Duggar, p. 347 (1909).
Mango
(Mangifcra indica, L.)
Anthracnose (Collelelolrichnm glceosponoides, Penz.).

McMurran Bull. U. S. Dept. Agr. No. 52 (1914).
Maple
(Acer spp.)
Anthracnose {Glaosporium apocrypium, Ell. & Ev.).

Descr., N. Y. Agr. .Exp. Sta., Rep. 14, 1895, pp. 531-532. (1896).
Treat, (rec), N. Y. Agr. Exp. Sta., Rep. 14, 1895, p. 532 (1896)
Decay, Fomes fomentarins (L.) Fr. Duggar, p. 467.
Gall, Pycnochylrium globosum, Schrot; Duggar, p. 139.
Heart-rot, Fomes igniarius (L.) Gill.; Duggar, p. 465.
Leaf-blotch, Rhytisma acerinum (Pers.) Fr.
Leaf-spot {Phyllosticta acericola, Cke. & Ell.).
Descr. Illus., Dep. Agr., Rep. for 1888, pp. 383-386 (1889).
U. S.
Powdery Mildew, Uncintila aceris (DC.) Wint.
White-rot, Polyporus squamosus (Huds.) Fr.; Duggar, p. 453.
Melon
(Cucumis melo, L.)
Anthracnose (Collclotrichnm lagenarium (Pass.) Ell. & Hals.).

Descr., U. S. Dep. Agr., Bot. Div., Bull, 8, p. 64 (1889).
Descr. Illus., Okla. Agr. Exp. Sta., Bull. 15, pp. 30-31 (1895).
Treat, (pos.), Md- Agr. Exp. Sta., Rep. 4, 1891, p. 387 (1892).
.
Anthracnosc {Collclolrichum oligochcetum, Cav.).

Bacteriosis or Wilt {Bacillus tracheiphilus, E. F. Sm.).
Downy Mildew {Plasmopara cubenis (B. & C.) Humph.).
Occ. Descr., Conn. Agr. Exp. Sta., Rep. 23, 1899, pp. 277-278 (1900).
Leaf-blight {Altcrnarla hrassica, Sacc, var. nigrescens, Regel.).
lUus., Ohio Agr. Exp. Sta., Bull. 73, pp. 235-236 (1897).
Treat, (pos.), Conn. Agr. Exp. Sta., Rep. 22, '9Sj#%). 229-235 (1899)-.
Cf. Conn. Agr. Exp. Sta., Rep. 23, 1899, pp. 270-573 (1900).
Leaf-spot {Phyllosticta cuctirbitacearum, Sacc. ?)
Scab {Scolecotrichutn melophlhorum, Pr. & Del.).
Soft-rot (Bacillus melonis, Gidd.) Vt. Bull. 148, 363-416, pis. 8 (Jan. 1910).
Southern Blight (Sclerotium Rolfsii, Sacc).
Wilt (Neocosmospora vasinfccta (Atk.) E. F. Sm.).
Mesquite
{Prosopis juliflora, DC.)
Anthracnose {Glaosporium leguminum (Cke.), Sacc).

Blight {Scleropycnium aureutn, Heald & Lewis). Trans. Amer. Micr. Soc, XXX F,
5-9 (June, 19 1 2).

Rust (Ravenelia arizonica, Ell. & Ev.).
Mignonette
{Reseda odorata, L.)
Leaf-blight {Cercospora reseda;, Fckl.).

Treat, (pos.), U. S. Dep. Agr., Rep. for 1889, p. 431 (1890).
Millet
{Panicum miliaceum, L.)
Purple-spot {Piricularia grisea (Cke.), Sacc).

Smut {Uslilago Cramcri, Korn.).
Mulberry
{Morus spp.)
Die-back {Myxos pari urn Dledickii, Syd.).

Chytridiose {Cladochytrium mori, Prunet.).
See U. S. Dept. Agr., Exp. Sta. Rec, VI-9, p. 830 (1895).
Eye-spot {Ccrcospora moricola, Cke.).

Leaf-spot {Ccrcospora missouriensis, Wint).
Rooi-iot (Hclicobasidium mompa,Ta.naka..=Scptobasid!nm mompa (Tanaka), Rac.«).
Mushroom
(Agariciis campcslris, L.)
Mold {Mycogone perniciosa, Magn.).
Nasturtium
{TropceoJtim majiis, L.)
Wilt {Pscudomonas solanaccarum, E. F. Sm.).

Journ. Agric. Research 4, pp. 451-457, pis. 64 (1915).
Leaf-blight {Alternaria sp., and Pleospora tropceoli, Hals.).
Descr., N. J. Agr. Exp. Sta. Rep. 13, 1892, p. 290-293 (1893).
Oak
(Quercus spp.)^
Anthracnose (Gnomonia veneta (Sacc. & Speg.), Kleb).

Pocketed-rot {Poly poms pilotce, Schw.).
Decay, or Brown-rot {Polyporus sulphureus (Bull.) Fr.).
Atkinson, Bull. 193, Cornell Agr. Exp. Sta. (June, 1901).
Heart- rot {Fames igniarius (L.) Gill.).
Honeycomb Heart- rot {Stcreum subpileatiim, B. & C).
Journ, of Agr. Research V; 421 (Dec. 6, 1915).
Leaf-curl {Taphrina caerulescens, Desv. & Mont.), Tul.
Leaf-spot {Marsonia quercus, Pk.}.
f {Armillaria mellea, Vahl). Bull. U. S. Dep. Agr., No. 89 (1914).
{Clitocyhe parasUica, Wilcox).
Root-rot
{Polyporus dryadetis, Fr.).
1^
{Rosellinia quercina, Hartig).
Soft Rot {Polyporus obhisus, Berk).
String and Ray-rot {Polyporus Berkeleyi, Fr.).
Straw-colored Rot {Polyporus frondosus, Fr.), Journ. Agr. Research I, 109 (1913).
Tar-spot {Rhyiisma erylhrosporum, Bri. & Cav.).
White-rot {Polyporus squamosus (Huds.) Fr.).
1 Consult VON SCHRENK, HERMANN and Spaulding, Perley: Diseases of De-

ciduous Forest Trees. Bull. 149, U. S. Bureau of Plant Industry, 1909.
Oats
(Aiena saliva, L.)
Blight (Bacterial) Pseudomonas avence, Manns.

Descr., Journ. Mycol., Vol. VI, p. 72; Ohio Bull. 210, Oct. 1909, pp. 91-167,
pis. IS (1890).
Leaf-spot {Phyllosticla sp.).
Descr., N.
Agr. Exp. Sta., Rep. 15, 1894, p. 319 (1895).
J.
Mildew {Helminthosporium inconspicuum, Cke. & Ell., var. hrillanicum Gr., and
Cladosporiiim herbarum (Pers.), Lk.).
Me. Agr. Exp. Sta., Rep. for 1894, pp. 95-96 (1895).
Descr.,
Rust {Puccinia coronata, Cda., and P. graminis, Pers.).
See Wheat (Rust).
Cf. U. S. Dep. Agr., Div. Veg. Phys. & Path., Bull. 16, pp. 45-52 & 60-65
(1899).
Smut (Uslilago avena (Pers.), Jens, and U. levis (Kell. & Sw.) Magn.).
Descr. lUus., Kan. Agr. Exp. Sta., Rep. 2, & 259-260 (1890).
1889, pp. 215-238
Treat (pos.), U. S. Dep. Agr., Farm. Bull. 75, pp. 11-16 (1898).
Okra
(Hibiscus esculentus, L.)
Root- rot (Ozonium omnivorum, Shear).

Wilt {Fusarium vasinfedum = Neocosmos pora vasinfcclum (Atk.), E. F. Sm.).
See U. Dep. Agr., Div. Veg. Phys." & Path., Bull. 17, p. 31 (1899).
S.
Cf. N. Car. Rep. 191 1, pp. 70-73, figs. 4.

{Verticillium albo-alrmn, Reinke & Berthold) Phytopathology IV, p. 393 (De-
cember, 1914).
Oleander
{Neritim oleander, L.)
Leaf-spot (Macrosporium nerii, Cke.).

Bull. 218, CaHf. Agric. Exper. Sta. (June, 191 1).
Olive
(Oka europcea, L.)
Anthracnose {Glosospormm olivarum, d'Almeida).

Fruit-mold or Dry-rot (Alternaria sp. and Macrospoxium sp.).
Descr. lUus. Cal. Agr. Exp. Sta., Rep. for '95-'97, pp. 235-236 (1898).
Knot (Pseudomonas Savastonoi, E. F. Sm.).

Cook Diseases of Tropical Plants, p. 144 (1913).
Rot (Bacterial).
Descr., Cal. Agr. Exp. Sta., Bull. 123, p. 19 (1899).
Scab, Peacock Leaf-spot {Cycloconium oleaginum, Cast.).
Descr., Cal. Agr. Exp. Sta., Rep. 1892-93, pp. 297-298 (1894).
See U. S. Dep. Agr., Exp. Sta. Rec, XI-6, p. 554 (1900).
Sooty- mold (Meliola sp., Syn. Capnodlunt cilri Berk. & Desm.).
Tuberculosis {Bacillus clece (Arcang.) (Trev.).

Descr. Illus., Cal. Agr. Exp. Sta., Bull. 120 (1898).
Treat, (rec), Cal. Agr. Exp. Sta., Bull. 120, pp. lo-ii (1898).
Cf. Cal. Agr. Exp. Sta., Rep. for '97-'98, p. 178 (1900).
Onion
{Allium cepa, L.)
Anthracnose or Rot {Vermicular ia circinans, Berk).

Descr. Illus., Conn. Agr. Exp. Sta., Rep. 13, 1889, p. 163 (1890).
Treat, (rec), Conn. Agr. Exp. Sta., Rep. 13, 1889, pp. 164-165 (1890).
Downy Mildew {Peronospora Schleideniana, deBy.).
Descr. Illus., Wis. Agr. Exp. Sta., Rep. i, 1883, pp. 38-44 (1884).
N. Y. Cornell Bull. 218, pp. 137-161, figs. 17 (Apr., 1904).
Treat, (rec), Vt. Agr. Exp. Sta., Rep. 10, 1896-97, pp. 61-62 (1897).
Mold {Macrosporium sarcinula, B., var. parasiticum, Thiim., and M. Porri, Ell.).
Descr. Conn. Agr. Exp.
Illus., Sta., Rep. 13, 1889, pp. 158-162 (1890).
Treat, (rec), Conn. Agr. Exp. Sta., Rep. 13, 1889, p. i6r (1890).
Rot (Bacterial).
Descr. Illus., N. Y. Agr. E.xp. Sta., Bull. 164, pp. 209-212 (1899).
Smut {Urocyslis cepula, Frost).
Descr. Illus., Conn. Agr. Exp. Sta., Rep. 13, 1889, pp. 129-146 (1890).
Ohio Bull. 122, pp. 71-84, figs. 4 (Dec, 1900).

Treat, (pos.), Conn. Agr. Exp. Sta., Rep. 13, 1889, pp. 147-153 (1890).
(By transplanting), Conn. Agr. Exp. Sta., Rep. 19, 1895, pp. 176-182
(1896).
Cf.U. S. Dep. Agr., Farm. Bull. 39, pp. 16-20 (1896).
N. Y. State Bull. 182, pp. 145-172, pi. i (Dec, 1900).
Orange
{Citrus aurantium; L.)
Anthracnose {Collelotrichum gloeosporioides Penz.).

Descr. Illus., Fla. Agr. Exp. Sta., Bull. 53, pp. 171-173 (1900). .
Fla, Agr. E.xp. Sta., Bull. 108, pp. 25-47 (Nov., 191 1).
Treat, (rec), Fla. Agr. Exp. Sta., Bull. 53, p. 173 (1900).
Black-rot {AUcrnaria citri Ellis & Pierce).
Colt, Citrus Fruits, p. 388 (1Q15).
Cottony- mold and Twig-blight {Sdcrotinia libcrliana, Fkl.).
Coit, Citrus Fruits, p. 382 (1Q15).
Diplodia Rot {Diptodia natalensis Evans), Coit, p. 397 (1915).
Flyspeck {Leptothyrium pomi (Mort & Fr.) Sacc.) Hume, Citrus Fruits and Their
Culture, p. 481 (191 1 ).
Foot-rot or Mal-di-gomma {Fusarium limonis, Bri.).

Descr. Illus. Treat, (rec), U. S. Dept. Agr., Div. Veg. Phys. & Path., Bull. 8,
pp. 28-31 (1896J.
Fla. Agr. Exp. Sta., Bull. 53, pp. 151-155 (1900).
Fruit-rot {Penicilliiim digitatum. (Fr.), Sacc. & Penicillium Ualicum, Wehm.).
Gum-disease {Bolrytis vulgaris, Fr.).
Coit, Citrus Fruits, p. 366 (1915).
Leaf-glaze {Strigula complanata, Fee).
Occ, Journ. Mycol., Vol. VII, p. 36 (1891).
Melanose {Phomopsis citri Fawcett).
Descr. Illus. Treat, (pos.), U S. Dept. Agr., Div. Veg. Phys. & Path., Bull. 8,
PP- 33-38 (1896).

Nail-head Rust {Cladosporium hcrharum (Pers.), Pk. var. citricolum Fawcett &
Berger).
Coit, Citrus Fruits, p. 395, 1915, Fla. Bull. 109, pp. 47-60 (May, 191 2).
Scab {Cladosporium citri Mass.).
Phytopath. 6, pp. 127-142 (1916).
See Lemon (scab.).
Sooty-mold {Meliola Penzigi, Sacc. and M. camellice (Catt.), Sacc.
Descr. Illus. Treat, (pos.), U. S. Dept. Agr., Div. Veg. Phys. & Path., Bull. 13
(1897).
Toadstool Root-rot (Armillaria mcllea Vahl.).
Coit, Citrus Fruits, p. 373 (1915).
Trunk-rot {Schizophyllum commune Fr.).
Coit, Citrus Fruits, p. 399 (191 5).
Wither-tip (Colletotrichum gloeosporioides, Penz.).
Coit, Citrus Fruits; 380 (1915).
Grossenbacher, J. G.; Some Bark Diseases of Citrus Trees in Florida, Phyto-
path. 6, pp. 29-50 (1916).
Orch.xrd Grass
{Dactylis glomcrala, L.)
Puccinia coronata, Cda.;Duggar, p. 420 (1909)

Puccinia graminis, Pers.; Duggar, p. 408 (1909^
Scolecotrichose {ScoJetotrichum graminis Fuckl.).
ORCHros
(Orckidacca)
Anthracnose {Glceosporium cincliim Bri. & Cav. Colletotrichum cinclum (Bri. & Cav.)
Stonem.).
Anthracnose {Glceosporium macro pus, Sacc).
Leaf-blight {Ccrcospora angrcci, Feuill & Roum.).
Osage Orange
(Toyxlon pomiferum, Raf.)
Rust {Physopella ficl (Cast), \rt\\. = U redo fici Cast.)

Blight {Sporodesmium maclurcr Thiim.
Palm
(Phoenix dactylifera, L.)
Leaf-spot (Graphiola plicenicis (Moug.) Poit.).

Bull. 218, Calf. Agr. E.xp. Sta. (June, 191 1).
Pansy
{Viola tricolor, L.)
Leopard Petal-blight {Colletotrichum, viola-iricoloris), R. E. Smith.

Smith, R. E., Bot. Gaz. 27, p. 203 (March, 1899).
Dry-up {Fusarium djo/(5 Wolf.).
Wolf, F. A.; Mycologia, 2, p. 19 (January, igio).
Papaw
{Carica papaya, L.)
Leaf-spot {Pucciniopis caricce Earle).
Parsnip
{Paslinaca saliva, L.)
Leaf-blight {Cercospora apU, Fres.j.

Occ, N. J. Agr. E.Kp. Sta., Rep. 15, 1894, p. 351 (1895).
Root-rot {Corticium vagum, Bri. & Cav., var. solani, Burt.).
Heald & Wolf, Plant Disease Survey in Texas (191 2).
29
4SO SPECIAL PLANT PATHOLOGY
Pea
[Pisum salivuni, L.)
Damping-off {Ascochyla pisi, Lib. and Pylhium sp.).

Occ, Conn. Agr. Exp. Sta., Rep. 23, 1899, pp. 280-281 (1900).
Ohio Bull. 173, pp. 231-246, figs. II (Apr., 1906).
Pod-spot {Ascochyta pisi, Lib.).
Treat. (rec.J, Del. Agr. Exp. Sta., Bull. 41, pp. 9-1 1 (1898).
Leaf-spot {Septoria pisi, West.).
-
Mold {Pleospora pisi (Sow.), Fckl.).
Peach
{Primus persica, Benth. & Hook)
Anthracnose {Glceosporium laeticolor. Berk.).

Occ. Ohio Agr. Exp. Sta., Bull. 92, p. 225 (1898).
Brown-rot {Sclerotinia cinerea (Bon.) Schrot.) Heald, F. W., Washington Agricul-
turist, VIII, No. 9, June, 191 5.
Crown-gall (Pseiidomonas tumejaciens, E. F. Sm. and Towns.).
Die-back {Valsa leucostoma (Pers.) Fr.).
Stevens & Economic Plants, p. 129
Hall, Diseases of (1910).
California Peach Blight {Coryneum Beijerinckii Oud.).
Oregon Stat. Biennial Report, p. 255 (1911-12).
Cal. Bull. 191, pp. 73-98, figs. 17 (Sept., 1907).
Frosty Mildew {Cercosporella persica, Sacc).
Fruit-mold or Twig-blight {Sclerotinia friicligena (Pers.) Schrot.).
Descr. Illus., Journ. Mycol., Vol. VII, pp. 36-38 (1891).
Ga. Agr. Exp. Sta., Bull. 50 (1900).
Treat, (pos.), Ga. Agr. Exp. Sta., Bull. 50, pp. 267-269 (1900).
Cf. Cherry (Fruit-mold and Twig-blight).
Pustular-spot {Helminthosporitim carpophilum, Lev.).
Occ, Mich. Agr. Exp. Sta., Bull. 103, p. 57 (1894).
Treat, (pos.), Ohio Agr. Exp. Sta., Bull. 92, p. 225 (1898).
Leaf-blight or Shot-hole {Cercosporella persica, Sacc).
Leaf-blight or Frosty Mildew {Cercosporella persica, Sacc).
Leaf-curl {Exoascus deformans (Berk.), Fckl.).

Descr. N. Y. (Corn, Univ.) Agr. Exp. Sta., Bull. 73, pp. 324-325 (1894)-
Illus.,

Treat, (pos.), N. Y. Cornell Bull. 276, p. 151-178, figs/s (Apr., 1910).
Powdery Mildew {Spharolhcca pannosa (Wallr.), Lev. ? and Podospkcsra oxyacantha
(DC), de By.).
Descr.Illus., N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 74, p. 381 (1894).
Root-rot {Fungus indet.?).
Ohio Agr. Exp. Sta., Bull. 92, p. 23s (1898).
Rust {Puccinia pruni-persicce Hori).
Phytopath. 2, p. 143-145, also Tranzschelia punctata (Pers.) Arth.
2d Biennial Crop Pest and Hort. Rep. Oregon (June, 1915).
See Cherry (Rust).
Scab {Cladosporium carpophilum, Thiim).
Descr. Illus., Ind. Agr. Exp. Sta., Bull. 19, pp. 5-8 (1889).
Del. Agr. Exp. Sta., Rep. 8, 1895-96, pp. 60-63 (1896).
Treat, (pos.), Del. Agr. Exp. Sta., Rep. 8, 1895-96, p. 63 (1896).
Cf. N. J. Agr. Exp. Sta., Rep. 15, 1894, pp. 328-330. (On leaves).
Conn. Agr. Exp. Sta., Rep. 20, 1896, pp. 269-271. (On twigs).
Bull. 395, U. S. Dept. Agric, 1917.
Shot-hole {Coryneum Beijcrinckii Oud.).
Stevens & Hall, Disease of Economic Plants, p. 129 (1910).
Stem-blight {Phoma persicce, Sacc).
Descr. Illus., Ohio Agr. Exp. Sta., Bull. 92, pp, 233-234 (1898).
Yellows.
Peanut
{Arachls hypogcea, L.)
Bacterial Blight {Bacillus solanacearum, E. F. Sm.).

Phytopathology IV; 397 (December, 19 14).
Leaf-spot' {Cerccspora personata (Bri. & Cav.), Ell. & Ev.).
Red-rot {Neocosmopora vasinfecta (Atk.) E. F. Sm.).
Phytopathology IV; 397 (December, 1914).
Sclerotial-rot {Sclerotium Rolfsii Sacc).
' Consult also Wolf, Frederick A. : Further Studies on Peanut Leaf-spot.
Journ. Agr. Res. 5, pp. 891-902, Feb., 1916.
{Pirns conimunis, L.)
Anthracnose {Collclotrichum sp.).

Bitter- rot (Glomcrella rufomaculans (Berk.), Spauld. & v. Schr.).
Stevens & Hall, Diseases of Economic Plants, p. 107 (1910)
RIack-rot {Spharopsis malormn, Berk.).
Brown-blotch (Macros porium Sydowianum, Farneti).
Circ. 52, N. J. Agr. E.xp. Sta.
Body-blight or Canker (Spkacropsis malar urn, Berk.).
Occ, N. Y. Agr. E.xp. Sta., Bull. 163, p. 203 (1899). .
Dry-rot {Thelephora pediceUata, Schw.).

Descr., Journ. Mycol., Vol. VI, pp. 113-114 (1891).
Treat, (pos.), Journ. Mycol., Vol. VI, p. 114 (1891).
Fire-blight {Bacillus amylovorus (Burr.), Trev.).
Descr. Illus., N. Y. Agr. Exp. Sta., Rep. 5, 1886, pp. 275-289 (1887).
U. S. Dept. Agr., Year-book for 1895, pp. 295-298 (1896).
N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 145, pp. 622-625, 1898.
Utah Bull. 85, Nov., 1903, pp. 45-52.
Vt. Rep. 1902, pp. 231-239.
Ark. Bull. 113, 1913, pp. 493-505.
Treat, (pos.), Phytopath. 6, pp. 152-158, 288-292 (1916).
Fly-speck {Leptothyriurn carpophilitm, Pass.).
N. J. Agr. Exp. Sta., Rep. 18, 1897, pp. 378-383 (1898).
Fruit Spot {Fabrcea maculatum, (Lev.), Atk.).
Leaf-blight {FabrcRa maculaium (Lev.), Atk. and Cercospora minima, Tracy and
Earle).
Del. Agr. Exp. Sta., Bull. 13, pp. 4-6 (1891).
N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 145, p. 611 (1898).
Heald and Wolf, Plant Disease Survey, San Antonio, Texas,
(1912).
Treat (pos.), R. L Agr. E.xp. Sta., Bull. 31, pp. 5-9 (1895).
Cf. Quince (Leaf-spot).
Leaf-spot {Septoria piricola, Desm.).
Descr. Illus., Treat, (pos.), N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 145, pp.
597-611 (1898).
Rust {Gymnos porangium globosum, Farl.).
Occ, Conn. Agr. Exp. Sta., Rep. 14, 1890, p. 98 (1891).
Scab {Fusicladinm pirinmn (Lib.), Fckl. = Ventnria pirina, Aderh.).
Descr. Illus., N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 145, pp. 616-620 (1898).
Treat, (pos.), Vt. Agr. Exp. Sta., Bull. 44, pp. 85-90 (1895).
Cf. Apple (Scab).
Shot-hole {CyHndrosporium padi, Karst.).

Bull. 212, Colo. E.xp. Sta. (October, 1915).
Pecan
(Iliioria pcain (Marsh.), Butt.)i
Anthracnose (Glomerella cingulata (Stonem) S. & S.).
Brown Leaf-spot {Ccrcospora fusca, Rand).

Crown-gall (Pseudomonas tumefaciens, E. F. Sm. & Towns.).
Kernel -spot (Coniothyrium caryogenum, Rand).
Leaf-blight {Septoria carya, Ell. & Ev.).
Leaf -blotch (Mycosphcerella convcxula (Schw.), Rand).
Phytopath. i, pp. 133-138 (iQ")-
Mildew {Micros phccra alni (Wallr.), Wint.
Nursery-blight {Phyllosticia caryce, Pk).
Scab {Fiisidadium effiisiim, Wint.).
Orton, W. A., Science, new ser. 21, p. 503 (March 31, 1905).
Peony
{Pceonia officinalis, L.)
Mold {Botryiis pceonia, Oud.)
Peppers
{Capsicum anmmm, L.)
Anthracnose {Colletolrichnm nigrum, Ell. & Hals, and Glmosporium piperalim, Eil.
& Ev.).
Descr. N. J. Agr. Exp. Sta., Rep. 11, 1890, pp. 358-359 (1891).
Illus.,
Cf. Agr. Exp. Sta., Rep. 13, pp. ^^2-7,31 (1893)-

N. J.
Fruit-rot {Glceosporium piperatum, Ell. & Ev.).
Mold {Macros porium sp.).
Occ, N. Agr. E.xp. Sta., Rep. 15, 1894, p. 351 (1895).
J.
Leaf-spot {Ccrcospora capsici, Heald & Wolf).
Persimmon
{Diospyros spp.)
Black Leaf-spot {Ccrcospora fuliginosa. Ell. & Kellem).

Leaf-spot {Ccrcospora kaki, Ell. & Ev.).
Fruit- rot {Phyllosticia biformis, Heald & Wolf).
1 Rand, Frederick V. : Some Diseases of Pecans, Journal of Agricultural Re-
search I, pp. 303-337, June ID, 1914.
{Agaricus.
Ccrcospora air a, EH. & Ev.
Glososporium diospyri, EH. & Ev.
See N. C. Agr. Exp. Sta., Bull. 92, p. 116 (1893).
Phlox
(Phlox spp.)
Leaf-spot {Scptoria divaricala, Ell. & Ev.).
Pine
{Pinus spp.)i
Blister-rust {Cronartium ribicola, Fisch. (= Peridermium slrobi (Kleb.), Spauld.).

Bull. 206, Bureau of Plant Industry, 191 1; American Forestry (Feb., Dec,
1916).
Bluing (Ceraloslomdlapilifera (Fr.)Wmt.).
von Schrenk, U. S. Bureau Plant Industry, Bull. 36 (1903).
Chalky Quinine Fungus (Fames laricis (Jacq.), Murr.).
Meinicke, 1914, p. 44.
Dry-rot (Trametes Pini (Brot.) Fr., and T. radiciperda 'H.axiig =Fomes annosiis
(Fr.), Cke.).
(1900).
Gray Leaf- tip (Hypoderma Desmazicri, de By.).
Leaf-blight (Lophodermium brachysporum, Rostr. = Hypoderma hrachysporum
(Rostr.), Tubeuf.).
Stevens & Hall, p. 445 (1910).
Needle Disease (Hypoderma deformans, Weir on Finns ponderosa, Laws. Journ. Agric.
Res. VI: 277-288, May 22, 1916).
Pine Gall (Peridermium Harknessii Moore =P. cerebrum Pk.).
Meinecke, E. P., Forest Tree Diseases Common in California and Nevada,
U. S. Forest Service (1914).
Punk-rot (Polyporus pinicola, Aik.^Fomes ungulalus (Schraeff) Sacc).
Bull. 193, Corn. Univ. Agr. Exp. Sta. (June, 1901).
Red-rot (Polyporus ponderosus, v. Schr.).
U. S. Bureau of Plant Industry, Bull. 36 (1903).
Root- rot (Polyporus Schweinitzii, Fr.).
Descr. Illus., U. S. Dep. Agr., Div. Veg. Phys. & Path., Bull. 25, pp. 18-24 (1900).
*The twelve species of Peridermium found in American pines are described by

Arthur and Kern in North American Species of Peridermium, Bull. Torr. Bot.
Club 33, pp. 403-438, 1906.
Rust {Coleosporium pini, GaX\=Gallowaya pinl (GalL), Arth. and Peridennium

piriforme, Pk.).
Descr., Journ. Mycol., Vol. VII, p. 44 (1891).
Wet- rot (Polyporus subacidus, Pk. ?).
Descr. Illus., U. S. Dep. Agr., Div. Veg. Phys. & Path., Bull. 25, pp. 44-49.
(1900).
Pink (Sweet William)
(Dianthus barbalus, L.)
Mold (Heterosporium echinulalum (Berk.), Cke.).

Rust {Puccinia arenarice (Schum.), Wint.).
Descr. lUus., N. J. Agr. Exp. Sta., Rep. 13, 1892, pp. 278-280 (1893).
Plum
{Primus spp.)
Bacterial Leaf-spot (Pêz^/owoHd^ />«<;»", E. F. Sm.).

Heald & Wolf, Plant Disease Survey, San Antonio, Texas (191 2).
Black-knot (Plourightla morbosa (Schw.), Sacc).
Descr. Illus. Treat., Ky. Agr. Exp. Sta., Bull. 80, pp. 250-256 (1899).
Cf. Cherry (Black Knot).
Canker (Neclria ditissima, Tul.).
Descr., See U. S. Dep. Agr., Exp. Sta. Rec, IX-8, pp. 761-762 (1898).
Die-back {Valsa leticostoma (Pers.), Fr.).
Heald & Wolf, Plant Disease Survey, San Antonio, Texas (191 2).
Fire-blight (Bacterial).
Occ, Conn. Agr. Exp. Sta., Rep. 18, 1894, pp. 117-118 (1895).
Fruit-mold {Sclerotinia frnctigena, Kze. & Schm.).
Descr. Illus., Oreg. Agr. Exp. Sta., Bull. 57, pp. 3-12 (1899).
Mo. Agr. Exp. Sta., Bull. 31, pp. 16-18 (1895).
Valleau, W. D.: Varietal Resistance of Plums to Brown Rot, Journ. Agr. Re-
search V, pp. 365-395 (1915)-
Cf. Cherry (Fruit-mold).
Leaf-curl {Exoascus mlrabllis, Atk.).
Treat, (pos.). Conn. Agr. Exp. Sta., Rep. 20, 1896, p. 281 (1897).
Leaf-spot {Cylindrosporium padi, Karst. and Phyllostida congesta, Heald & Wolf).
Descr. N. Y. Agr. Exp. Sta., Rep. 5, 1886, pp. 293-296 (1887).
Illus.,
N. Y. Agr. Exp. Sta., Rep. 6, 1887, pp. 347-35© (1888).

Treat, (pos.), U. S. Dep. Agr., Div. Veg. Path., Bull. 7, p. 30 (1894).
N. Y. Agr. Exp. Sta., Rep. 15, '96, pp. 384-401 (1897).
Cf. Cherry (Leaf -spot).
Plum-pockets {Exoascus pruni, Fckl.).

Descr. lUus., U. S. Dep. Agr., Rep. for 1888, pp. 366-369 (1889).
N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 73, pp. 329-330 (1894),
Treat, (rec), N. C. Agr. Exp. Sta., Bull. 92, p. iii (1893).
Powdery Mildew {Podosphccra oxyacanthcB (DC), de By.).
See Cherry (Powdery Mildew).
Root-rot {Armillaria mellea, Vahl).
Bull. 59, pp. 14 (1903).
Rust {Puccinia pruni spinosa, Fers. = Tranzschelia punclala (Pers.), Arth.).
Treat, (pos.), Journ. Mycol., Vol. VII, pp. 356-362 (1894).
Cf. Cherry (Rust).
Scab {Clados pori urn carpophilum, Thiim).
Descr. Illus., Iowa Agr. Exp. Sta., Bull. 23, pp. 918-920 (1894).
Cf. Cherry and Peach (Scab).
Shot-hole (Cylindros porium padi, Karst).
Pomegranate
{Punka granatum, L.)
Leaf-spot {Ccrcospora lylhraccarum, Heald & Wolf).
Pomelo
(Citrus decumana, Murr.)
Anthracnose (CoUctotrichum glceosporioides, Penz.).

Fla. Bull. 74, pp. 159-172, jdIs. 4 (August, 1904).
Canker {Pscudomonas citri, Hasse).
Journ. Agr. Research VI, pp. 69-99 (April, 1916).
Poplar
{Populus spp.)
Anthracnose (Marssonia populi (Lib.), Sacc).

Leaf-spot (Septoria musiva, Pk.) and {Septoria populicola, Pk.).
Rust (Melampsora populina (Jacq.), Lev.).
Treat, (pos.), Mass. Agr. Exp. Sta., Rep. 7, 1894, p. 20 (1895).
Potato :
{Solanum tuberosum, L.)
Anthracnose {Vcrmicularia, sp.).
Black-leg {Bacillus phylophlhorus, Appel).

Orton, W. A., Potato Tuber Diseases, Farmers' Bull. 544 (1913).
Blight (Bacillus solanaccamm, E. F. Sm.).

Descr. Illus. Treat, (rec), U. S. Dep. Agr., Div. Veg. Phys. & Path., Bull. 12
(1896).
Chytridiose or Black Scab (Synchytriutn cndohioikum (Schilb.) Vcxa\a.l = Chryso-
phlyclis cndohioiica, Schilb.)
Downy Mildew or Rot {Phytophthora infcslans, de By.).
Descr. Illus., U. Dep. Agr., Rep. for 1888, pp. 337~33S (1889).
S.
N. H. Agr. Exp. Sta., Bull. 22, pp. 3-5 (1894).

Vt. Bull. 168, pp. 100, pis. 10, figs. 10 (August, 191 2).
Conn. Rep., 753-774 (1909)-
pt. 10, pp.
Melhus, I. E., Hibernation in the Irish Potato, Journ. Agr.
Research V, pp. 71-102 (1915).
Treat, (pos.), U. S. Dep. Agr., Farm. Bull. 91 (1899).
Dry-rot {Fusarium solani (Mart.) Sacc. and F. radicicola, WoUenw.).
Occ, 111. Agr. Exp. Sta., Bull. 40, p. 139 (1895).
Internal Browning (Bacterial ?).
Descr., Agr. Exp. Sta., Bull. 40, pp. 138-139 (1895).

111.
N. Y. Agr. Exp. Sta., Bull. loi, pp. 78-83 (1896).

Leaf-blotch {Cercospora concors (Casp.) Sacc).
Leaf-mold or Early-blight (AUernaria solani (Ell. & Mart.), Jones & Grout).
Descr. Illus., Del. Agr. Exp. Sta., Rep. 4, 1891, pp. 58-59 (1892).
N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 140, p. 393 (1897).
Vt. Agr. Exp. Sta., Bull. 72, pp. 16-25 (1899).
Treat, (pos.), U. S. Dept. Agr., Farm. Bull. 91, pp. 5-7 (1899).
Wise. Rep., pp. 343-354, figs. 7 (1907)-
Leak (Pylhium de Baryaniim, Hesse.)
Journ. Agr. Research VI, pp. 627-640, pi. i (1916).
Powdery^ Scab {Spongospora siihkrranea).
W.
Orton, Potato Tuber
A., Farm. Bull. 544 (1913)-
Diseases, U. S.
U. S. Dept. Agr., No. 82 (1914).
Bull.,
Powdery Dry-rot {Fusarium Iriclwlhecoides Wollenw.).
Orton, W. A., U. S. Farm. Bull. 544 (1913).
Pratt, Journ. Agric. Res., VI: 817-831, Aug. 21, 1916.
Root-rot {Entorrhiza solani, Fautr.).
See U. S. Dept. Agr., Exp. Sta. Rec, VII-io, p. 873 (1896).
Scab (Actinomyces chrcmogenes, Gasp.).
1 Carpenter, C. W.: Some Potato Tuber-rots Caused by Species of Fusarium,

Journal of Agricultural Research V, pp. 183-209 (Nov. i, 1915).
^ Consult Melhus, I. E., Rosenbaum,
J. and Schultz, E. S.: Studies of Spon-
gospora subterranea and Phoma tuberosa of the Irish Potato, Journ. Agr. Research
Vn, pp. 213-253, October, 1916, also IV, pp. 265-278.
Cf. W. Va. Agr. Exp. Sta., Sp. Bull. 2, pp. 97-111 (iSqS).
Treat, (pos.) Journ. Agr. Research IV, pp. 129-133 (1915).
(Cor. Sub.) Mich. Agr. Exp. Sta., Bull. 108, pp. 38-45 (1894).
Ind. Agr. Exp. Sta., Bull. 56, pp. 70-80 (1895).
Conn. Agr. Exp. Sta., Rep. 19, 1895, pp. 166-176 (1896).
(Formalin) U. S. Dep. Agr., Farm. Bull. 91, pp. 9-10 (1899).
Scurf (Rliizoctonia solani, Kiihn = Coriicium vagum, B. & C, var. solani, Burt.).
Silver-scurf (S pondylocladium alrovircns, Harz.).
Orton, U. S. Farm. Bull. 544 (1913).
Journ. Agr. Research VI, pp. 339-350 (June, 1916).
Stem-blight (Fusarium acuminatum, Ell. & Ev. ?).
Descr., N. Y. Agr. Exp. Sta., Bull. loi, p. 85 (1896).

Cf. N. Y. Agr. Exp. Sta., Bull. 138, pp. 632-634 (1897).
Stem-rot {Coriicium vagum, Bri. & Cav., var. solani, Burt.).
Cal. Bull. 70, pp. 1-20, pis. 12 (March, 1902).
Tuber-rot {Fusarium oxysporum, Schlecht).
Orton, U. S. Farmer's Bull. 544 (1913).
Bull., U. S. Dep. Agr., No. 64 (1914).
Wart {Synchytritim endohioticiim (Schilb.), Percival).
Orton, W. A., Potato Tuber Diseases, U. S. Farmer's Bull. 544 {igi^,).
Wet-rot {Bacterial).
Descr., Del. Agr. Exp. Sta., Rep. 4, 1891, pp. 54-57 (1892).
Wilt {Bacillus solanacearum, E. F. Sm.).
Yellow-blight {Sclerotinia liberliana, Fckl.; Syn. Peziza postuma, Berk. & WUs. ?).
Primrose
{Primula, spp.)
C
Phyllosticta primulicola, Desm.
Ramularia primulce, Thm.
Miscellaneous Fungous Diseases.
CoUetolrichum primulce, Hals.
[ Ascochyta primula. Trail.
See N. J. Agr. Exp. Sta., Rep. 15, 1894, pp. 377-380 (1895).
Privet
{Ligustrum vulgare, L.)
Anthracnose {Glceosporium cingulatum, Atk.).

Descr. Illus., N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 49, PP- 306-314 (1892).
Leaf-spot {Cercospora adusta, Heald & Wolf, C. Hgustri, Roum and Phyllosticta
ovalifolia, Brun.)
Quince
{Pirus cydonia, L.)
Black-rot {Sphosropsis malorum. Berk.).

Descr. Illus., N, J. Agr. Exp. Sta., Bull. 91, pp. 8-10 (1892).
Treat, (rec), Conn. Agr. Exp. Sta., Bull. 115, pp. 6-7 (1893).
Fire-blight {Bacillus amylovorus (Burr.), Trev.).

See Apple and Pear (Fire-blight).
Leaf-blight {Entomosporium maculatum, Lev=Fabraea maciihUum (Lev.) Atk.
Descr. lUus., See Pear (Leaf-spot).
Mold {Sclerotinia cydonice, Schellenb.).
Pale-rot (Phoma cydonia, Sacc. & Schulz.?).
Descr. N. J. Agr. Exp. Sta., Bull. 91, pp. lo-ii (1892).
Illus.,
Ripe-rot or Anthracnose {Glceosporiiim fructigenum, Berk.).

See Apple and Grape (Ripe-rot).
Rust {Gymnosporangimn clavipes, C. & P., Syn. Rcestelia aurantiaca, Pk.).
Descr.Îllus., N. J. Agr. Exp. Sta., Bull. 91, pp. 2-5 (1892).
Mass. Hatch Rep., pp. 61-63 (1897).
Radish
{Raphantis sativus, L.)
Club-root {Plasmodiophora brassica;, Wor.).

Occ, N. J. Agr. Exp. Sta., Rep. 11, 1890, pp. 348-349 (1891).
Downy Mildew (Peronospora parasitica (Pers.) deBy.).
White-rust {Cyslopus candidus (Pers.), Lev.).
Treat, (rec), N. J. Agr. E.xp. Sta., Rep. 11, 1890, p. 350 (1891).
Raspberry
(Rubus spp.)
Anthracnose {Glceosporiiim venclum, Speg. = Gl. necator, Ell. & Ev.).

Descr. Illus., U. S. Dept. Agr., Rep. for 1887, pp. 357-360 (1888).
Ohio Agr. Exp. Sta., Bull. IV-6, pp. 124-126 (1891).
N. Y. Agr. Exp. Sta., Bull. 124, pp. 262-264 (1897).
Treat, (pos.), Conn. Agr. Exp. Sta., Rep. 23, '99, pp. 274-276 (1900).
Black-blight {Fnsarium, sp. ?).
Blue-stem {Acrostolagmiis caulophagus, Lawrence.).

Washington Bull. 108, pp. 30, figs. 28 (October, 191 2).
Cane-blight {Coniothyrimn Fuckelii, Sacc).
Descr. N. Y. Agr. Exp. Sta., Bull. 107, pp. 305-307 (1899).
Crown-gall (Possibly identical with Crown-gall of Peach, q.v.).
See Ohio Agr. Exp. Sta., Bull. 79, pp. 108-112 (1897).
Fire-blight (Bacterial).
Descr., Ohio Agr. Exp. Sta., Bull. IV-6, pp. 128-129 (1891).
Leaf-spot {Scploria riihi, Westd).

Mushroom Root- rot (Armillaria hirllca Vahl).
Ore. Sta. Bien. Rep. (1911-12).
Orange-rust {Gyntnoconia interstitial is).

Rust {Gymnoconia intcrstltialis (Schl.) v. Lagerh.).
Spur-blight {Spharclla rubina Pk.).
Wilt {Leptosphccria coniothyriiini (Fckl.) Sacc).
Red Gum
{Liqiiidamhar styracljlua, L.)'
Sap-rot {Polyslictus versicolor, (L.) Fr.).

von Schrenk, Diseases of Deciduous Forest Trees, U. S. Bur. Plant Industry,
Bull. 149 (1909).
Red Top
(Agrostis alba, L.)
Sclerotial Disease {Sclerotium rhizodes, Auersw.).

Conn. E.xp. Sta., Rep., p. 23 (1914).
Rice
{Oryza saliva, L.)
Blast {Piricularia oryza, Cav.).

Cook, Diseases of Tropical Plants, p. 99 (1913).
Smut (Tilletia corona, Scrib.).
Descr. Illus., S. Car. Agr. Exp. Sta., Bull. 41, pp. 7-1 1 (1899).
Treat, (rec.;, S. Car. Agr. Exp. Sta., Bull. 41, pp. 15-29 (1899).
Rose
{Rosa spp.)
Anthra'cnose {Glososporium rosce, Hals.).

Cane-blight {Coniothyrium Fuckelii Sacc).
Downy Mildew {Peronospora sparsa, Berk.).
'von Schrenk, Hermann: Sap-rot and other Diseases of the Red Gum, U.
S. Bureau of Plant Industry, Bull. 114, 1907, where all the important diseases
are considered.
1
Leaf-blotch {Actinonema rosa (Lib.), Fr.).

Treat, (pos.), N. J. Agr. Exp. Sta., Rep. 13, 1892, p. 281 (1893).
Leaf-spot {Splitrrella rosigena, EIL).
Occ.
Mildew {Peronospora sparsa, Berk.).
Powdery Mildew {Sphcerotheca pannosa (Wallr.), Lev.).
Descr., N. J. Agr. E.xp. Sta., Rep. 13, 1892, p. 281 (1893).
Treat, (pos.), N. J. Agr. E.xp. Sta., Rep. 13, 1892, pp. 281-282 (1893).
Rust {Phragmidium subcortkium (Schrank) Wint. and P/k speclomm, Fr.).
Descr. Illus., U. Dept. Agr., Rep. for 1887, pp. 369-372 (1888).
S.
Treat, (pos.), See U. S. Dept. Agr., Exp. Sta. Rec, X-7, p. 651 (1899).
Twig-blight {Botrylis clnerea, Pers.).
Rye
{Secalc ccrcale, L.)
Anthracnose (Collciotrichum gramincola (Ces.) Wilson).

Ergot (Claviceps purpurea, (Fr.) TuL).
Descr. Illus., S. Dak. Agr. Exp. Sta., Bull. 33, pp. 40-43 (1893).
Rust (Black-stem, Puccinia graminis, Pers., and Orange-leaf, P. rubigO'vera (DC),
Wint.).
See U. S. Dep. Agr., Div. Veg. Phys. & Path., Bull. 16, pp. 42-45 & 60.
Smut {Urocystis occulta (Wallr.), Rabh.).
Occ. Mass. Agr. E.xp. Sta., Rep. 9, 1891,
Illus., p. 247 (1892).
Treat, (pos.), see Oats and Wheat (Smut).
Stem-blight {Leptosphceria herpotrichoidcs, de Not).
Salsify
{Tragopogon porrifoliits, L.)

Rot (Bacterial).
Descr., N. J. Agr. E.xp. Sta., Rep. 11, 1S90, p. 351 (1891).
White-rust {Cystopus iragopogonis, (Pers.), Schrot.).
Occ, N. J. Agr. E.Kp. Sta., Rep. 15, 1894, p. 355 (1895).
Rust {Puccinia tragopogoni (Pers.), Cda.).
Scrub Pine
{Pinus virigitiiana, Mill)
Burl Disease {Cronarlium quercus (Brand.) Schrot).

Graves, A. H., Phytopathology IV (February, 1914).
Heart-rot {Trameles pint (Brot.) Fr.).
Leaf-cast {Galloivaya pini (Gall.), Arth.).

Rust (Coleosporium inconspicuum (Long), Hedg.).
Shaddock or GRAPE-FRtni
{Citrus decumana, Murr.)
See Lemon and Orange

Snapdragon
{Antirrhinum niajiis, L.)
Anthracnose {Colletotrichum antirrhini, Stewart).

Descr. Illus. Treat, (pos.), N. Y. Agr. Exp. Sta., Bull. 179 (1900).
Root-rot {Thielavia basicola, Zopf.).
Rust {Puccinia antirrhini, Diet. & Holway.).
Stem-rot {Phoma sp.).
Descr. Treat, (rec), N. Y. Agr. Exp. Sta., Bull. 179, pp. 109-110 (1900).
Sorghum
{Sorghum vulgare, Pers.)
Blight {Bacillus sorghi, Burrill).

Descr., Kan. Agr. Exp. Sta., Rep. i, 1888, pp. 281-301 (1889).
Treat, (rec), Kan. Agr. Exp. Sta., Rep. i, 1888, pp. 301-302 (1889).
Head-smut {Sorosporlum reilianum (Kiihn) McAlpine).
Journ. of Agr. Research II, pp. 340-371 (Aug. 15, 1914).
Descr. Illus., Kan. Agr. Exp. Sta., Bull. 23, pp. 95-96 (1891).
111. Agr. Exp. Sta., Bull. 57, pp. 335-347 (1900)-
Treat, (pos.), 111. Agr. Exp. Sta., Bull. 57, pp. 345-346 (1900).
Kernel-smut {Sphacelotheca sorghi).
BuU. 212, Colo. Agr. Exp. Sta. (October, 1915).
Journ. Agr. Research II, pp. 339-371, pis. 7 (1914).
Soy
{Soja hispida, Moench.)
Wilt-disease {Fusarium tracheiphilum, E. F. Sm.), Journ. Agric. Res. 8: 421-439,

with I pi., Mch. 12, 191 7.
Spinach
{Spinacia oleracea, Mill.)
Leaf-blight {Cercospora beticola, Sacc).

Cf. N. J. Agr. Exp. Sta., Rep. 18, 1897, p. 303 (1898).
Anthracnose {CoUetotrichum spinacecs, Ell. & Hals.).
Downy MUdew {Peronospora effiisa, (Grev.), Rabenh.).

Miscellaneous
Leaf-spot {Phyllostida chenopodii, Sacc.)."
Fungous Diseases.
Scab {Cladosporium inacrocarpum, Preuss).
^
White Smut (Entyloma Ellisii, Hals.) .
Descr. N.
Illus., J. Agr.' Exp. Sta., Bull. 70 (1890).
Treat, (rec), N. J. Agr. Exp. Sta., Bull. 70, pp. 13-14 (1890).
Spruce
{Picea spp.)i
Blight of Seedlings (Ascochyta piniperda, Lindsin = Dlplodina parasitica (Hart, Prill),

and Sderotinia Fuckeliana, deBy).
Graves, A. H., Phytopathology IV (April, 1914).
Brown-rot {Polyporus sulphureus (Bull.) Fr.).
Dry-rot (Trametes pini (Brot.) Fr. and T. abietis
Karst.).
Heart- rot {Polyporus borealis (Wahl.) Fr.). Descr. Illus., U. S. Dep. Agr.
Atkinson, BuU. 193 (Corn. Univ.) Agr. Exp. Sta. Div. Veg. Phys. & Path.
(June, 1901). Bull. 25 (1900).
Root-rot {Polyporus Schweinitzii, Fr.).
Wet-rot {Polyporus suhacidus, Pk. ?).
Squash
{Cucurbit a spp.)
Anthracnose {CoUetotrichum lagenarium (Pass.), Ell. & Hals.).

Bacteriosis or Wilt {Bacillus tracheiphilus, E. F. Sm.).
Downy Mildew {Plasmopara cubensis (Bri. & Cav.), Humph.).
Fruit-mold {Macrosporium sp.).
Powdery MUdew {Erysiphe cichoracearum, DC. and E. polygoni, DC).
Descr. Illus., See Cucumber (Powdery Mildew).
Treat, (pos.), N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 35, p. 330 (1891).
Leaf-spot {Cercospora cucurbitce, Ell. & Ev.).
Strawberry
{Fragaria spp.)
Blight {Micrococcus sp. ?).
Descr., Mass. Agr. Exp. Sta., Rep. 9, 1896, pp. 59-61 (1897).
Leaf-blotch {Ascochyta fragaria, Sacc).
Descr. Illus., N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 14, pp. 182-183 (1889).
Treat, (rec), N. Y. (Corn. Univ.) Agr. Exp. Sta., BuU. 14, p. 183 (1889).
^ For species of Peridermium on spruce consult Arthur & Kern, North American
Species of Peridermium, Bull. Torr. Bot. Club 2>2„ PP- 403-438, 1906.
.
Leaf-spot {Aposphceria sp.).

Treat, (rec.) N. J. Agr. Exp. Sta., Rep. 14, 1893, pp. 331-332 (1894)-
Leaf-spot {Mycosphcerclla fragarice, (Tul.) Lindau).
N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 14, pp. 171-181 (iJ
Oregon State Biennial Rep. p. 268 (1911-12).

Leaf-spot {Mycos phcerella fragarice (Tul.), Lindau).
Treat, (pos.), U. S. Dep. Agr., Rep. for 1890, p. 397 (1890).
Conn. Agr. Exp. Sta., Bull. 115, p. 14 (1893).
Powdery Mildew (Sphcerotheca Castagnei, Lev.).
Descr., N. Y. Agr. E.xp. Sta., Rep. 5, 1886, pp. 291-292 (1887).
Descr. Mass. Agr. Exp.
Illus., Sta., Rep. 10, 1892, p. 239 (1893).
Treat, (rec), Mass. Agr. Exp. Sta., Rep. 10, 1892, pp. 243-245 (1893).
Rot (Sphceroncemella fragarice, Stev. & Pet.).
Phytopath. VI, pp. 258-266 (1916).
Sugar-C.4ne1
{Saccliarum officinarum, L.)
Bundle-blight {Pseudomonas vascularum (Cobb.) E. F. Sm,).

Cacao Disease (Diplodia cacaoicola, Henn).
Cook, Disease of Tropical Plants, p. 85 (1913).
Iliau {Gnomonia iliau, Lyon).
Cook, p. 85 (1913)-
Phytopath. 3, pp. 93-98 (191 3).
Leaf-spot {Ccrcospora longipes, Butler).
Cook, p. 89 (1913).
Macrosporium gramimim, Cke.
Miscellaneous Diseases. ,,. ^^..,
1 .. ,„ ^ ,^7 o -ht ^
Uromyces Kuhnii (Krug.j, Wakk. & Went.
.. 1 1
Pineapple Disease {Thielaviopsis ethacelicus, Went).

Red- rot {Collelotrichum falcatum, Went).
Rind Disease {Trichosphceria sacchari, Mass.).
See U! S. Dept. Agr., Exp. Sta. Rec, X, pp. 56-57, '98, and XI, p. 759 (1900).
Ring-spot {Leptosphceria sacchari, de Haan)
Cook, p. 89 (1913).
Smut {Ustilago sacchari, Rabenh.).
Stool Disease (Marasmius sacchari, Wakker).
Cook, p. 92 (1913).
1
Cf. Edgerton, C. W.: Some Sugar-cane Diseases, Bull. 120, La. Agric. Exper.
Sta., July, 1910; Cobb, N. A.: Fungous Maladies of the Sugar Cane, Bull. 6, Exper.
Sta., Hawaiian Sugar Planters Assoc, 1906.
Sunflower
(Helianthtis anntius, L.)
Black-rot {SphcBronema fimbriatum (Ell. & Hals.) Sacc).

Duggar, p. 348 (1909).
Dry-rot (Phoma batata, Ell. & Hals.).
Duggar, p. 344 (1909).
Root-rot {Cortkium vagum, B. & C. var. solani, Burt.).
Duggar, p. 444 (1909).
Rust (Puccinia lielianiki, Schw.).
Sweet Pea
{Lathyrus odoratus, 'L.y
Anthracnose {Glomcrella rufomacidans (Berk.), Spauld. & v. Schr.).
Powdery Mildew (Erysiphe polygoni DC).

Root-rot (Thielavia basicola, Zopf.; Rhizoctonia {Corticium vagum, Bri. & Cav.
var. solani Burt); ChcBlomiiim spirochete, Pall.; Fusarium lathyri, Taub.
& Manns).
Stem or Collar-rot {Sclerotinia libertiana, Fckl.).
Streak {Bacillus lathyrii, Manns & Taub.).
Sweet Potato
{Ipomceo batatas, Lam.)
Black-rot (Sphoeronema (Ceratocystis) fimbriata (Ell. & Hals.), Sacc).

Journ. Mycol., Vol. VII, pp. 1-9 (1891).
Treat, (pos.), Md. Bull. 60, pp. 147-168, figs. 17 (March, 1899).
(rec), U. S. Dept. Agr., Farm. Bull. 26, p. 21 (1895).
Charcoal-rot {Sclcrollum bataticola, Taub.).
Foot- rot (Plenodomus destruens. Hart.).
Phytopathology. 3, pp. 242-245 (1913).
Taubenhaus & Manns, Bull. 109, Del. Agr. Exp. Sta., May, 1915.
Journ. Agr. Research I, p. 251
Java- rot {Las iodiplodia tuber icola, Ell. & Ev.).
Soil-rot {Acrocystis batatae, Ell. & Hals.).
Descr. Illus., N. J. Agr. Exp. Sta., Bull. 76, pp. 14-18 (1890).
Treat, (pos.), N. J. Agr. Exp. Sta., Rep. 20, 1899, pp. 345-354.
N. J. Spec. Bull. 5, February, 1900, pp. 22-31, pis. 3 (1900).
1 Consult Taxjbenhaus, J. J.: The Diseases of the Sweet Pea, Bull. 106, Del.
Agric. Exper. Sta., November, 1914.
30
.
Stem-wt {Fusanum hyperoxysporum Wollenw.).

U. S. Farmers' Bull. 714, March 11, 1916.
Phytopath. 4,pp. 277-303 (1914).
Dry-rot {Phoma batatce, Ell. & Hals, conidial stage of Diaporlhe
batatatis (Ell. & Hals.), Hart & Field).
Leaf-spot {Phyllosticla bataticola, Ell. & Mart.)
U. S. Farmers' Bull. 711.
Scurf {Monilochates infuscans (Ell. & Hals.) Hart).
Miscellaneous N. J. Bull. 76, pp. 25-27 (Nov., 1890).
Diseases. | Soft-rot {Rhizopus nigricans, Ehrb.).
Phytopath. 4, pp. 305-320.
Trichoderma Rot (Trickoderma Koningi, Oud.).
Taubenhaus & Manns, Bull. 109, Del. Agr. Exp. Sta., May,
1915-
White-rot {Penicillium sp.).
White-rust {Cystopus ipomaos-pandurance (Schw.), Farl.).

Descr. Illus., N. J. Agr. Exp. Sta., Bull. 76 (1890).
Md. Agr. Exp. Sta., Bull. 60 (1899).
Treat, (rec), U. S. Dept. Agr., Farm. Bull. 26 (1895).
Vine- wilt {Fusarium batatatis, Wollenw.).
Taubenhaus and Manns, Bull. 109, Del. Agr. Exp. Sta., May, 1915.
Wollenweber, H. W., Journ. Agr. Research 2, pp. 251-283 (191 1).
Sycamore
(Platanus occidentalis, L.)
Anthracnose {Glceosporinm nervisequum (Fckl.), Sacc, stage of Gnomonia veneta

(Sacc. & Speg.) Kleb.).
Blight {Gnomonia veneta (Sacc. & Speg.), Kleb.).
Descr. U. S. Dep. Agr., Rep. for 1888, pp. 387-389 (1889),
Illus.,

Cf. Journ. Mycol., Vol. V, pp. 51-52.
Gar. and For., X-488, pp. 257-258.
Tea
{Thea chinensisY
Bark Disease (Corticium javanicum, Zimm. = C Zimmermani, Sacc. & Syd,).

Blister- blight {Exobasidium vexans, Massee.).
Copper-blight {Loestadia tliece, Show).
Grey- blight (Pestalozzia guepini, Desm.).
Horsehair-blight (Marasmius sarmentosus. Berk.).
Internal Stem Disease {Massaria theicola, Petch.).
Red-rust {Cephaleus mycoidea, Karst.).
1 For all consult Cook, Diseases of Tropical Plants, pp. 170-180 (1913).
.
Root Fungus {Roscllinia radicipcrda, Massee.)-

Soot- blight (Capnodium Footii Berk, and Desm.).
Thread- blight (Stilbum nanum, Massee.).
Teosinte
{Euchlcena mexicana, Schrod.)
Smut {Ustilago zeoe (Beckm.), Ung.).
Timber
Decay (Stereum fmstulosum (Pers.), Fr.).

von Schrenk, Diseases of Deciduous Forest Trees, U. S. Bureau of Plant Industry,
Bull. 149 (1909).
Sap-rot {Dadalea quercina (L.), Pers.)-
Mainly on oak timber, von Schrenk (1909).
Timothy
{Phleum pratense, L.)
Ergot {Claviceps purpurea (Fr.), Tul.).

Phytopath. 4, pp. 20-22 (1914).
Rust {Puccinia phlei-pratensis, Eriks & Henn.)
Phytopath. 4, pp. 20-22 (1914).
Smut {Ustilago stricefomits (West.), Niessl.).
Tobacco
{Nicoliana labacum, L.)
Black- rot {Sterigmatocystis nigra v. Tieg.)

Wise. Res. Bull. 32, pp. 63-83, figs. 7 (June, 1914).
Blue Mold {Fungus indet.). .
Brown-spot {Macros poritim longipes, Ell. & Ev.)

Cf. U. S. Dept. Agr., Exp. Sta. Rec, XII-4, p. 359 (1900).
"Damping-o2" {Altertiaria tenuis, Nees).
Downy Mildew
f '^ hyoscyami, deBy.).
1^
J^f'^^f
| {Phytophthora nicottanoe, de Haan).
Leaf-blight {Cercospora nicotianoe, Ell. & Ev.).
Descr. Conn. Agr. E.xp.
Illus., Sta., Rep. 20, 1896, pp. 273-277 (1897).
Mosaic, Bull. U. S. Dept. Agr., p. 40 (1914).
. .
Pole-burn (Fungi and Bacteria).

Powdery Mildew {Erysiphe cichoracearum, DC, Syn. E. lamprocarpa (Wallr.), Lev.).
Root-rot {Thielavia basicola, Zopf.).
Gilbert, W. W., Bull. 158, U. S. Bur. of Plant Industry (1909).
Conn. Rep., pt. 5, p. 342 (1906.)
Phytopath. 6, pp. 167-181 (,1916).
Stem-rot (Botrylis longihradiiata, Oud.)
Cook, Diseases of Tropical Plants, p. 149 (191 3).
Treat, (rec). Conn. Agr. Exp. Sta., Rep. 15, 1891, pp. 185-186 (1892).
White-speck {Macros porium tabacinum, Ell. & Ev.).
Cf. Conn. Agr. Exp. Sta., Rep. 20, 1896, p. 276 (1897).
Tomato
{Lycopersicum esculentum, Mill.)
Anthracnose {Collctotrichum phomoides (Sacc), Chester).

Cf. Del. Agr. Exp. Sta., Rep. 6, 1893, pp. 111-115 (1894).
Nebr. Rep., 1907, pp. 1-33, figs. S3-
Me. Agr. Exp. Sta., Rep. for 1893,
Treat, (rec), p. 155 (1894).
Blight {Pscudomonas solanacearum, E. F. Sm.).
Descr. Illus., See Egg-plant (Blight).
La. BuU. 142, pp. 1-23, figs. 3 (October, 1913).
Treat, (pos.), Md. Agr. Exp. Sta., Bull. 54, pp. 123-125 (1898).
Fla. Agr. Exp. Sta., Bull. 47, pp. 133-136 (1898).
Blight {Sclerotium sp.).
Descr., Fla. Agr. Exp. Sta., Bull. 21, pp. 25-27 (1893).
Ala. Agr. Exp. Sta., Bull. 108, pp. 28-29 (iQoo)-
Treat, (pos.), Fla. Agr. Exp. Sta., Bull. 21, pp. 32-36 (1893).
Downy Mildew {Pliyloptliora infestans (Mont.), deBy.).
Fruit-rot {Macrosporium solani, E. «fe M. and Phoma dcstructiva, (Plowr.),
Jamies.)
Descr., Ala. Agr. Exp. Sta., Bull. 108, pp. 19-25 (1900).
Cf. N. Y. Agr. Exp. Sta., Rep. 3, 1884, pp. 379-380. 1885.
Cf. N. Y. Agr. Exp. Sta., Bull. 125, pp. 305-306. 1897.

Journ. Agr. Research 4, p. i (April 15, 1915)-
Leaf-blight {Cylindros porium sp.)

Descr.,N. Y. Agr. Exp. Sta., Rep. 14, 1895, p. 529 (1896).
Treat, (rec), N. Y. Agr. E.xp. Sta., Rep. 14, 1895, pp. 530-531 (1896).
Leaf-mold {AUernaria solani (Ell. & Mart.), Jones & Grout). ,
.
Descr., Fla. Agr. Exp. Sta., Bull. 47, pp. 124-125 (1898).
Treat, (pos.-), Fla. Agr. E.xp. Sta., Bull. 47, pp. 125-127 (1898).
Leaf-spot {Septoria lycopersici, Speg.).
Descr. Illus., Del. Agr. Exp. Sta., Rep. 7, 1894-95, p. 123 (1895).
Ohio Agr. Exp. Sta., Bull. 73, p. 241 (1897).
Treat, (pos.), Va. Bull. 192, pp. 16, figs. 9 (April, 191 1).
Ala. Agr. Exp. Sta., Bull. 108, pp. 32-33 (1900).
Rust (Macrosporium solani, Ell. & Mart.).
Scab (Cladosporium fulvum, Cke.).
Treat, (pos.), U. S. Dep. Agr., Sec. Veg. Path., Bull. 11, p. 47 (1890).
Ala. Agr. Exp. Sta., Bull. 108, p. ^^ (1900).
Wilt {Fusarium lycopersici, Sacc).
Trumpet Creeper
(Tecoma radicans (L.) Jass.)
Leaf-blight (Cercospora sordida, Sacc.) '

.
Duggar, p. 315 (1909).

Leaf-spot {Septoria lecomce, Ell. & Ev.).
Tulip Tree
{Liriodendron tulipifcra, L.)
Leaf -blight {Glceosporium liriodendri, Ell. & Ev.).

Sap-rot {Pclyst ictus versicolor (L.), Fr.).
von Schrenk, H., Diseases of Deciduous Forest Trees, U. S. Bur. of Plant In-
dustry, Bull. 149 (1909).
Turnip
{Brassica campcstris, L. and B. rapa, Linn.)
Brown-rot (Fseudomonas campestris (Pam.), E. F. Sm.).

Descr. Illus., Iowa Agr. Exp. Sta., Bull. 27, pp. 130-134 (1895).
Club-root (Plasmodiophora brassicce, Wor.).
Descr. Illus.,- See Cabbage (Club-root).
Treat, (pos.), N. J. Agr. Exp. Sta., Rep. 20, '99, pp. 354-367 (1900).
Downy MUdew {Peronospora parasitica (Pers.) deBy.).
Occ, Mass. Agr. Exp. Sta., Rep. 8, 1890, p. 222 (1891).
Dry-rot (Plioma brassicce, Thiim ?).
See U. S. Dept. Agr., E.xp. Sta. Rec, XII-3, p. 256 (1900).
Conn. Exp. Sta., Rep., p. 355 (191 2).
. .
Leaf- mold {Macros porium herculeum, E. & M.).

Descr. Illus., N. Y. Agr. Exp. Sta., Rep. 15, '96, pp. 451-452 (1897).
Powdery Mildew (Erysiphe polygoni, DC).
Occ, N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 61, pp. 305-306 (1893).
White-rust (Cystopus candidns, (Pers.) Lev.).
Occ, Mass. Agr. Exp. Sta., Rep. 8, i8go, p. 222 (1891).
Tree of Heaven
{Ailanthus glandulosa, Desf.)
Shot- hole (Ccrcospora glandiilosa, Ell. & Kell.).
Verbena
{Verbena sp.)
Powdery Mildew {Erysiphe cichoraccarum, DC).

Occ, N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 37, p. 405 (1891).
Treat, (pos.), N. Y. (Corn. Univ.) Agr. Exp. Sta., Bull. 37, p. 405 (1891).
Vetch
( Vicia spp.)

Duggar, p. 227 (1909).
Rust {Uromyces pisi (Pers.) de By).
Duggar, p. 398 (1909).
Violet .
{Viola odorata, L. and V. tricolor, L.)
Anthracnose {Glceosporinm violce, B. & Br.).

Anthracnose {Colletotrichum violce-iricoloris, Smith).
Descr., Mass. Agr. Exp. Sta., Rep. 11, '98, pp. 152-153 (1899).
Treat, (pos.), Mass. Agr. Exp. Sta., Rep. 11, '98, p. 153 (1899).
Gall or Chytridiose {Cladochylrium violce, Berl.).
See U. Dep. Agr., Exp. Sta. Rec, XI-3, p. 261 (1899).

S.
Downy Mildew {Peronospora violas, de By.)

Dry-rot {Merulius lacrymans (Jacq.) Fr.)
See U. S. Dep. Agr., Exp. Sta. Rec, XI-io, p. 947 (1900).
Leaf-bhght {Cercospora violce, Sacc).
Occ Illus., N. J. Agr. Exp. Sta., Rep. 15, 1894, pp. 3S4-386 (1895).
Treat, (rec) N. J. Agr. Exp. Sta., Rep. 15, 1894, pp. 386-389 (1895).
. 1
Leaf-mold or Spot Disease {AUernaria violcc, GaH. & Dors.).

Descr. Illus. Treat., U. S. Dep. Agr., Div. Veg. Phys. & Path., BulL 23 (1900).
Leaf-spot {Phylloslida viola;, Desm. and AUernaria viola;, GaU. & Dors.).
Descr., Mass. Agr. Exp. Sta., Rep. 10, 1892, pp. 231-232 (1893).
Treat, (rec.) Mass. Agr. Exp. Sta., Rep. 10, 1892, pp. 232-235 (1893).
N. J. Agr. Exp. Sta. Rep. 15, 1894, pp. 286-389 (1895).
Root-rot (Tkielavia basicola, Zopf).
Descr. Conn. Agr. Exp. Sta., Rep. 15, 1891, pp. 166-167 (1892).
White Mold (Zygodcsmus albidiis, Ell. & Hals.).
Virginia Creeper
(Ampelopsis quinquefolia, Michx.)
Leaf-spot {Phylloslida ampdopsidis, E. & M.) =Laestadia Bidwellii (Ell.). V. & R.
Walnut
(Juglans regia, L.)
Bacteriosis {Pseudomonas jnglandis, Pierce).

Ore. Sta. Rep., p. 260 (1911-12).
See U. S. Dep. Agr., Exp. Sta. Rec, Vol. XI, p. 261 (1899).
Cal. Bull. 231, pp. 320-383, figs. 19 (August, 191 2).
Leaf-blight {Marsonia juglandis (Lib.) Sacc. oi Gnomonia leptosiyla (Fr.) Ces. & deN.
Leaf-spot {Ascochyta juglandis, Boltsh. and Phleospora muUimaculans, Heald &
Wolf.).
Leaf Disease (Cylindrosporium juglandis, Wolf.)
Mycologisches Centralblatt 4, p. 65 (1914).
Watermelon ;
(Cilrullus vulgaris, Schrad.)
Anthracnose (Colletotrichum lagenarimn (Pass.), Ell. & Hals.).

Treat, (neg.), Del. Agr. Exp. Sta., Rep. 5, 1892, p. 79 (1893).
Cf. N. J. Agr. Exp. Sta., Rep. 13, pp. 326-330 (1892).
Del. Agr. Exp. Sta., Rep. 5, pp. 75-79 (1892).
Downy MUdew (Plasmopara cubensis (Bri. & Cav.), Humph.).
See Cucumber (Downy Mildew).
Leaf-blight (Cercospora citrullina, Cke.).
Occ, Ohio Agr. Exp. Sta., Bull. 105, p. 232 (1899).
Leaf-mold (AUernaria brassica, Sacc, var. nigrescens, Regel.).
See Melon (Leaf-mold).
Leaf-spot {Phyllosticta sp. and (?) Spharella sp.).
Descr. Ulus., DeL Agr. Exp. Sta., Rep. 5, 1892, pp. 75-78 (1893).
Wheat
{Trilicum viilgare, L.)
Blight (Mysirosporium abrodens, Neum.).

Chytridiose {Pyroctontim spharicum, Prunet).
See U. S. Dept. Agr., Exp. Sta. Rec, VI-3, pp. 226-227 (1894).
Ergot (Claviceps purpurea, (Ft.) Tul.).
See Rye (Ergot).
Foot-rot [Ophlobolus & Leptospharia).
See U. S. Dept Agr., Exp. Sta. Rec, IX-ii, p. 1057 (1898).
U. S. Dept. Agr., Exp. Sta. Rec, X-7, p. 650 (1899).
Leaf-spot {LeptosphcBria eustoma (Fr.), Sacc, var. tritici, Garov.).
Leaf-spot (Seploria graminum, Desm.).
See U. S. Dept. Agr., Exp. Sta. Rec, X-s, p. 452 (1899).
Mildew {Erysiphe graminis, DC).
Iowa Bull. 104, pp. 245-248 (July, 1909).
Mold (Cladosporiutn hcrbarum (Pers.), Lk.).
Rust (Black-stem, Puccinia graminis, Pers. and Orange-leaf, P. rubigo-vcra (DC.),
Wint., also P. glumarum (Schum.), Eriks. & Henn.).
Descr. lUus., Ind. Agr. Exp. Sta., Bull. 26 (1889).
Kan. Agr. Exp. Sta., Bull. 38, pp. 1-3 (1893).
Treat, (rec), Idaho Agr. Exp. Sta., Bull. 11, pp. 33-34 (1898).
Cf. U. S. Dept. Agr., Div. Veg. Phys. & Path., Bull. 16 (1899).
Scab {Cladosporium herbarum (Pers.), Lk.).
Scab (Fusarium culniorum (E. F. Sm.), Sacc.=i^. rubiginosuni, Appel & Wollenw.).
Scab {Gibberella Saubenetii (Mont.), Sacc, Stage oi Fusarium roseum, Lk.).
Descr. Illus., Ohio Agr. Exp. Sta., Bull. 97, pp. 40-42 (1898).
Stinking-smut {Tilletia fcetens (Bri. & Cav.), Schrt, T. tritici (Bjerk.), Wint.).
Phytopath. 6, pp. 21-28 (1916).
Loose-smut (Uslilago tritici (Pers.), Jens.).
Descr. Kan. Agr. Exp. Sta., Rep. 2, 1889, pp. 261-267 (1890).
Illus.,
N. Dak. Agr. E.xp. Sta., Bull. 1, pp. 9-20 (1891).

U. S. Dept. Agr., Farm. Bull. 75, pp. 6-8 (1898).
Treat, (pos.), Ohio Agr. Exp. Sta., Bull. 97, pp. 60-61 (1898).
U. S. D^pt. Agr., Farm. Bull. 75, pp. 11-14 (1898).
Willow
(Salix spp.)
Black-spot {Rhytisma salicinum (Pers.), Fr.).

Duggar, p. 209 (1909).
Crown-gall {Pseiidomonas tumefaciens, E. F. Sm. & Towns.).
Duggar, p. 114 (1909).
Decay, or Brown-rot (Poly poms stdphHreus (Bull.), Fr.).
Duggar, p. 457 (1909)-

Powdery Mildew {Undnula salicis (DC), Wint.).
Duggar, p. 230 (1909).
White-rot (Polyporus sqnamosus (Huds.), Fr.)-
Duggar, p. 453 (1909).

Rust {Mdampsora salickaprce (Pers.), Wint.) =M. Jariiiosa (Pers.), Schrot.
Zinnia
{Crassina ckgam (Jacq.) Kze.)
Leaf-spot {Ccrcospora atrkincta, Heald & Wolf).

BIBLIOGRAPHY OF SPECIFIC PLANT DISEASES

That the foregoing list may be made as useful to American students as possible,
a partial bibliography of some of the publications dealing with specific diseases of
our economic plants is herewith given.
Arthur, Joseph C. and Kern, F. D.: North American Species of Peridermium on

Pine. Mycologia, vi: 109-138, May, 1914.
Clinton, G. P.: The Smuts of Illinois Agricultural Plants. Univ. 111. Agric. Exper.
Sta., Bull. 57, March, 1900.
Cook, Mel T.: Potato Diseases in New Jersey, N. J. Agric. Exper. Sta., Circular 33.
Cook, Mel T.: Common Diseases of the Peach, Plum and Cherry. N. J. Agric.
Exper. Sta., Circular 45.
Cook, Mel T.: Common Diseases of Apples, Pears and Quinces. N. J. Agric.
Exper. Sta., Circular 44.
Duggar, B.' M.: Some Important Pear Diseases. Cornell Univ. Agric. Exper.
Sta., Bull. 145, February, 1898.
Duggar, B. M.: Three Important Fungous Diseases of the Sugar Beet. Cornell
Univ. Agric. Exper. Sta., Bull. 163, February, 1899.
Edgerton, C. W.: Some Sugar Cane Diseases. La. Agric. Exper. Sta., Bull.
120, 1910.
Eugerton, W.: Disease of the Fig Tree and Fruit. La. Agric. Exper. Sta.,
C.
March, 1911.
Bull. 126,
Freeman, E. M., and Johnson, E. C: The Loose Smuts of Barley and Wheat.
Freeman, E. M., and Johnson, E. C: The Rusts of Grain in the United States,
.
Freeman, E. M. and Stakman, E. C: The Smuts of Grain Crops. Minn. Agric.
Exper. Sta., Bull. 122, February, 1911.
Harter, L. L.: Sweet Potato Diseases. U. S. Farmers' Bull. 714, March 11, 1916.
Hesler, Lex R., and Whetzl, Herbert H.: Manual of Fruit Diseases, 191 7.
The MacMillan Co.

Johnson, E. C: Timothy Rust in the United States. U. S. Bureau of Plant In-

dustry, Bull. 224, 191 1.
Orton, W. a.: Some Diseases of the Cowpea. U. S. Bureau of Plant Industry,

Bull. 17, 1902.
Orton, W. A.: Tomato Diseases, from Tomato Culture by Will W. Tracy,
1907, Orange Judd Co.
Orton, W. A.: Potato Tuber Diseases. U. S. Farmers' Bull. 544, 1913.
Pool, Venus W.: Some Tomato Fruit Rots during 1907, 1908.
Reed, Howard S. and Crabill, C. H.: Notes on Plant Diseases in Virginia ob-
served in 1913 and 1914. Va. Agric. Exper. Sta., Tech. Bull. 2, April, 1915.
Selby, a. D.: Some Diseases of Orchard and Garden Fruits. Ohio Agric. Exper.
Sta., Bull. 79, 1897.
Selby, A. D.: Prevalent Diseases of Cucumbers, Melons and Tomatoes. Ohio
December, 1897.
Agric. Exper. Sta., Bull. 89,
Shear, C. L. Cranberry Diseases. U. S. Bureau of Plant Industry, Bull. 1 10, 1907.
:
Stevens, F. L.: Fungous Diseases of Apple and Pear. N. C. Agric. Exper. Sta.,
Bull. 206, March, 1910.
Stone, Geo. E.: Tomato Disease. Mass. Agric. Exper. Sta., Bull. 38, June, 191 1.
Taubenhaus, J. J.: Diseases of the Sweet Pea. Del. Agric. Exper. Sta., Bull.
106, November, 19 14.
VON Schrenk, Hermann: Two Diseases of Red Cedar caused by Poly poms
jtmiperinus and P. carneus. U. S. Div. Veg. Physiol. & Pathol., Bull. 21, 1900.
VON Schrenk, Hermann: The Decay of Timber and Methods of Preventing It.
U. Bureau of Plant Industry, Bull. 14, 1902.

S.
VON Schrenk, Hermann, and Spaulding, Perley: The Bitter Rot of Apples.
Wilcox, E. Mead: Diseases of Sweet Potatoes in Alabama. Agric. Exper. Sta.
of the Ala. Polytechnic Institute, Bull. 35, June, 1906.
CHAPTER XXXIV
DETAILED ACCOUNT OF SPECIFIC DISEASES OF PLANTS
This section of the book will be devoted to a consideration of the
specific diseases of plants, and the treatment of the subject has been
made possible by a selection of nearly loo parasitic and non-parasitic
diseases. In this selection, several things have been kept in view, viz.,
the importance of the disease over wide geographic areas, the system-
atic relationship of the fungus in order to connect up the practical and
the systematic parts of the book, because our knowledge of the disease
warrants its inclusion in the descriptive part which follows. As a con-
sideration of the remedial measures used to combat the disease was
omitted largely in the description of plant diseases in general, it is intro-
duced incidentally with the study of specific plant diseases. The chief
reference to such remedial substances and found in one
their use will be
of the appendices in the back of the book, where the manufacture of
sprays and washes and their recommended use may best be made with
the consideration of a spray calendar. A regular spraying program
is now considered a necessity by every successful plant-grower, the
expense of which, treated as insurance, can no more be escaped than the
outlay for cultivation, manures, or pruning. In the control of plant
enemies, including both insect pests and fungous parasites, there are
essential points in practice which may not be evaded or neglected,
namely: To
spray at the correct time (hence the need of a calendar) to
use the proper form and strength of spray (hence the need of formulae)
and to make a thorough covering of the parts sprayed. Hence that
important branch of phytopathology known as therapeutics will be
mentioned incidentally in part III and treated in detail in the latter part
of part IV.
The description of each disease will be given in condensed form pur-
posely, so that the student of plant pathology who wants to know more
about the specific diseases of some particular crop in which his interest
has been aroused will be compelled to study the literature and thus gain
475
access to the most important work which has been done. In this inves-
tigation, the student should write descriptions of the diseased host
plants and parasitic organisms concerned, according to the method out-
lined in part IV, pages 639 to 642, and together with this detailed de-
scription he should compile a bibliography.

Pedagogically it is a mistake to give too full details in a text-book,
because the student learns to depend on the statements in the book
rather than on original observations of his own. The compilation of a
bibliography becomes an important adjunct to all successful phyto-
pathologic work. "Study things, not books" is a truism in this depart-
ment of scientific knowledge, as in other departments of natural science.
The teacher should so guide and stimulate the class of students that each
member of the class will be led to independent study and investigation,
so that they may be able to apply individually the modicum of knowledge
which the strictures of the time allotted to the subject in the college has
permitted them to obtain. Unless this independence of thought and
action is secured, the results of the teaching have not been satisfactory.
It is, therefore, hoped by the writer of this text-book that what has been
included in its pages will be directive and helpful to teacher and student
rather than a work The subject of phytopa-
of encyclopedic value.
thology is would be impossible without the coop-
such a vast one, that it
eration of a large number of specialists to make a work which would be

of encyclopedic value. The design of this text-book has been to give an
outline of the subject, so that the attention of the student may be direc-
ted to the important phases of the subject of phytopathology.
Alfalfa {Mcdicago sativa L.)
Leaf-spot {Pseudopeziza medicaginis (Lib.), Sacc). —The fungus

which causes this widely prevalent disease, where alfalfa is grown,
belongs to a genus in which the apothecium is formed beneath the epi-
dermis and as it grows it breaks through the epidermal covering and
emerges as a shallow, relatively simple structure with asci that contain
eight one-celled spores. It is related to a similar fungus Ps. trifolii,
which attacks the leaves of clovers. It forms small brown, or black,

spots on the upper leaf surface usually. These spots, which are about
2 mm. in diameter, represent the sessile apothecia, which are sprinkled
pretty copiously over the leaf surface in the latter part of summer.
DETAILED ACCOUNT OF SPECIFIC DISEASES OF PLANTS 477
The unicellular spores measure 10 to 14/i in length. No practical

method has been devised for controlling the alfalfa leaf-spot disease.
—
Rust {Uromyces striatus Schrot.). The aecidiaof this rust are found
on Euphorbia cyparissias in Europe and in Great Britain the uredinea
and telia occur on a clover Trifolium minus. In California, it forms
reddish-brown, dusty pustules on the surfaces of alfalfa leaves and
in wet weather it may be destructive to the crop, but in dry weather
it usually disappears. The spots are on close examination seen to be
cinnamon-colored, due to the presence of globose to ellipsoid, faintly
echinulate, yellowish-brown uredospores, which measure 15 to 22//
with a spore wall i and with four to six germ pores each
to 2^ thick,
with a small cap. The telia are darker in color, and the teliospores are
globose to ovate with a minute papilla striated from apex to base with
lines of brown warts and measure 18 to 24 by 15 to 20/x with an epir
spore 1 3^ to 2/i thick. The best way of combating this disease is to
cut and burn badly affected crops. Frequent close mowing is useful
in checking leaf-spot.
Apple {Pyrus malus L.)
Bitter-rot {Glomerella cingulata (Stonem.) S. & V. S.). —This

fungus, which in some text-books is known as G. rufomaculans (Berk.)
Spauld. & von Sch., causes one of the most serious losses in the apple-
growing United States (Fig. 190). It is distributed
districts of the
widely, particularly eastward of the arid portions of the country and
its effects are seen during July and August and later, especially during
warm rainy weather, which produce sultry conditions of the atmos-
phere, when the age of the fruits is such as to render them especially
susceptible. Cold weather acts as a check to the spread of the dis-
ease. The fruit is attacked chiefly, but the branches may also become
diseased.
The disease first appears as a small brown spot beneath the skin of
the apple, which increases gradually in size, keeping nearly a circular
outline with a well-defined margin. The central part of the spot soon
becomes sunken and this is accompanied by the spread of the fungus
throughout the fruit and the formation of pustules. Decay soon sets
in and the products of the decay are invariably bitter. The fruits, if
attacked on the tree, later fall off, but sometimes, they hang on and
become mummified. Two stages in the life history of the fungus have
been discovered. The gleosporial, or imperfect stage, usually develops

on the fruit, while the ascigeral stage is occasionally produced on a
fruit or twig, and in artificial cultures is readily obtained. Early in-
fection of the fruit is probably due to the spores produced in pustules

on the areas which have become cankered through the attack
of stem,
of the bitter-rot Such cankers are sunken areas upon twigs
mycelium.
or limbs, accompanied by a cracking and breaking of the bark over such
regions. The pustules, which accompany the rot of the fruit, are formed
beneath the apple skin as ccmdensed masses of the mycelium known
as stroma and these emerge as a cone-shaped mass of erect hyphae,
which are the conidiophores, which cut off conidiospores that emerge
as a pink waxy strand, later becoming of a gray color. The ovate to
oblong conidiospores, which measure in extreme cases 6 to 40 by 3.5
to 7/1, more usually 12 to 16 by 4 to 6/i, are imbedded in a gelatinous
matrix which dissolves in water setting the spores free. These spores
germinate freely and become septate in doing so. Infection of apple
fruits may be through the uninjured skin, but a slight abrasion facilitates
the entrance of the germ tube of the spore. Berkeley, who first
described this stage, named it Gleosporium Jructigenum and under this

scientific name the disease is frequently quoted.
Clinton discovered the perithecial stage in 1902, and as it is readily
obtained in cultures on any of the ordinary nutrient media its character-
istics are well-known. The perithecia which are developed contain
oblong-clavate asci, 55 to 70 by g/x, which develop eight curved asco-
spores, usually uniform in size, 12 to 22 by 3.5 to 5/1. The pomologist,
who wishes to control the disease, should prune away all cankered limbs
and keep the orchard free of diseased fruits. The spraying of the trees
with Bordeaux or lime-sulphur (3-3-50) has been found efficacious,
and the crop returns from sprayed trees, as contrasted with unsprayed
trees, have abundantly repaid the trouble which the orchardist has
taken in the application of Bordeaux mixture. The first application
of the spray should be in the form of a mist about a month after the
petals have fallen and subsequent applications should be made about
two weeks apart until at least five sprayings have been made.
Black-rot {Spharopsis malorum Berk.). Although the apple is—
one of its host plants, the black rot fungus attacks other pomaceous
trees, producing cankers so that the description of the disease and
fungus, as applied to the apple, will serve with certain modifications for
the other pomaceous trees as well, and this may be said of several of
the other diseases treated of here that the description of a disease as
specifically affecting a certain host, might equally apply to several
other host plants. The
black-rot fungus not only causes a fruit
decay of apples, quinces and pears, but it causes the formation of
canker on the limbs of these trees. The fruit rot is the generally
recognized form of the disease. The disease begins as a small spot
sometimes near the bud end of the fruit and it spreads until the whole
fruit is involved. The apples do not shrink, as in the former disease.
The canker form of the disease on the bark of the trees is accompanied
by either a roughening of the bark in mild forms of the disease, or in
more virulent forms by a destruction of the bark with the formation
of depressed areas about which local swellings of the limbs occur.
The sooty brown, or olivaceous, mycelium penetrates the bark
of the tree, hardly extending into the wood. It soon forms pycnidia
which are erumpent and surrounded by the remnants of the epidermis.
The pycnospores are oblong-elliptic, 22 to 32 by 10 to 141JL, brown in
color, and their size varies with the host plant on which the fungus lives.
Artificial cultures of the fungus have successfully produced spores.
Lime-sulphur solution has been found useful in combating the disease,
but pruning and scraping the trees should not be neglected.
—
Scab {Venturia inequalis (Cke.) Wint.). The scab also appears on
the pear, but mycologists now consider that the scab fungus of the
apple is specifically distinct from that of the pear. Earlier mycologist^
were familiar with the conidial forms fungi, and they
of the two
were placed under the genus Fusicladium, as F. dendriticum and
F. pyrimim, but since the perfect stages have been discovered the
species have been put in the genus Venturia. The perithecial stage
is saprophytic. Scab is found wherever the apple is grown from
Maine to California.
The fungus mainly attacks the fruit and leaves of the apple, but
it has been found on the flowers, flower stalks and twigs. The leaf
spots are more abundant on the lower surface, but sometimes also on
the upper surface, as a velvety, olivaceous, superficial growth, occasion-
ally accompanied by a curling of the leaf. The fruit spots are at first
small and olivaceous, and as the mycelium spreads the epidermis is
killed and the scabby areas arise (Figs. 164 and 165). Nearly all varie-
tes of apple and pear are susceptible, but there is a varietal difference
in this susceptibility.
48o SPECIAL PLANT PATHOLOGY
The hyphae grow beneath the epidermis and between the epidermis
and cuticle spreading slowly. The erect conidiophores, which are
produced, rupture the epidermis, giving the characteristic velvety,
Fig. 164. — Two apples affected with scab {Venliiria inequalis), showing spots,
deformation and reduction in size of the fruit. {After Heald, F. D., Bull. 35 {Sci.
Ser. 14), Univ. of Tex., Nov. 15. 1909.)
Fig. 165. —Two apples affected with scab {Vcnluria inequalis), showing spots,
deformation and reduction in size of the fruit. {After Heald, F. D., Bull. 135 {Set.
Ser. 14), Univ. of Tex., Nov. 15, 1909.)
olivaceous character to the spotted surface, and as the scabby areas

are formed, the epidermis disappears. Conidiospores arise at the tips
of the conidiophores and in concatenation. These spores are ovate,
DETAILED ACCOUNT OF SPECIFIC DISEASES OF PLANTS
truncate at the base and measure 28 to 30ju by 7 to 9/1. According to

Clinton, they do not retain their vitality long. An investigation of
perithecial formation indicates that perithecia begin to form in
October, or even later, and reach maturity in the following April,
when mature ascospores have been
found especially on the under sur-
faces of the leaves. They are im-
bedded in the leaf tissues and are
slightly pyriform in shape, includ-
ing clavate slightly curved asci
measuring 55 to 75// by 6 to 12^1.
Each ascus contains eight two-

celled ascospores, which are olive-
brown in color with the following
dimensions: 11 to 15/iby 5 to 7/x.
They germinate readily in water.

Spraying with lime-sulphur
mixture 32° Beaume, 1-40, before
the time of flowering has been rec-
ommended for Scab, followed by
a second, or even a third spraying
after the petals fall, and at least
two or three weeks after the
second.
Ash (Fraxinus americanus, L.)
Heart-rot (Fomes fraxinophilus

(Pk.) Sacc). —In the Mississippi
Valley, white ash trees of all ages Fig. 166.—An old sporophore of Fomes
are attacked by this bracket (Polyporus) fraxinophilus on white ash.
(After Herynann von Schrenk, Bull. 32,
fungus, which is a tree wound U. S. Bureau of Plant Industry. 1903.)
parasite, entering usually the stub
of a branch, which has been broken off by the wind, or by snow. From the
point of entrance, the mycelium grows into the heartwood of the trunk.
The wood at first turns darker in color, later the disease is marked
by a bleaching of the color in the spring wood of the annual rings, which
turn to a straw color and then become blanched. The whole woody
31
—
Fig. 167. Disease of ash caused by Fames (Polyporus) fraxinophilus. i, Cross-
section of ash wood; 2, of medullary ray; 3, medullary ray, showing later stage of
attack; 4, 5, of wood cells; 6, starch grains from medullary ray cell; 7 diseased
wood; 8, transection from entirely rotted wood. (After von Schrenk, Hermann,
Bull. 32, U. S. Bureau of Plant Industry, 1903)
tissue becomes straw-colored and finally transformed into a loose

spongy mass of fibers, which readily absorbs water (Fig. 167).
The fruiting brackets, or sporophores, make their appearance from
the mycelium at the base of the stubs, or from wounded surfaces,
either alone, or a number together (Fig. 166). The mature sporophore,
according to von Schrenk,^ is nearly triangular in cross-section with a
broad rounded edge, which at first is white, turning gradually darker
until it becomes straw-colored (Fig. 167). The older portions of the
upper surface are dark brown, or black, and are very hard and woody,
its upper surfaces obscurely zoned, pale brown and rust colored. Wound
protection, as outlined in the section on prophylaxis, is an important
method of preventing the white heart-rot from killing white ash trees.
Asparagus (Asparagus officinalis, L.)
—
Rust {Puccinia asparagi DC.) The asparagus rust is well-known,
having been investigated by a number of mycologists in this country,
notably Halsted, Sirrine, Smith and Stone. ^ In Europe the disease is
of little consequence, but in America it threatens the asparagus growing
of our country, spreading rapidly, especially during times when dew is
abundant, for Smith says: "The amount of rust varies directly and
exactly with the amount of dew, and so long as there is little or no dew,
there can be no rust." During dry summers rust is largely absent.
All of the spore forms are found on the stems and twigs of the culti-
vated asparagus and on several wild species of the genus. The uredi-
nia and telia appear also on the leaf-like branches of the plant. The
aecidia appear as long light-green cushion-like patches. They have a
white peridium and are short cylindric, inclosing the orange-colored
aeciospores, which are 15 to iS/z in diameter, and retain their power of
germination for several weeks. Stomatal infection probably is the rule.
Associated with these secia are spermagonia in small, yellow clusters.
Early summer ushers in the red rust (uredo) stage of the disease with
the deep brown sori more or less scattered at first, later becoming con-
fluent. The urediniospores are yellowish-brown, thick-walled with
four germ pores and measure 21 to 24^. The clothing of a person
Won Schrenk, Hermann and Spaulding, Perley: Diseases of Deciduous
Forest Trees. Bull. 149, U. S. Bureau of Plant Industry.
''Smith, Ralph E.: Asparagus and Asparagus Rust in California. Calif. Agric.
Exper. Sta., Bull. 165: 1-95, 1905.
rubbing against the plant may be colored owing to the abundance pro-
duced. Later in the season the black rust stage appears with the forma-
tion of elliptic two-celled teliospores, 30 to dofx by 21 to 28/i, and with
a thickened apex and long pedicels. Infection of asparagus plants in
cultivated fields is, according to Duggar/ through the aeciospores pro-
duced on wild or escaped plants and not directly from the germination
of the teliospores, which remain in or about the soil. Bordeaux mix-
ture, used as a spray alone, has not been very successful. A more
successful treatment has been obtained by adding a resin mixture to
the Bordeaux solution. Sirrine recommends the following: Bordeaux
mixture, 5-5-40 formula, 40 gallons; resin mixture, 2 gallons, diluted
10 gallons. The resin mixture consists of resin 5 pounds; potash lye i
pound; fish oil i pint; and water 5 gallons. Under certain climatic con-
ditions in California it has been found efficient to dust the young tops
with dry powdered sulphur on a dewy morning at the rate of one
and a half sacks of sulphur per acre, followed in a month by a
second application, using two sacks of sulphur per acre.
Banana {Musa sp.)
Bud-rot {Bacillus musce, Rorer). —Bud rots of the banana have

been reported from the greater Antilles (Cuba, Jamaica) from Central
America and Trinidad. The disease in Trinidad has been investigated
by a mycologist from the United States, J. B. Rorer, the mycologist
of the island government, and he has proved that an organism which
he has isolated and named Bacillus musce is the responsible parasite.
However, the bud-rots of the banana are probably due to the same
cause, but the matter has not been investigated satisfactorily outside
of Trinidad. The disease usually appears on the young plants, attack-
ing the young leaves and the core, which become brown. The tissues
disorganize and a putrid rot sets in with the death of the parts
attacked.
March is the month in which the disease usually begins and in
three or four months its destructive effects are seen.
Beet (Beta vulgaris, L.)
Leaf -spot (Cercospora beticola, Sacc). —This disease is distributed

widely in America and Europe and the red garden beet is seldom free
^ DuGGAR, B. M.: Fungous Diseases of Plants, 406.
from The leaf-spots are very small brown with reddish-purple

it.
borders, when they first appear, and later, when about 4 mm. in
diameter, they become ashen gray at the center with the usual margin.
They are scattered over the blade and eventually the leaves blacken
and dry up, and as the lower leaves die, new ones are formed above
until a characteristic elongated crown may be produced. The gray
color of the spots is usually associated with the formation of conidio-
sphores- and conidiospores. The conidiophores are clustered, arise
from a few-celled stroma, and push through the leaf stomata. The
conidiospores are elongated and needle-shaped, multicellular, 75 to
200/1 by 3.5 to 4.SAi, and under moist conditions, the average length may
be exceeded. They germinate readily in ordinary nutrient media
and the submerged mycelium in agar grows as a dense colony oliva-
ceous in color, while the aerial portion is grayish-green. The disease
fortunately can be controlled by the use of Bordeaux mixture (4-4-50),
and as the spores retain their vitality for some time, early spraying
is important and frequent after sprayings.
—
Rust {Uromyces betcB (Pers.), Tub). The beet rust is known only
from California. It is common in Australia and not unusual in
Europe. Klihn thinks that the mycelium may be biennial in the host,
forming aecia throughout the year. The spermogonia are found in
small yellow groups associated with the Eecia, which are white and
saucer-shaped with aecidiospores 17 to 36/i in diameter, filled with
orange-colored contents. The uredinia and telia are irregularly scat-
tered over the leaf surfaces. The urediniospores are obovate, 21 to
24|U by 35M with echinulate and two opposite germ pores. The
walls,
short pedicellate obovate -teliospores are 18 to 24/i by 25 to 32/x,
with an apical germ pore piercing a wall scarcely thicker at the apex.
Cabbage (Brasska oleracea, L.)
Black-rot (= Pseudomonas brassicce. (Pam.), Sm., Bacterium cam-

pestris (Pam.), Sm.) —The cause of the black-rot of cabbage and other
cruciferous plants is a yellow, uni-flagellate microorganism, which causes
a yellowing of the cabbage leaves accompanied by a black stain in the
vascular system, forming a conspicuous black network on a yellowish,
or light-brown, background. The badly diseased leaves are shed, so
that the stem may have a terminal tuft of badly distorted leaves.
A stem section shows a browning of the vascular ring and the vessels
are found occupied by bacteria (Fig. i68). When the cabbage plant
is attacked early in the season, it is killed outright, or else it fails
to form the characteristic head. Infections may take place through

injury of the surface, but the greater part of them are through the
water pores, which exude drops of water, which collect during cool
Fig. i68. — Brown-rot of turnip (Pseudomonas brassica). Cross-section from

middle of turnip root showing small bundle fully occupied by the bacterial organism.
{After Smith, E. F., Bull. 29, U. S. Bureau of Plant Industry, Jan. 17, 1903.)
nights, and in natural infection slugs are responsible carriers of the

organism.
Russell has found that the cauliflower is the most susceptible plant,
while turnips and rutabagas are not very susceptible. Edwards reports
that the Houser cabbage is practically immune to black-rot under field
conditions. The period of incubation is variable. In some cases with
needle punctures, the first signs of the disease appear in seven to
twenty-eight days on leaves and in from nine to thirty-one days on

stems. E. F. Smith obtained with needle punctures first signs of
disease in fourteen to twenty-one days. In a study of the morbid
anatomy of the cabbage, been found that the parasite is
it has
confined for some time to the vascular system and even to particular
leaf traces or bundles, especially to the spiral and reticulated vessels,
which are very often filled with incalculable numbers of this organism.
Later, as the walls of the vessels are destroyed, the organism finds its
way into the parenchyma. Pseudomonas brassicoe is

surrounding
sometimes motile, especially when taken from the plant, and is
examined in a hanging drop of water. Its measurements are 0.7 to
3.0/i by 0.4 to 0.5^. It is often somewhat irregular in shape. The
flagella is several times the length of the cell and arises at or near the
end. The organism is wax-yellow, changing to a dirty yellow-brown
in old cultures.
The treatment of this disease falls principally under the head of
restriction and prevention.
Seasonal variations are found and the
organism thrives well in cool, moist lands. Underdrainage of soils
might prove advantageous in wet seasons. The diseased plants should
not find their way manure heap, but all refuse should be de-
into the
stroyed. As E. F. Smith puts it, "Avoid infected seed, soil and manure;
destroy insect carriers of infection, if the plants are attacked." Crop
rotation is advantageous. Soaking the seed for fifteen minutes in a
solution of mercuric chloride (one tablet to a pint of water) should be
practiced.
Club-root {Plasmodiophora brassiccB, Wor.) This disease, which
has been known for a hundred years, has received a number of
names, such as fingers and toes, Anbury, Hanbury (England), Kohl-
hernie (Germany), maladie digitoire (France) Kapoustnaja Kila
Russia). (The organism causes unsightly and destructive root dis-
ease of cruciferous plants, such as cabbage, Brussels sprouts, turnips,
rutabagas, radishes and certain mustards (Fig. 169). The parasite is
a slime mould (Myxomycetes) named by Woronin {Plasmodiophora

hrassicce).It lives in the parenchymatous cells, often in the vicinity
of the cambium, and an abnormal development of phloem is notice-

able. The infested cells are grouped together into packets and their
contents are at first fluid, then turbid and granular, assuming the
amoeboid form with distinct nuclei. The amoeba are increased by
division, and by a sort of gemmation. The myxamoeba are provided

with several nuclei. The formation of spores soon begins by the suc-
cessive simultaneous divisions of the myxamoebae, so that each nucleus
and surrounding mass of cytoplasm is differentiated, as a spore by the
formation of a spore wall about them. The diseased cells are crammed
full of such spores, which escape
only when the root disintegrates.
The liberated spores will germi-
nate in water in from four to
twenty-four hours and later the
parasite gains entrance to the
roots of the cabbage plant. The
•organism causes an excessive
formation of new cells so that a
gall, or canker results.
In order to check the organ-
isms, soils have been treated with
lime, sulphur and other fungi-
cides. Liming, using two tons of
c|uicklime to the acre eighteen
months before planting, has been
found the most reliable with the
destruction of the refuse of pre-
vious crops by burning.
Carnation (Diani/ms
caryophyllus, L.)
Fig. 169. — Cabbage roots showing club- Alterniose {Alter naria

root caused by a parasitic slime mould,
Plasmodiophora
Microbiology.
brassicce.
Second
{From Marshall, diantki, Stev. & Hall).
edition, p. 609, after ^ >•
Through
.
— •-!-.•
y ^ ^ Connecticut,
t-> 1
Pennsylvania, Dis-
Woronin.)
trict of Columbia and North
Carolina this disease of the cultivated carnation has been recently quite
troublesome. The leaves and stems, especially at the nodes, are dis-
colored with spots of ashen whiteness with a central black fungous
growth. The spot is dry, shrunken and thinner than the surrounding
healthy parts of the leaf, and is somewhat elongated
either circular, or
in Hne with the long axis of the leaf. The nodal spots involve the leaf
bases as well,and the mycelium finally grows into the stem killing its
tissuewhich becomes soft and broken down (Fig. 170). The variety
known as Mrs. Thomas W. Lawson is especially susceptible.
Rust (Uromyces caryophyllinus (Schrank). Wint. This disease —
was practically unknown in the United States prior to 1890, but now it
Fig. 170.— Carnation alternariose {Allernaria dianlhi). i, Branched, septile my-

celium; 2, hyphas below surface of stroma; 3, spore formation; 4, compound spores,
5, young clustered hyphse; 6, older cluster. {After Stevens, F. L., and Hall, J. C;
Bot. Gas., 47: 409-413, May, 1909.)
is prevalent wherever the carnation grown commercially. The dif-

is
ferent varieties of cultivated carnations differ to amarked degree in

susceptibility. Enchantress and Lawson have a high degree of resist-
ance to rust, while Scott and Jubilee are very susceptible.
The fungus is by its urediniospores, which are

largely propagated
ellipsoid to spheric inform and measure 24-35^1 by 21-26/i. The
spore wall is thick and spinulose. The teliospores resemble in form
the urediniospores and measure 20-35JU by 18-25/x. Their walls are
chestnut-brown and uniformly thickened with terminal germ pores
and are papillate. As the adult plants may be infected, the disease
may spread rapidly during the growing season.

The disease can be controlled undoubtedly by growing rust-resistant
varieties of carnations. The leaves should be kept away from the
moist by simple V-shaped wire mesh supports and lastly fungi-
soil
cides, such as a solution of copper sulphate (i pound copper sulphate

to 20 gallons of water), might be used with success, Duggar also rec-
ommends the use of potassium sulphide i ounce to a gallon of water.
Sub-irrigation has been practised.
Cacao (Theobroma cacao, L.)
—
Brown-rot (Thyridaria tarda, Bancroft). A number of different
organisms have been thought at different times to cause the brown rot
of the chocolate pods, but Bancroft in 191 1, an authority on the sub-
ject, ascribed the disease to the above-named fungus. Circular brown
patches appear on the chocolate fruits along the grooves that seam
the surface. The disease spreads rapidly and the fruit falls in from six
to ten days from the time that it is first infected. When the spots are
2 cm. in diameter, their center becomes marked by wounds in which
a brownish-gray mycelium appear. Wounded fruits are especially
open to infection through the abraded surface and the seeds, or beans,
are sometimes involved and are destroyed completely. The disease is
widely spread in the eastern and western tropics (in Jamaica, Santo
Domingo and the Philippines). It may be controlled to some extent
by burning all diseased fruits, busks and prunings.
Pink Disease {Corticium lilaco-fuscum, Berk & Curt.). The genus —
Corticium belongs to the family of Thelephorace^, which includes
the smothering fungi of the genus Thelephora. The leathery hymeno-
phore of Corticium is membranous, fleshy, waxy with clavate basidia
with four sterigmata. The basidiospores of our cacao fungus are
sessile on the basidia. It attacks the younger branches of the chocolate
tree covering them with a pinkish incrustation, which spreads over
1
the bark and into the bark crevices, causing the bark to crack and
peel. Later a new bark forms under the old. The new bark is not
sufficiently resistant to the attacks of species of Diplodia and Neclria,
so that these fungi may enter and complete the work of destruction.
Corticium lilaco-fuscum grows more rapidly in damp, shady places,
and it usually refuses to grow in sunny places, hence opening up the
growth is beneficial.
Cherry (Prunus spp.)
Leaf-curl (Exoascus ccrasi (Fckl.), Sadeb.).^ —This fungus produces

witches' brooms out and when the leaves on
of the twigs of the cherry,
affected twigs are parasitized, they become somewhat reddish and
curled. The asci develop on the leaves and measure according to
Sadebeck, 35 to 50)uby 7 to lo/x, or in specimens studied by Atkinson,
25 to 2)2)1^ by 6 to 9/1. The asci are naked and arranged in rows over the
leaf surface. Spraying, if done at all should be done when the buds
begin to develop in the Spring, and again when the asci are mature
and ready to discharge their spores.
Powdery Mildew (Podosphcera oxyacanthcB (DC), deBy). This —
disease, although found on a number of other rosaceous plants,
such as plums and hawthorns and the like, is especially destructive to
apples and cherries. The leaves become mildewed with large spots
of white mycelium from which arise the perithecia, which are 65 to 90)u
in diameter surrounded by the dichotomously branched hyphal append-
ages four to thirty in number, which are usually five times as long as
the diameter of the perithecium. A single ascus usually contains 8
ascospores. recommended to spray with lime sulphur
It is (1-40)
or dust with powdered sulphur in combating this disease.
Chestnut (Castanea dentata (Marsh.) Borkh.)
Blight {Endothia parasitica (Murrill), Anderson).—When the chest-

nut blight fungus was first described by Murrill he called it Diaporthe
parasitica, but by the studies of Anderson and others it has been trans-
ferred to the genus Endothia, where it seems rightly to belong,^ On
account of its virulency and its rapid spread through the chestnut
1 Shear, C. L., Stevens, Neil E., and Tiller, R. J.: Endothia parasitica
and Related Species. Bull. 380, U. S. Dept. Agric.
Fig. 171. — Canker lesion that nearly surrounds the chestnut branch, sunken on
one side and enlarged on the other. (,Photo by Wm. Ciirrie, Bull. 5, Penna. Chestnut
Tree Blight Com., 1913.)
forests of the eastern United States, it has been the subject of much
legislation and also a copious bibliography has been formed by the
appearance of papers on its parasitism, life history and the remedial
measures to be taken to combat it. The chestnut blight funguswas
Fig. 172. —Perithecial pustules of chestnut blight fungus (Endothia parasitica)

in the crevices of bark of a fallen chestnut tree. (Photo by Wm, Currie, Bull. 5,
Penna. Chestnut Tree Blight Com., 1913.)
discovered by Merkel in 1904 on American Chestnut trees {Castanea

dentata) in the New York Zoological Park. It was studied by Murrill
during 1906 by pure culture and by inoculation on healthy chestnut
trees, and an account was published of the fungus as a new species in
Torreya (6 186-189) '^^ 1906.
:
The rapidity of spread has been phenomenal, and the completeness

of destruction is without parallel in the annals of plant pathology. It
is now found from New Hampshire to Albemarle County, Virginia, in
the South. Summer is the best time to study the symptoms of the
disease, which are manifested in the brown shriveled leaves, which
may be seen at a distance. The dead leaves hang on the tree over
winter, and if on the blighted branches, the girdling is completed while
the burs are maturing. Burs smaller than usual, and unopened, re-
FiG. 173.— Chestnut blight pustules producing gelatinous threads with summer
spores. (After pictorial card issued by Penna. Chestnut Tree Blight Com., 1912.)
main attached to the tree through the winter months and well into the
next spring. If, however, the girdling takes place after the leaves and
burs are shed and before the leaves open in the spring, the leaves do
not attain their full size, but are pale and distorted and this is a com-
mon symptom during May and June. Dead limbs without attached
leaves, or burs, are often indications of the canker disease. Water
sprouts, or suckers, may develop just below the cankered regions of
the branches or stem and thick clumps of suckers on the trunk and
branches, or at the base of the tree, are evidences that the trees are
attacked by the chestnut blight fungus.
The cankers on smooth bark are especially marked, and with a
reddish-brown color in contrast with the healthy bark can be seen for a
considerable distance (Fig. 171). As sunken, or swollen diseased areas
of the bark, they occur on branches and generally the cankers
of all sizes
up and down the stem (Fig. 171).
are ellipsoidal with the long axis
The cankered areas of bark become covered with numerous small
pimples (Fig. 172) from which emerge in wet weather long twisted
Fig. 174. —Chestnut blight fungus, iiMffoi/u'o parasitica. A, Pustules on bark;

B, escape of pycnospores as gelatinous cords; C, D, magnified views of the cord-like
masses of pycnospores. {From Gager after Murrill.)
yellow horns of a gelatinous nature (Figs. 173 and 174). As the

canker ages the bark splits and cracks, and in a year or two it peels
off from the tree leaving the wood exposed to the weather (Fig. 127).
The mycelium forms mats in the bark and cambium
thick, fan-Uke
of the tree and it spreads both longitudinally and circumferentially
(Fig. 175) until, having completed its growth around the stem, or
branch, and killed the cambium and bark, the part of the tree above
the girdled portion succumbs and the next year leafless branches
show the irreparable damage done to the tree by the blight fungus
(Fig. 127).
Fig. 175.— Fan-shaped mycelium of chestnut blight fungus (Endothia parasitica)

from rough bark of a chestnut tree. (Photo by E. T. Kirk, after Anderson, Bull. 5,
Chestnut Tree Blight Com., IQIS-)
Morphology. — On smooth bark, especially in summer, the outer cork

layer is raised into numerous little blisters, with slender, yellow, waxy
twisted horns emerging from a pore in their apices. A section across
each blister reveals a somewhat globose pycnidium surrounded by a
scanty loose mass of whitish, or yellowish hyphae, which merge with
the tangled hyphae that make up the pycnidial wall. The conidio-
phores arise inside the pycnidium, as a dense brush-like fungi and pro-
ject into the fruit cavity (Figs. 174 and 176). They range in length
from 20 to 40/1. From these conidiophores, spores (pycnospores) are
abstricted, and as the cavity is filled with the hyphal stalks, the pyc-
nospores are forced out at an opening in the top of the pycnidium in
the form of a twisted slimy cord (Figs. 173 and 174). The smooth
hyaline pycnospores are held together by a sticky material and they
measure 1.28 by 3.56/i in size, and are oblong cylindric with rounded
ends sometimes slightly curved. Heald and Gardner^ find that the
pycnospores are to a considerable degree resistant to desiccation in
soil in the field and that a large number may retain their viability
during a period of 2 to 13 days of dry weather (Fig. 177). They
found that with indoor desiccation a large number of spores survived
two months and that in 5 out of 12 samples not all of the spores had
succumbed after three months of drying. The longevity limit varies
from 54 to 119 days, the average being 81 days. Studhalter and Rug-
gles^ by experimental methods obtained some interesting results as to
insects as carriers of the chestnut blight fungus. Tests were made
with twenty-one ants in certain laboratory and insectary experiments
in which they had been permitted to run over chestnut bark bearing
1 Heald, F. D. and Gardner, M. W.: Longevity of Pycnospores of the Chestnut

Blight Fungus in Journal Agricultural Research II: 67-75, April 15, 1914.
Soil.
Additional facts in the life history of the chestnut blight fungus are presented
in the following: Heald, F. D., and Walton, R. C: The Expulsion of the
Ascospores from the Perithecia of Endothia Parasitica (Murr.), Amer. Jour.
Bot., 1:449-521, Dec, 1914; Heald, F. D., and Studhalter, R. A.: Seasonal
Duration of Ascospore Expulsion of Endothia parasitica. Amer. Journ. Bot.,
2: 429-448, Nov., 1915; Ibid., The Effect of Continual Desiccation on the Expul-
sion of Ascospores of Endothia Parasitica. Mycologia, 7: 126-130; Ibid., Lon-
gevity of Pycnospores and Ascospores of Endothia Parasitica under Artificial
Conditions. Phytopath, 5:35-44; Stevens, Neil E. Some Factors Influencing
:
the Prevalence of Endothiagyrosa. Bull. Ton. Bot. Club, 44: 127-144, Mch., 191 7.
2 Studhalter, R. A. and Ruggles, A. G.: Penna. Dept. of Forestry. Bull. 12,
April, 1915.
32
spore horns or active perithecial pustules of Endothia parasitica. They

found that five of the twenty-one ants were carrying spores. Tests
with other insects demonstrated that they were carrying spores. The
number of viable spores carried varied from 74 to 336.960 per insect,
and the last number was obtained on Leptostylus Macula, one of the
beetles, which feeds on the pustules of the blight fungus. During
these experiments, it was proved that the spores of Endothia parasitica
were easily shaken from the body of the beetle during its own move-
ments. Heald and Studhalter^ undertook to determine whether birds
carried the spores. They found on
birds shot on blighted chestnut
and wings of each bird were scrubbed
trees after the bill, head, tail, feet
with a brush and poured plates were made from the wash water, which
was retained and centrifuged for its sediment, that in the case of the
36 birds tested, 19 were found to be carrying the spores of the chestnut-
blight fungus. The viable spores carried by two downy woodpeckers
numbered 757,074 and 642,341 respectively, while a brown creeper
carried 254,019, and that the highest positive results were obtained
from birds shot two to four days after a period of considerable rain-
fall. Analyses of spore traps at West Chester and Martic Forge^
showed that viable pycnospores of the chestnut blight fungus were
washed down the trees in enormous numbers during every winter
rain.
The mature stromata on older cankers have numerous projecting
on the surface. The black speck at the tip of each papilla is
papillae
the opening of a perithecium, which is a bottle-shaped depression with
a long neck-like, black canal opening at the surface. There are com-
monly fifteen to thirty perithecia in a stroma. The mature perithecia
(Fig. 176) measure about 350 to 400^1 in diameter, and are mostly
spherical. The neck is usually four to six times the diameter of the
perithecium and its black wall is composed of densely interwoven,
septate, heavy-walled hyphae running parallel with the long axis of
the neck. The asci are clavate, or oblong, and contain eight ascospores
imbedded in an epiplasm. The ascospores are two celled and measure
1 Heald, F. D. and Stud halter, R. A.: Birds as Carriers of the Chestnut Blight
Fungus. Journal of Agricultural Research II: 405-422, Sept. 21, 1914.
2 Heald, F. D. and Gardner, M. W.: The Relative Prevalence of Pycnospores
and Ascospores of the Chestnut Blight Fungus during the Winter. Phytopathology
3: 296-305, December, 1913.
—
4.5 to 8.6/i in size (Fig. 177). The walls are thicker than those of the
pycnospores. Expulsion of the ascospores is dependent upon tempera-
-^f
•^'^^:!d^>^
Fig. 176. A, Vertical section of a pycnidial pustule. The filaments lining the
cavity produce the spores that ooze out as "spore-horns;" B, vertical section of a
perithecial pustule.'
Several of the perithecia are cut so as to show the fuUlengths
of the necks in the chestnut blight fungus (Endoihia parasitica). (After Heald,
F. D., Bull. 5, Chestnut Tree Blight Com., 1913.)
ture, as well as moisture. There was no expulsion of ascospores under

field conditions from late November until the rain of March 21, when
temperature conditions were favorable. Ascospores were not expelled

during the warm winter rains, but during the summer rains ascospores
Fig. 177. — Spore-sacs or asci with eight two-celled ascospores of chestnut blight
fungus (Endoihia parasitica). Below diagram showing relative size of pycnospores
(left) and ascospores (right). (After Heald, F. D., Bull. 5, Chestnut Tree Blight
Cojn., 1913.)
are forcibly expelled in large numbers from the perithecia during and
after each warm rain in case the amount of rain is sufficient to soak up
Fig. 178.— Photograph showing svtccessive stages in the germination of asco-

spores and pycnospores of the chestnut blight fungus {Endolhia parasitica). Left,
ascospore series from 8 to 22 hours at hourly intervals; right, pycnospore series from
8 to 22 hours, taken every two hours. {After photo by Wm, Currie, Bull. 5, Penna.
Chestnut Tree Blight Com., 1913.)
the pustules.^ and wind transport

All of the experiments point to air
of the ascospores, as one of the very important methods of dissemina-
tion. Infection is by means of wounds produced mechanically, as by
insects and other animals (Fig. 178). It is still to be demonstrated that
the parasite can enter without visible breaks in the bark.^ In the
control of this disease inspection of nursery stock should be made
and the use of gas tar following removal of diseased branches.
Leaf Mildew {Phyllactinia corylea (Pers.), Korst). The under leaf —
surfaces of the chestnut are marked frequently by irregular patches
of mycelium, which constitute the mildew fungus (Fig. 53). Typical
haustoria are absent, but there are special setalike branches which
penetrate the leaf tissues. The subglobose perithecium is large and
is garnished with rigid needle-like appendages with a swollen base
(Fig. 53). There are many included asci usually containing two
spores, occasionally three. It is a fungus of wide geographic distribu-
tion throughout the temperate regions of the world.
Clover (Trifolium spp.)
Rust, Uromyces trifolii (Hedw.), Liv.-—The common clovers of our

cultivated fields, such as the red clover, alsike clover, white clover, and
crimson clover, are attacked by this rust, which causes serious disease
conditions (Fig. 70, E and F). The prevalence of the disease varies
greatly with the season. The clover rust fungus is autoecious, all of
the stages being found on the same host plant. All of the stages
occur on the white clover (T. repens). In general the spermagonia
and aecia are not met with on the red clover, the host upon which the
other stages are perhaps more frequent. The mycelium is local in its
occurrence in the plant, from and spermagonia arise in the early
it secia
spring, or at almost any time during an open winter. They occur on

the under leaf surfaces and on the leaf stalk. The aeciospores are 14
to 23JU in diameter and germinate readily in water.
Heald, F. D., Gardner, M. W. and Studthalter, R. A.: Air and Wind

Dissemination of Ascospores of the Chestnut Blight Fungus. Journal of Agricul-
tural Research iii: 493-526, March 25, 1916.
For numerous other details consult Anderson, P. J. and Rankin, W. H.:
'^
Endothia Canker of Chestnut. Bull. 347, Cornell University Agricultural Experi-

ment Station, June, 1914.
The urediniospores are about 22-26/x by 18-20/x, and repeated

crops of these may be produced. The teliospores are formed in sori
with the urediniospores, as the season advances. They are one-
celled, thick walled and measure 20-35/x by 15-22^- The teliospores
germinate in the ordinary way by the formation of a four-celled
basidium each producing a basidiospore. No satisfactory method

of controlling clover rust is known.
Coffee (Cojfea arabica, L.)
Leaf-spot {Cercospora caffeicola, B. —

& €.). The leaves and fruits
of coffee plants in the Dutch East Indies, Mexico, Cuba, Jamaica,
Trinidad and Brazil are attacked by the leaf-spot fungus, which causes
large blotches at first visible only on the upper leaf surface. The spots
are dark brown at first, later becoming grayish above and clear below.
The center of these blotches die and here the spores are borne. The
disease causes the leaves to fall, thus reducing the vigor of the plant
and preventing the proper maturing of the coffee berries. Infected
berries fall before ripening.
Rust {Hemileia vastatrix, Berkeley & Broome) — The coffee rust
.^
is
widely spread through the coffee-growing regions of the old world,

and it has been reported from the American tropics, but there is some
uncertainty about reports. It is the most destructive disease of the

coffee plant and American coffee growers should be on the lookout
for it.
Orange-red spots appear on the leaves, which finally wither and drop,
and frequently parts or whole plants die, especially during the rainy
season, when the red spots increase in number. The spots appear as
shghtly transparent discolorations, which are not easily observed until
the leaf is held up to the Hght. An older spot is yellow in color and then
a bright orange color. They vary in size, but are usually circular in
outhne, and increase in number during June and July, when the disease
reaches its culmination, if the weather conditions are favorable. The
spores are produced in great abundance in the orange-red spots and on
being set free are carried by the wind and insects to other coffee plants
on the leaves of which they germinate sending a germ-tube into the
leaf through the stomata. The urediniospores 35 to 40/x by 25 to 28ju
are single, usually egg-shaped, provided with a papilla and without
germ-pores. The teliospores are unicellular. As a remedial measure

the use of tobacco water or Bordeaux mixture isrecommended.
Corn (Zea mays, L.)
Dry-rot {Diplodia zees (Schw.), Lev.). —The dry rot fungus attacks
the dry ears of corn soon after silking and does not usually manifest
itself until husking time, when the kernels are found to be covered with
a whitish mycelial growth, which dips down between the individual
grains of corn. The become shrunken, loosely
grains so attacked
attached to the cob, lighter in weight, darker in color, and more brittle
than the healthy grains. Pycnidia may be found imbedded in the
mycelium, especially between the kernels. In the open field, these
pycnidia may be formed in such numbers as to impart a black color
to the grains of corn. Of course the feeding value of the corn is gone
and some physicians even ascribe pellagra to the use of such moldy
corn. When the fungus attacks the stalks, it forms small dark specks
under the epidermis near the nodes and even on three-year-old stalks
pycnidia have been found. Infection takes place through the roots
and the fungus which enters in this way Ear
finally reaches the stem.
infection may also occur by wind-blown spores which

through the silk
come from old diseased stalks left in the field, so that by destroying
the corn trash the disease can be controlled to some extent. Rotation
of crops is probably more efficacious.
—
Smut {Ustilago zem (Beckm.), linger). The smut boils of Indian
corn, or maize, are found not only on the ears as with most smuts, but
also on the husks, on the tassels of male flowers, on the leaves, and even
on the stem (Figs. 179 and 180). The attack first begins on any young
and tender part of the plant. If the leaves are the part attacked, they
assume a pale yellow hue and are puckered with smaller, or larger
bladder-like swellings. The swellings are made up of masses of the
hyphae of the smut fungus and their surface is covered with a smooth
skin-like covering. Later the hyphae divide up into innumerable
rounded which develop into the smut spores, or chlamydospores.
cells,
Finally, the silvery-white skin having beenmore and more stretched

bursts, and the black chlamydospores are set free, as a powdery
mass. The echinulate chlamydospores measures 8 to 12/i, and they
readily germinate in manure-water giving rise to a four-celled basidium,
each cell ofwhich produces a basidiospore. Infection of the nascent

tissue at any part of the growing corn plant is accomplished by the
Fig. 179. —
Smut boil of UsUlago zea on ear of corn, developed from one infected
kernel. {After Jackson, F. S., Bull. 83, Del. Coll. Agric. Exper. Slat., December,
1908.)
basidiospores and not by the chlamydospores (Fig. 181). Wet weather

is essentia] for the growth of the corn and the smut also.
5o6 SPECIAL PLANT PATHOLOGY
The disease may be controlled by removing the smutted plants

from the field and destroying them and also by a rotation of crops.
Fig. i8o. — Corn smut on tassels of sweet corn. (After Jackson, F. S., Bull. 83, Del.
Coll. Agric. Exper. Slat., December, 1908.)
As the fungus may infect the adult plant, the treatment of the seed
corn with fungicides has been unsuccessful. Rotation of crops also
assists in keeping smut in check.
DETAILED ACCOUNT OF SPJOCli'lC DISEASES OF PLANTS 507
Wilt (Fseudomouas SkwaHl, Smith). —This is a specific communi-

cable disease of sweet corn and other races of maize, caused by a yellow,
polar-flagellate organism discovered in 1895 by F. C. Stewart. The
disease has been found on Long Island, in New Jersey, Washington,
D. C, Maryland, Michigan, Virginia and West Virginia. One of the
first signs of the disease in well-grown plants is the whitening (drying
—
Fig. 181. Germination of the chlamydospores of corn smut (Ustilago zece); i.
Various stages in germination from corn 3 days after being placed in water; 2, spores
germinated in contact with air; 3, several days after spores were placed in 1/20 per
cent, acetic acid, formation of infection threads, a. Spores; h, propiycelia; c, basidio-
spores; d, infection threads; e, detached pieces of mycelia. {After Bull. 57, Univ.
III. Agric. Exper. Stat., March, 1900.)
out) of the male inflorescence. The leaves then dry out and the plant
is dwarfed, later the stem dries. If the leaves or the stem be chosen and
broken across, sHmy yellow contents ooze out. A cross-section of
the stem shows that the organism fills the vessels of the host plant and
the wilting is due to the stoppage of the water suppHes by the trachei'd
plugging.
The greatest pains should be taken to secure only sound seed corn,
but in the present indififerent state of the seed-trade, even the best
should be treated with mercuric chloride before planting. On fields
subject to the disease, only resistant varieties should be planted.

Manure containing corn stalks from diseased fields, or gathered from
animals pastured in such fields, should never be used on land designed
for corn.^
Cotton (Gossypium sp.)
Boll Anthracnose (Glomerella gossypii (Southw.) Edg.) (= Colleto-

trichum gossypii, Southw.),- — The same fungus causes an anthracnose
of stem and boll of the cotton plant, especially in the Gulf states. The
disease is more important when it attacks the boll, or the seedlings.
The ^bolls lose their green color and become dull red, or bronzed.
If the boll is nearly- mature when attacked, it may mature and
open in the usual manner, but if attacked early, it may cause a prema-
ture dying of the carpels and an unequal growth of the boll, which is
liable to crack open and expose the immature lint to the action of the
weather. The first evidence of the disease is a minute reddish spot,
which later becomes black in the center and depressed with a reddish
border, and these spots may run together.
Two types of conidiophores break out from the stroma within the
tissues. Some of the conidiophores are hyaline and abstrict conidio-
spores that measure 4.5 to 7^1 by 15 to 20/x, while other conidiophores in
the form of setae arise from the dark colored cells of the stroma. The
setse are clusteredand bear ovate, basally pointed spores. Spores and
setae together form an acervulus. The spores germinate readily and
produce a myceUum which grows vigorously in culture, is hyahne,
flexuous and abundantly septate and it may give rise to appressoria.
Proper remedial measures have not been discovered, and a field of
experimentation is opened up along these lines. Use resistant
varieties.
Rust (Uredo gossypii, Lager.). —This is the uredo stage of Kuehneola
gossypii (Lagerh.) Arth. which occurs on the cotton plant in Cuba,
Puerto Rico, Florida and Guiana, ^cia are wanting in the life cycle,
1 Smith, Erwin F. : Bacteria in Relation to Plant Diseases, Volume III: 89-150,

1914,^ where full details of the experimental study of the disease and the causal
organism will be found.
while the other spore forms are represented by urediniospores and

teliospores. All parts of the green cotton plant may be rusted,
spreading to the new leaves as they are formed. Small rounded, or
angular, purplish-brown spots appear on the upper leaf surface and the
urediniospores are borne in pustules just beneath the epidermis on the
under which finally ruptures and sets them free. The
leaf surface,
grown in the Southern United States are partially
varieties of cotton
immune, while the tropic varieties are more susceptible. It is rec-
ommended that the cotton grower destroys all rubbish in his fields
and adopts a system of field culture in which only vigorous plants will
be obtained.
Cranberry {Vaccinium macrocarpon, Ait.)
Gall (Synchytrium vaccina, —

Thomas) (Fig. 230). The fungus which
causes cranberry gall is a very much reduced phycomycetous one, which
attacks the young stems and leaves, as well as flowers and fruit of the
cranberry. It also lives on other ericaceous plants. The galls are
small in size, reddish in color and are produced in great numbers on the
parts affected. The fungous body is much reduced, consisting of a
single cell which becomes a zoosporangium. The presence of this
parasitic cell in the tissues of the host is to produce a small gall. Later
the zoosporangium develops a mass of swarm spores, or zoospores,
which escape into the water. Infection, therefore, probably takes
place when water is abundant.
Scald {Guignardia vaccinii Shear). ^ —The scald fungus (Figs. 182
and 183) may attack the very young fruit and even the flowers of the
cranberry and annually does considerable damage to the growing crop,
as the annual loss has been estimated at $200,000. The pycnidia
are usually found upon such parts. The berries are characterized by
watery spots, which may remain small under certain conditions, while
under others it spreads quickly, often concentrically until the whole
berry becomes soft. The leaves are also spotted with irregular brown
spots within which the pycnidia are found.
The pycnidial stage is a characteristic Phoma, or Phyllosticta,
measuring 100 to i20)u in diameter. These are scattered over the
affected surface and abundant hyaline, obovoid pycnospores are formed,
1 Shear, C. L.: Cranberry Diseases. U. S. Bureau of Plant Industry. Bull,
no: 1-64, 1907.
5IO SPECIAL PLANT PATHOLOGY
Fig 182. — Cranberryscald {Guignardia vaccinii Shear). {After Shear; Bull, no,
U. S. Bureau Plant Industry, pi. i, 1907.)
DETAILED ACCOUNT OF SPECIFIC DISEASES OF PLANTS 51I
Fig. 183. — Details of cranberry scald fungus (Guignardia vaccinii). i, A cran-

berry leaf, showing pycnidia of Guignardia vaccinii thickly scattered over the under
surface; a, a cranberry blossom blasted by Guignardia vaccinii, showing pycnidia on
calyx, corolla, and pedicel; b, a blasted fruit, showing pycnidia. 2, Avertical section
of a single pycnidium of Guignardia vaccinii from a cranberry leaf, showing pycno-
spores in various stages of development. 3. An immature pycnospore of the same
fungus, showing the partially formed appendage; a, the same, showing a little later
stage of development; h and c, fully developed pycnospores and appendages. 4, 5,
6, 7, 8, and 9, Various stages in the germination and growth of pycnospores oiGuig-
nardia vaccinii grown in weak sugar solution; 4, 5, 6, and 7, 72 hours after sowing;
8 and 9, 86 hours after sowing. 10,A vertical section of a perithecium of Guignardia
vaccinii, showing asci, from a cranberry leaf collected in New Jersey. 11, Three
asci, with ascospores showing variations in length of the stipe and the arrangement
of the spores; a and b, from perithecia on a leaf; c, from a pure culture. 12, A fresh,
which measure 10.5 to 13. 5m by 5 to 6ju. The ascigeral stage is less com-
mon. The perithecium has a rather dense wall inclosing a number of
clavate asci, which are 60 to 80^1 long (Fig. 183). The ascospores are
hyaline, elliptic to sub-rhomboidal in form with granular contents.
The fungus has been grown successfully in artificial culture media, but
after a few generations, it seems to lose in vitality.
Preventative measures consist in an occasional renovation of the
bag and in the proper regulation of the water supply. Spraying at
least six times with Bordeaux mixture (5-5-50) is used with success;
especially, adhesive substances (4 pounds resin
if fish oil soap) are
added to the mixture.
Grape {Vitis spp.)
Black-rot (Guignardia Bidwellii (Ell.) V. & R.). — Wherever the

grape is grown this American fungus is a constant menace to the suc-
cessful prosecution of the industry. It attacks not only the fruits, but
also the leaves, fruit pedicels and stems. The disease, which mostis
important on the berries (.Fig. 184), begins as a small circular brown spot
which enlarges until it is 5 to 10 mm. in diameter, when the center of the
spot will be found to show a few black pimples which are the openings
of the pycnidia, which have now appeared beneath the skin. The
spots become darker in color and spread until more than one-half of
the fruit surface is involved, when the fruit begins to lose its spheric
contour and to shrivel, persistently hanging on the vine sometimes
throughout the season. Nearly all of the dark colored grapes are
susceptible, such as the universally grown Concord, while some light
colored varieties are more resistant. The Scuppernong is apparently
entirely resistant.
As with many which attack our cultivated plants, the
of the fungi
different stages were known before the complete
life cycles were de-
termined and therefore, these stages received scientific names, which

are relegated to synonymy, when the life history becomes known
mature ascospore, showing the usual condition, in which the protoplasm is very
coarsely granular. 13, An old ascospore from a dried specimen, having its contents
homogeneous. 14, a, A portion of the coarse brown mycelium from the interior of
a scalded berry, from which a culture was made December 23, producing pycnidia
and ascogenous perithecia of Guignardia vaccinii; b, a portion of younger, lighter
colored hyph^ from the same berry. (After Shear, C. L., Bull, no, U. S. Bureau of
Plant Industry, 1907.)
— —
thoroughly. So it has been with tlie black-rot fungus. The pycnidial

stage on the grape leaves (Fig. 185) was called PhyUosticla lahrusca,
while on the fruit it was called Phoma uvicola. These have been
determined to be merely stages of one and the same fungus, Guignardia
Bidwellii. The mycelium of the black-rot fungus is never abundant in
the outer portions of the berries where Here a stromatic
it is found.
mass of hyphse arises beneath the grape skin and develop the pycnidia,
which are broadly elliptic, thick-walled and
beakless depressions from the inner walls of
which the pycnidiophores arise which abstrict
off the ovate to elliptic pycnidiospores (pycno-
spores) 8 to 1 0/1 by 7 to S^t. These are pushed
out in twisted masses and can germinate im-
mediately.
Spermagonia-like pycnidia of smaller size
are also found. These produce filiform con-
idiophores, which cut off minute, slightly
curved microconidia. The ascigeral' stage,
discovered in 1880, may be had on fruit,
which has been covered with grass and leaves

in the dried up state. The perithecia are
globose and bear broadly clavate asci con-
taining eight unicellular ascospores, measur-
ing 12 to i7Aiby 4.5 to 5M.
The black-rot grape disease can be con-
trolled by Bordeaux mixture (4-4-50). The Fig. 184. Black-rot
fungus, Guignardia Bidwellii,
first application should be made in the spring, attacking green grapes. Cold
Spring Harbor, L. I., July
just
'
as the buds begin
c to swell, followed by
J >
20, 1915.
a second spraying, as the buds unfold. Sub-
sequent sprayings, always before rain storms, to the number of five
or six, should be made two weeks apart during the season. After
July 20 use' 4-2-50 Bordeaux, or ammoniacal copper carbonate.
Downy Mildew (Flasmopora viticola (B. & C.) Berl. & DeTon).
The consensus of opinion among mycologists is that the downy mildew
fungus American origin, and it is now widely spread in Europe and
is of
eastern North America, where it probably did not originate. It has
been noted on practically every variety of cultivated and wild grapes,

and it attacks stems, leaves and berries. Usually it confines its attack
3.5
514 SPECIAL PLANT PATHOLOCxY
to the grape" leaves (Fig. i86), where it produces under ordinary

conditions spots of mildew, especially on the lower leaf surface. In
bad cases, the whole lower leaf surface may be covered with the downy,
or cottony mass of hyphae which gives the fungus its common name.
The parasitic hyphse live in the intercellular spaces of the host and
send into the host cells small knob-like haustoria. The presence of
the mycelium seriously interferes with the normal physiologic activity
of the host. In light cases, the areas of upper leaf surface immediately
overlying the hyphse turn brown in the form of angular spots. Through
Fig. 185. — Black-rot fungus {Guignardia Bidwellii). a. Portion of an affected

grape showing pustules; b, section of pustule (pycnium) showing pycnospores; c,
ascus with ascospores; d, ascospores. {After Quaintance, A. L., and Shear, C. L.,
U. S. Farmers' Bull. 284, 1907-)
the stomata emerge stifif projecting conidiophores which form short

stub-Hke branches from which fall ellipsoidal conidiospores. These
conidiospores are virtually zoosporangia for their protoplasmic con-
tents divide into a number of biciliate zoospores which escape and
swim about in the rain water which covers the washed
leaf or stem, or are
down, or splashed from plant to plant during a dashing rain storm.

When the fungus appears on the fruit, it has been called gray rot, and
occasionally, the berry may be completely covered with a downy mass
of hyphae.
The oogonia and antheridia are not so common as the conidiospores.

If the shriveled parts of the leaves are examined in September, the
Fig. 186. — Grape leaf attacked by mildew, Plasnwpara vilicola, Cold Spring Harbor,
L. I., Aug. 2, 1915.
oogonia will be found as spheric organs attached to the intercellular

hyphae by a short stalk. One or several filamentous curved antheridia
are formed near the oogonia to the surface of which they become ap-
5l6 SPECIAL PLANT PATHOLOGY
plied. A germ tube is formed through which the antheridial con-

tents pass over into the oogonium. A single large central egg-cell, or
oosphere, becomes differentiated in the protoplasm of the oogonium;
this contains a single nucleus in a central position, while the remaining
nuclei pass into the peripheral layer of protoplasm (periplasm). A
single male nucleus passes through the antheridial beak into the
oosphere, which becomes surrounded by a cell wall. Nuclear fusion
now takes place and the oosphere becomes an oospore with a single
central nucleus. The oospores are about 30/x in diameter.
Bordeaux mixture is the most important fungicide used in combating
the downy mildew disease. It is applied as in black-rot.
CHAPTER XXXV
DETAILED ACCOUNT OF SPECIFIC PLANT DISEASES
(CONTINUED)
Hemlock (Tsiiga canadensis Carr)
Heart-rot (Polyporus borealis (Wahl.), Fr.). — This bracket fungus is
distributed widely in North Temperate regions. As a wound para-

site, it occurs on hemlocks, pines and spruces, entering these trees
through the stubs formed by the breaking ofif of branches. The

mycelium gradually grows into the heart of the trees and from there
downward into the roots and upward into the tops. It advances in
definite directions through the wood in the form of cords, or strands,
which run radially, or tangentially, in the channels dissolved by the
action of the enzyme, which formed by the living hyphae. The
is
wood shrinks and the mycelial strands begin to dry up, and the wood
is separated into cuboidal blocks marked off by the channels formed
by enzyme action. If the mycelium attacks the cambium, the trees

die. The bracket-like fruit bodies are soft and spongy and last only a
season. They are, according to Atkinson, lo to 20 cm. (4 to 8 inches)
by 6 to 15 cm. broad. Several of these sporophores may be joined
together. The upper surface is rough, shaggy and has a sodden ap-
pearance. The pores on the under side are quite regular with rounded
openings in some specimens, or irregular, elongated and sinuous in other
samples.
Hollyhock (AlihcBa rosea Cav.) (Fig. 187)
Rust {Puccinia malvacearimi, Mont.).' —

This fungus was introduced
into France about 1868 from where it is native, and in the
Chili,
summer of 191 5, the writer found it very destructive to the hollyhocks
in the gardens on the Island of Nantucket off the southern coast of New
England. It spread rapidly over Europe and came to the United
States in 1886 upon infected seed. The leaves are spotted with the
yellowish-brown sori slightly raised above the leaf surface (Fig. 72), or
they are found on the stem in the form of small wart-like elevations.
The leaves dry up, as if blighted, and during August of 1915 on Nan-
517
5j8 SPECIAL PLANT PATHOLOGY
Fig. 187. —Hollyhock rust, Puccinia malvacearum. i, Typic mature telio-

spore; 2-6, different stages in growth of promycelium (basidium); 7, forked promy-
celium; 8, basidium dividing into 4 cells; 9, basidium resembling a germ tube; 10-12,
cells breaking apart; 13-16, germination of promycelial cells; 17, empty cell; 18,
mature basidiospores; 19, 20, same in germination; 25, 26, formation of chlamydo-
spore-Hke bodies in old promycelia. {After Taubenhaus, J. J.: Phytopath. I, April,
1911.)
DETAILED ACCOUNT OF SPECIFIC PLANT DISEASES 519
tucket only a few host leaves were left on a row of garden hollyhocks,
all of the other leaves having fallen off. The sori consist of light-
colored teliospores which are two-celled and measure 17 to 241J, by 35 to
63M (Fig. 187).

Bordeaux mixture (4-3-50) has been found efficient, as a spray, in
controlling the hollyhock rust. Others recommend sponging the dis-
eased parts with permanganate of potash, two tablespoonfuls of
saturated solution diluted with one quart of water.
Larch (Larix spp.)
Canker {Dasyscypha Willkommii, Hartig). —The life history of this

destructive fungus of larch trees has been studied by German plant
pathologists, so that it is pretty well known. In the moist, marsh
meadows mountains of Europe where the larch has been planted
in the
in pure forests, the fungus has been frequent in past years. The
mycelium attacks the bast elements of the stem and its insidious char-
acter is manifested in the death of the bark, which peels off. Pro-
nounced cankers soon develop and the fungus lives perennially in the
tree spreading rapidly when the larch tree is comparatively inactive,
viz., autumn and winter. The diseased area, represented by wounded
tissue, may heal over during the growing season, but when the fungus
regains its activity the disease progresses until the branch is com-
pletely girdled and its terminal part dies.
Creamy white stromatic tufts appear, where the bark has been killed
and on this superficial mycelium minute conidiophores arise, which
bear simple hyaline conidiospores. As these probably do not germinate
they have no influence in the spread of the canker. Short-stalked
apothecia may appear on the canker areas later in the year. They
are somewhat yellow on the outer surface and orange within. The
cylindric asci (i20;u by gn) bear light ovoidal, unicellular ascospores.
Filiform paraphyses are found between the asci. No efficient remedial
measures are known.
Dry-rot (Trametes pini (Brot Fr.). —This fungus is very common
in the forests of New England, Canada and Newfoundland.
It grows
on nearly all coniferous trees; white pine, red spruce, white spruce,
hemlock, balsam fir and larch attacking the living trees after they
begin to form heartwood. In the tamarack, or larch, the decay goes
much beyond that of the spruce and balsam fir. In the early stages,
according to von Schrenk, small white spots appear, which occupy the
entire width of an annual ring. Two or more of these spots soon join,
at first in a longitudinal direction, then laterally also, so that one or

more rings of woods are transformed to cellulose. The rings are thus
separated from adjoining ones so that a series of easily separable
tangential plates are formed. The line of separation between the
rings is always at the point where the summer wood stops and the
spring wood of the following year begins.
The progress of decay is marked by the attack of more and more
sound wood fibers which are reduced to loose fibers of cellulose until the
wood has disappeared, when black lines appear, scattered irregularly.
The tangential plates become ultimately extremely thin and they then
consist of the resistant summer wood cells more or less infiltrated with
resin. The whole of the former woody cylinder becomes a mass of
separate fibers which can be pulled out individually.
The fruiting organfound commonly on all of the afi"ected trees.
is
It is readily distinguished from allied forms by the light red-brown

color of the hymenial surface, and the regular small round pores. The
form of the pileus varies greatly. Sometimes the brackets are large
on the larch, lo cm. (4 inches) in width laterally, 7 cm. (2.8 inches) from
front to back, and 5 cm. (2 inches) in thickness, and are formed at the
ends of old hard stubs and at scattered points on the bark. Some-
times sessile sheets are formed inside of the brackets. The basidia,
which form the hymenial surface that lines the pores, are smaller at
the apex and form from slender, spore-bearing sterigmata. The
basidiospores are brown at maturity.
Lemon {Citrus limonum, Risso.)
Brown-rot {Pylhiacystis citriophora, R. E. Smith). — The disease is
characterized by a copious exudation of gum from the trunk just above

the bud union. A certain area of the bark surrounding the part which
shows gummosis dies, becomes hard and dry without any evidence of
the fungous parasite. It appears especially destructive on the fruits
after packing, and is recognized as a brownish, or purplish, discolora-

tion of the rind, which is lighter green than on the ripe fruits. It
spreads rapidly from fruit to fruit, and is also characterized by its

peculiar odor and the presence of small flies attracted to it. The
—
DETAILED ACCOUNT OP SPECIFIC PLANT DISEASES 521
mycelium penetrates the lemon rind and consists of much-branched

extensive hyphae of irregular diameter. Conidiospores which repre-
sent zoosporangia appear under fayorable conditions. They measure
20 to 60 by 40H to gofx and are lemon-shaped with a pronounced protu-
berance at the apex. Upon opening a number of biciliate zoospores
are liberated.
Infection of the fruit usually takes place in the orchard and also
during the operation of washing the lemons preparatory to packing
them. The wash water, therefore, should be treated with copper
sulphate, formalin, or potassium permanganate. In using formalin,
il is made up in one part to ten thousand parts of water, or i pint to
about 1200 gallons. Where more
the cheaper copper sulphate is
available, ipound should be dissolved in 250 gallons of water.

Sooty Mold (Meliola Penzigi, Sacc, and M. camellice (Catt.) Sacc).
This fungus is widely distributed in those districts where citrus fruits
are grown. It is most injurious to the orange, but occurs on the
lemon as well, appearing on both leaves and fruits. The mycelium
forms a sooty black covering on the leaves, twigs and fruits and is
usually associated with various scale insects and aphids, which exude
a honey dew upon which and the dead bodies of the scale "insects the
fungus feeds as a saprophyte. The mycelium consists of large branched
threads, which are closely septate, and the branches are cemented
together to form a false stratum, which lives purely as a superficial
saprophytic growth without penetrating into the tissues of the citrus
plant on which it is found. Certain hyphal branches flatten out and
probably serve as appressoria. The reproductive cells are of various
kinds, such as stylospores in pustules, pycnidia with pycnidiospores
(pycnospores) and perithecia. The stylospores arise from small
conidiophores within peculiar, elongate, flask-shaped structures. The
pycnidia are' small and scattered. The perithecia are spheric and in
close asci with eight dark elliptic, three- to four-septate spores.
The most effective substance for the treatment of sooty mold has
been found by Webber to be the resin wash.^ The mixture consists of
Resin 20 lb.
Caustic soda (98 per cent.) 4 lb.
Fish oil crude 3 lb.

Water to make 15 gal.
1 DuGGAR, B. M.: Fungous Diseases of Plants: 215.

Webber prepares the mixture as follows: Place the resin, caustic

soda and a large kettle, pour over them 13 gallons of water,
fish oil in
and boil until the resin is thoroughly dissolved, which requires from
three to ten minutes after boiling has commenced. While hot, add
enough water just to make 15 gallons. It is advised to make about
two sprayings when the white fly (Aleyrodes) is in the larval stage.
In Florida winter sprayings are important, but a spraying in May is
also often desirable. In all cases dilute the stock solution with 9
parts of water.
Lettuce (Ladtica saliva, L.)
Drop {Sclerotinia libertiana Fckl.). —This is one of the most disas-

trous of the sclerotium-producing fungi to garden and greenhouse
plants, being widely distributed and difficult to control. It attacks
greenhouse lettuces, causing at first flagging, then indications of

water-soaked areas over the stem and basal part of leaves, finally fol-
lowed by the collapse of the whole plant into a formless mass. The
mycelium may grow on the surface of the lettuce leaves and black
sclerotia may be formed there commencing as white condensations
which finally turn black. Conidiospore formation is not certainly
known in the lettuce-drop fungus. Sclerotia, however, are commonly
formed which measure 3 cm. in length and these are formed even on
artificial culture media. The apothecia are wineglass-shaped with
long black stalks. The
formed on the upper depressed side of the
asci
apothecia are cylindric and measure 130 to 13 5^ by 8 to loû, while the
ascospores are small, 9 to 13/i by 4 to 6.5/1.
dead and diseased lettuce plants should be destroyed by fire

All
and the ground where they grew soaked with some suitable fungicide
so as to confine, or practically exterminate the disease. The soil
should be sterilized with steam before planting.
Lilac {Syringa vulgaris, L.)
Powdery Mildew {Microsphcsra alni (Wallr.) Wint.).— During the

summer months and late in the autumn, the upper surface of the leaves
of the lilac will be found covered with a whitish mildew which consists
of interlacing hyphae, which form a cobwebby, superficial growth.
Short haustoria are produced which grow into the epidermal cells.
The mycelium develops upright vertical conidiophores which abstrict

off conidiospores in chains.These conidiospores no doubt account
for the rapid spread of the disease, which is never very serious to the
Hlac shrubs, but no doubt to some extent interferes with the normal
physiologic processes of the leaves. Subsequently perithecia are
formed which are spheric in shape, almost jet black in color, and which
are surrounded by a circlet of hyphae known as appendages, which are
curved or dichotomously hooked at the extremities. Each perithecium
produces 3 to 8 asci, and each ascus contains 4 to 8 relatively small
ascospores, which measure 18 to 23/x by 10 to 12/x (Fig. 54).
Maple (Acer spp.)
Decay {Fomes fomentarius (L. Fr.) (Fig. 188). —The sporophores

of this fungus are hoof-shaped and appear first as small rounded knobs
on the surface of the trunk, or at branch stubs. The upper surface is
smooth and more or less definitely marked by concentric ridges. The
older fruit bodies owing to the action of the weather are uniformly
gray and appear as if powdered. The lower surface is reddish-brown
in color and shows numerous, small round pores. The margin of the
new layer is grayish white and very soft and velvety. The sporo-
phores are found usually singly, althoughby proximity of two, or several,
they may appear grouped together. The decay produced in the wood
of deciduous trees by Fomes fomentarius begins in the outer alburnum
immediately beneath the barky layers, and extends inwardly, until
it reaches the pith of the tree. The rotten wood is distinguished by
a large number of irregular black fines outlining areas of sound wood.
Wholly decayed wood is extremely soft and spongy, fight yellow and
crumbles into numerous separate wood fibers when rubbed. The
tinder fungus, Fomes fomentarius, found in the deciduous forests of
is
Michigan, Minnesota, New England, New York, Wisconsin and in

other states. It grows rapidly in dead wood and the mycefium will
form large masses if the infected timber is kept under moist conditions.
Leaf-blotch (Rhytisma acerinum (Pers.), Fr.). —The tar spot of the

maple is found about Philadelphia usually on the silver maple to which
it does slight injury. The black irregular spots are, however, alwavs of
interest to the laymen and questions are asked frequently about their
cause. The spotbegins, as a yellow thickened area, when the maple
leaves are expanded fully. The epidermis

pushed up by short conidio-
is
phores which arise from a hyphal stroma beneath. These conidio-

phores produce unicellular, curved conidiospores which serve to dis-
tribute the fungus. Formerly this stage was called Melosmia. Later
r-^
Fig. 1 88. —
Cross-section of branch of dead beech rotted by Fomcs fomenlarius.
(After von Schrenk, Hermann, Bull. 149, U. S. Bureau of Plant Industry, pi. viii,
1909.)
as the season advances, the hyphae become massed into a sclerotium-

like area black without, but white within, and this persists after the fall
of the leaf. Sometime the next spring, there arise from these sclerotia
complex apothecia often 1.5 cm. broad, which break through at irregular
fissures. The club-shaped between

asci bear eight acicular ascospores
which are found paraphyses with hooked These ascospores tips.
measure 65 to 8o^t by 1.5 to 3/i and are ejected forcibly from the ascus.
As the disease is not a serious one, usually no remedial measures are
necessary. If the owner of maple shade trees wishes to keep it in
check, he should burn the dry maple leaves which litter the ground
about his place.
Melons, Squashes, Watermelons {Cnciirbita spp.)
Anthracnose (Colletotrichum lagenarhim (Pass.), Ell. & Hals. — As

an illustration of a disease-producing fungus included among the Fungi
Imperfecti, we may describe briefly the anthracnose of cucumbers,
squashes, watermelons, Colletotrichum lagenarium, which attacks
both leaves and fruits. The leaves are found with brown spots which
cause their early maturity. If the fungus attacks the fruits, it produces
sunken water-soaked spots in which the acervuH appear. The acervuH
produce numerous conidiospores sticking together to form viscid
masses of a pink color. During moist weather, the hyphae may grow
out, superficially covering the fruit with a mold-Uke growth. The
fungus eventually causes a complete decay of the fruit. The disease
has been prevalent in Nebraska and New Jersey. If the disease
appears in greenhouse culture, it is well to sulphur the greenhouses
thoroughly when they are empty, and to clean and whitewash all the
walls and woodwork to destroy any funguses present. Spraying with
Bordeaux mixture (3-6-50) should begin when the vines begin to trail
over the ground. Subsequent sprayings should be made every ten
days, if the weather is dry.
Wilt (Bacillus tracheiphilus, E. F. Sm.). — This serious disease of
cucurbitaceous plants was first reported by Erwin Smith about 1893.
It was first known in the northeastern states, but it is now common in
the middle west and Rocky Mountain Although pumpkins

regions.
and squashes may be attacked by wilt, yet cucumbers and melons are
most susceptible. This microorganism, which is a rod-shaped bacillus
two or three times as long as broad, is actively motile by wavy cilia
only when young. It measures 1.2 to2.5;u by 0.5 to 0.7/i. It causes a
progressive wilting of the host which it attacks. Whether the whole

plant dies depends upon the point of infection, which is usually ac-
complished by^biting insects. If a leaf is attacked, it dies back to the

stem. If the basal part of the stem is infected, the plant rapidly
succumbs. This rapid wilting due to the fact that the organism
is
lives in masses in the vessels of the xylem by which the water taken
up by the roots is distributed throughout the plant, hence any occlusion
of these spiral and pitted vessels stops the water supply and the plant
suffers. Advanced stages of the disease may be characterized by the
disintegration of the vascular system and the formation of cavities in
the adjacent parenchymatous tissue. Smith sums up the cultural
characteristics of this organism, as follows: Stains readily; smooth;
white; viscid; glistening; slow grower on media; surface colonies small,
round, discrete; nogrowthat 37°C.orat 6°C. (i6days); aerobic; faculta-
tive anaerobic (with grape-sugar, cane-sugar or fruit-sugar); usually
it grays potato after a time; clouds peptone-bouillon and Dunham's
solution thinly; growth retarded in acid juice of cucumber-fruits;
also retarded or inhibited by juice of many vegetables, e.g. table-beet,
sugar-beet, turnip, etc.; grows on many media at 25°C., carrot, coco-
nut, etc.; thermal death point 43 °C.;optimum for growth 25° to 3o°C.,
maximum, 34° to 35°C.; easily killed by dry-air, sunlight, freezing;
ammonia production moderate, in litmus milk persistent growth without
reduction or distinct change in color of litmus; killed readily by acids.
Group No. 222, 232, 2023. As the disease is distributed by insects,
the grower of cucurbits should endeavor to reduce the number of
these pests by the use of kerosene, or arsenate spray, and trap plants
should be grown to attract the insects away from the more valuable
plants.
Oak {Quercus spp.)
Decay {Polyporus sulphureus (Bull.) Fr. Figs. 189 and 190).- The —
decay induced by Polyporus sulphureus is often called the red heart-rot.
It attacks not only oaks, but also the chestnut, maples, black walnut,
butternut, alder, locust, etc. It is widely distributed in North America
and Europe. The sporophores of this fungus form a series of superim-
posed, fleshy brackets of a sulphur-yellow color, weighing in the aggre-
gate at times almost one hundred pounds (Fig. 189). The color some-
times may vary an orange-red. The under surface is usually a light
to
yellow color and beset with numerous minute pores. At maturity, the
fruit bodies lose their soft character and become harder and more brittle.
and frequently, become the prey of maggots which riddle them with
holes and burrows. It is also eagerly gathered by mycophagists who
know it to be an excellent article of food.
The mycelium of the fungus may live in the dead wood of a tree
after it has been killed for a number of years, so that the same tree may
produce successive crops of edible fruit bodies. The destruction, which
the mycelium works, is characteristic. The heartwood is reduced to a
crumbly brown mass which resembles charcoal in its fracture, but is
Fig. 189. — Fruiting body of Polyporus siilphureus. {After von Schrenk, Hermann,
Bull. 149, U. S. Bureau of Plant Industry, pi. iv, 1909.)
red-brown in color. The decayed wood shows concentric and radial

cracks extending irregularly through it (Fig. 190). As the wood is at-
tacked and destroyed by the spreading mycelium, these cracks increase
and in them are found leathery compact sheets of mycelium, which can
be isolated by reducing the decayed wood to a fine powder by the blows
of a hammer. The wood decays uniformly and is converted into a
brittle brown substance, which can be rubbed to a fine powder between
the fingers. Von Schrenk found that the youngest trees in which the
red heart-rot occurred were about 50 years old. The removal of dis-
eased trees seems to be the only efficient method of checking the spread
of Polyporus siilphiireus.
Honeycomb Heart-rot (Stercum subpileatnm, W. H. Long). —The

pocketed, or honeycomb, heart rot has been found on the following.
Fig. 190. —
Cross-section of a living post oak tree rotted by Polyporus sul-
phureus. {After von Schrenk, Hermann, Bull. 149, U S. Bureau of Plant Industry,
.
pi. iv, 1909.)
nine species of 02i\i's,:Querciis alba, MichauxU,

Q. lyraia, Q. marilandica, Q.
Q. minor, Q. palustris, Q. texana, Q. velutina and Q. virginiana}
The first indication of this honeycomb heart-rot in white oak is a
slight discoloration of the heartwood, which assumes a water-soaked
appearance, which may extend from i to 6 feet beyond the actual decay.
1 Long, W. H.: A Honeycomb Heart-rot of Oaks caused by Skrcuin subpileatnm,
Journal of Agricultural Research V: 421-428, Dec. 6, 1915.
DETAILED ACCOUNT OF SPFCIPIC PLANT DISEASES 529
The water-soaked heartwoocl becomes tawny in color when dry.

Light-colored, isolated areas now appear in the discolored wood and
these areas originate the pockets. The rot spreads in all directions into
the surrounding tissue, but more rapidly in the summer wood of the
annual ring of the preceding year, so that the bulk of the pocket lies
in the summer wood of one year and the spring wood of the succeeding
year. Delignification now follows in which delignified wood fibers
appear in patches in the light-colored areas, and this delignification
spreads rapidly until white, oval to circular pockets are formed.
These lens-shaped pockets are at first filled with white cellulose, which
is later absorbed, leaving cavities. The diseased area increases in size
until the pockets reach a large medullary ray, which seems to check the
activity of the enzyme, so that the larger medullary rays become the
radial walls of the pockets. All the cellulose finally disappears, leaving
the pockets either (i) empty, (2) containing the shrunken white
membranes of the included vessels, or (3) more or less filled with myce-
lium. The last stage of the rot is characterized by the very light and
honeycombed nature of the wood. The pockets are longer than
they are broad, and all of the wood has disappeared, except the thin
walls around the pockets, which remain distinct and usually involve the
heartwood uniformly. The rotted wood is, therefore, in the shape of a
cylinder and there is a brownish discoloration of the heartwood on the
outer edges of the affected area.
The growth of the mycelium seems to be preceded by the enzymes
which cause the disintegration of the wood. A few of the larger vessels
show hyphal threads and these become more 'numerous, as delignifi-
cation advances, until they become stuffed with small, intricately
branched, colorless hyphas. When the hyphae are exposed to the air,
they become brown ia color. The sporophores are found on dead

trees, or the dead areas of living trees. The sporophores are thin
shelving bodies formed in. the cracks of the bark, sometimes assuming
a conchate shape. They sometimes form in parallel lines, and range up
to 5 cm. in width. These sporophores may be formed on the dead tree
for a number of years. This fungus is widely distributed in the southern
states and ranges as far north as Ohio. The only method of control
is to prevent the infection of trees by eliminating forest fires, by pre-
venting the formation of the sporophores, and the destruction of all
diseased timber which has the rot.
Root-rot (Armillaria mellea, Vahl).' —The "hallimasch" of the

Germans, or the so-called honey mushroom, is a fungus of considerable
interest to the forester (Fig. 15). The spores, ifblown to an exposed
branch stub, may germinate and produce a mycelium which works up and
down the tree. Infection may be also by the mycelium growing across
from the roots of a diseased tree to a healthy one through the soil of
the forest. In either case, the young mycelium grows into the cambial
and finally completely encircles the trunk
layer, attacks the living cells,
ofan infected tree. Later the hyphae are converted into strands, which
show a characteristic apical growth, thus providing for the elongation
of the strands through the host. The strands of hyphae turn a deep
chocolate-brown color and are known as rhizomorphs (Fig. 15), which
may anastomose under the bark of the tree. Ultimately, as the tree
dies, the bark splits off and the rhizomorphs are found flattened against
the woody cylinder of the tree. If such trees are used as mine props,
the strands may keep on growing under the moist even temperature of
the mine and there they may hang down in long streamers into the mine
galleries, as specimens of such in the botanic museum of the Univer-
sity of Pennsylvania indicate. The effect of the mycelium in the tree
is to kill its top with the ultimate death of the entire tree. The
rhizomorphs formerly known as Rhizomor pha subterranea grow out
into the root system of the tree, which they kill, and here they may
live for a number of years, endangering the nearby healthy trees,
because they extend out into the toward other tree roots. It is
soil
this subterranean growth, which makes the honey mushroom an ex-

tremely dangerous oife to the hardwood forests, where it is found.
The fruiting bodies of this fungus usually occur grouped in considerable
numbers about the base of the affected tree arising from the dark-brown
rhizomorphs, which thus serve to connect together isolated groups of
the sporophores. The sporophores produced most commonly from
September to November are honey-colored, i.e., yellow to orange-
brown, and their umbonate tops have a more or less viscid character
with small black spicules scattered over the surface. The stipes are
slightly swollen at the base and a short distance below the pileus is
found the ring, or annulus. The lamellae are dirty-white and from
each pyriform basidium four white basidiospores fall until surround-
'LoNG, W. H.: The Death of Chestnuts and Oaks due to Armillaria mellea.
Bull. U. S. Dept. Agric. No. 89, 1914.
1
ing leaves and mosses may be coated with a mealy powder derived from
the gills of several sporophores directly over them.
Oat (A vena saliva, Linn.)
—
Rust {Puccinia coronifera, Kleb) . The oat rust, or crown rust, affects
oats and also several other grasses. The summer stage appears on
oats just prior to the period of ripening where it forms an elongated
uredinium of an orange color on the leaves and sheaths. The globular
spores germinate readily. The teliospores are formed later as black
spots around the edge of the uredosori. As the teliospores bear at
their apex a crown of blunt projections, or processes, the common name
of "crown rust" has been applied. Such winter spores remain in a
resting condition until the following spring, when they germinate in the
usual way. The basidiospores, which are formed from the basid-
ium, or promycelium, begin growth on the leaves of the buckthorn,
Rhamnus cathartica, where within eight to ten days cluster cups
(yEcidium catharticce) appear. The aeciospores germinate readily and
are blown to the oat and other grasses, such as perennial rye grass,
Yorkshire fog, so that at least eight forms of the species limited to
certain hosts have been distinguished. The measurements of its spores
are as follows: ^ciospores, orange, vermiculose, 16 to 25/x by 12 to 20^1;
Uredospores globose to obovate, echinulate yellow, 18 to 2 7yuby 16 to
24/i; teliospores brown, two-celled, crowned with rough projections;
approximately 35 to 60^1 by 12 to 22)u.
Smut (Ustilago avencB and U. levis). The a-ppearance of this dis-
ease is illustrated in the figures (Fig. 191).
Onion {Allium cepa, L.)
Smut (UrocysHs ceptdcB, Frost). — This fungus, probably of Ameri-

can origin, is found in the onion growing districts of the eastern United
States where it has been known for about 50 years. The smut fre-
quently appears soon after the first leaf appears, and is first in the form
of dark spots at the base and on succeeding leaves, as
of the first leaf
they make These spots are followed by longitudinal
their appearance.
cracks, which show, a granular spore powder associated with threads
of fibrous tissue. The spore powder under the microscope is found to
consist of the spore balls, which number several compacted cells, the
central one of which contains cytoplasm, being surrounded by an

envelope of sterile cells. Such spore balls are 17 to 25)11 in diameter
Fig. 191. — Smut of oats. A, UsLilago avence; B, Usltlago levis {After Jackson,
H. S., Bull. 83, Del. Coll. Agric. Exper. Slat., December, 1908.)
and may retain their capacity for germination in the soil for a period of
12 years.
As the spores occur in the soil, it is useless to treat the onion seeds
with chemic bodies. The most successful method of prevention is to
transplant the seedlings into beds known to be free from smut. Some
growers place sulphur (100 pounds to the acre) and air-slacked lime
(50 pounds) in the drills as the seeds are planted.
Orange (Citrus aurantium, L.)
Black-rot {Alter naria citri). —Only navel oranges are subject to

black rot which is recognized by the premature ripening, large size of
the fruit and its deep red color. The fungus gains entrance through
the navel end, because there imperfections of the skin occur. There
soon arises a black area of decay under the peel which remains isolated
for some time without spreading, therefore, the disease is not very
virulent. In Alter naria, the conidiophores are in bundles, always
unbranched and short. The conidiospores are club-shaped to flask-
shaped, divided and united into chains by thinner cells.
Fruit-rot {Penicillium italicum, Wehm.). —A large part of the decay

of the orange and other fruits of the genus Citrus is due to blue and
green molds. These molds usually cannot enter uninjured fruits, and
so their attacks usually follow a bruise occasioned by careless handling,
or when the fruit falls from the orange tree. Penicillium italicum seems
to be more common than the other species, P. digitatum. Pure cultures
of this fungus can always be secured from decaying oranges in the
market, which have the blue-green areas of rot just beginning to appear
upon them. These areas are usually blue-green in the center sur-
rounded by white areas which are grouped usually into little white
patches toward the vegetative margin and the whole superficial colony
surrounded by an area of soft watery rot. Sometimes, as the colonies
become older, P. digitatum mixes with P. italicum.
The conidiophores are short (looju), or very long (6oo;u) and black
in media containing sugar. They average about 250^ in length. The
conidial fructifications are up to 300/1 or more in length, consisting usu-
ally of a main branch and one lateral branch, each producing a whorl
of branchlets bearing crowded verticils of conidiospores, 12 to 14^1
by 3/i. The chains of conidiospores are cylindric to elliptic, slightly
ovate, clear green by transmitted light and measure 2 to 3/x by 3 to 5)u.
Decay of this sort can be prevented by careful handling of the fruit in
field and packing house.
Pea {Pisum sativum, L.)
Pod-spot {Ascochyta pisi, 'Lih.). —The horticulturist, who attempts

to grow the garden pea, will find that the leaves and pods become
spotted with conspicuous, circular, sunken spots 3 to 6 mm. in diameter,
which are dark bordered, pale in the centers and slightly pinkish when
mature. Pycnidia are associated with these spots and out of their
porous opening under favorable conditions the spore masses may be
seen issuing. When the leaves are affected, it is usually the lower
leaves which become diseased first, and such soon die. If the stems
are attacked, the spots sometimes penetrate through the woody part.
Different races of peas differ as to their susceptibility. The variety
Alaska is slightly affected, while the varieties American Wonder,
French June and Market Garden are frequently badly diseased.

According to Stevens, the pycnidia consist of angular cells, 5 to 7/1 with
a rounded ostiole and reddish-brown surface. The conidiospores are
constricted slightly at the septum, are oblong and measure 12 to i6m by
4 to 6ju. The mycelium perennates in affected seeds, reduces their
power of germination and carries the fungus over to the next crop.
Selby has indicated that healthy peas may be grown by spraying with
Bordeaux mixture, and it has been suggested, that a two years' rotation
of non-susceptible crops lessens the prevalence of the disease, if another
pea crop is raised.
Peach {Amygdalus persica, L.)
Leaf Curl (Exoascus deformans (Berk.), Fckl.) (Fig. 192). —This

disease is by the French Cloque du pecker, by the Germans
called
Krauselkrankheit and by Americans and English peach leaf curl. It
is widely distributed through America, Europe, China and Japan and
in Africa and Australia, so that it is practically cosmopolitan.

The disease is most prevalent and most disastrous to the leaves and
tender shoots of the peach, when the spring months are damp and cool,
for records show that such conditions prevailed during April of the
year 1893, 1897 and 1899, when peach leaf curl was especially abundant
in Ohio and New York. Warm and relatively dry springs seem to be
unfavorable to its occurrence. The susceptibility of the host plants
differs to a marked extent, some being susceptible, others less so.
The presence of the disease may be detected when the leaf buds
unfold, for the coloring of the young leaves is heightened, and as they
open out, the curling and arching of the blades become manifest.
The curling may be confined to a small portion of a leaf, or it may be
general and all of the leaves of a tree may be affected, as well as the
young stem on which \hey are found. The green, or reddish, color of
Fig. 192. — Peach leaves deformed by leaf curl {Exoascus deformans). (After Heald,
F. D., Bull. 135 (Set. Ser. 14), Univ. of Tex., Nov. 15, igoQ-)
the leaves is lost as they mature, and they become pale, or slightly
discolored. Diseased shoots may grow to twice their normal diameter
and assume a characteristic paleness. The diseased leaves finally
turn brown and drop off the tree, and if this defoliation is excessive
the crop of peaches may be nil. The twig affection is sometimes

associated with gummy exudations, particularly when the enlargement
is terminal. It is doubtful whether the mycelium perennates in the
twigs, as was supposed in former years. Infection must generally
occur as the buds unfold.
The mycehum of the fungus may be studied most advantageously
in the leaf before the fungus has appeared on the surface. At that time,
the hyphze show a greater protoplasmic content and sections reveal
the fact that the intercellular inycelium is distributed through the
mesophyll and cortex of the young stems. Pierce distinguishes vege-
tative hyphae, distributive hyphae and fruiting hyphae. The latter
push up between the epidermal cells and a series of short hyphal cells
are formed, as ascogenous which form an almost continuous layer

cells,
beneath the cuticle. The ascogenous cells give rise to the asci, which
push through the cuticle. An ascus is usually truncate at the exposed
end and it gives rise to four to eight ascospores, which may bud within
the ascus.
may be controlled by the use of lime-sulphur solution
Leaf curl
(1-20), Bordeaux mixture (5^5-50) and copper sulphate in water
(2-50), for the use of which the practical man is referred to the spray
calendar given in the subsequent pages of this book.
Pear (Pynis commurds L.)
Fire-bhght (Bacillus amylovorus De Trev. Toni).^ This

(Bun.), —
bacterial disease is found on the apple, pear and quince, but more
especially on the pear, so that it has been termed pear blight. It was
first reported from the northeastern United States, but now it is dis-
tributed throughout the country from the Atlantic to the Pacific
oceans. The disease first makes its appearance in the early part of the
season, when it appears in the form of a twig blight throughout the time
of blossoming of apples and pears, when the blossoms and tips begin
to wilt and show signs of blackening. This results in the complete
blackening and death of all the short branches, or spurs, upon which
flower clusters have been borne. The fire blight disease may continue
to extend down the twig, or the branch, the branch being entirely
killed, as it progresses. Under conditions more favorable to the host
1 Orton, C. R. and Adams, T. F. : Collar-blight and Related Forms of Fire-

blight. Bull. 136. Penna. Agricultural Experiment Station August, 1915.
the blight may extend only a short distance, which results in tip prun-
ing. The bark of the tree indicates the progress of the disease, for
the soft bark assumes a water-soaked appearance followed by a blacken-
ing and shriveling. When the organism ceases to spread rapidly in
the tissues, there appears a sharp Hne of separation between the dead
and the healthy tissues. The bark is broken and through the bark
cracks appear gummy, or gelatinous, drops which vary in color from
white to brown, or black.
Bacilhis amylovorous was described first by Burrill in 1877, a dis-
covery full of significance to plant pathology, because it established
the first bona fide case of a plant disease due to bacteria. It has been
established, that infection takes place through the visits of insects,
especially bees, to the pear flowers. From the floral nectary, the
bacillus spreads to the softer tissues of bark and cambium, where
it is very largely confined, and where it winters over, spreading to
other blossoms the next spring. Bacillus amylovorus is an oval
microorganism 1.5^ to 2^1 long, growing singly, or several attached
end to end, and is motile in fresh cultures. On agar, the cloudy
and white surface colonies appear the second day, and attain a di-
ameter of 2 to 3 mm. by the fourth or fifth day. Cloudiness appears in
bouillon after twenty-four hours, and in milk, thickening of the medium
begins at the third or fourth day, which increases until the fifth, or
sixth day, when the product is finally partially gelatinous with a clear
acid liquid above, changing to slightly alkaline.
The work of Waite has shown that pear blight can be controlled
by pruning out the blight during winter, so as to eliminate the source
of infection during the next year, and if this pruning is done thoroughly,
the disease can be kept in check. The stubs should be disinfected
with corrosive sublimate (i-ioo).
Pine (Pi litis spp.)
Blister-rust (Cronartium ribicoliim, Fisch & Waldh. = Pcrl-

dermium strobi, Klebahn).^ —This disease, as it appears on white pine,
Spaulding, Perley: The White Pine Blister Rust Situation, American
^
Forestry 22, pp. 137-138, March, 1916; The BHster Rust of White Pine, Bull. 206,
U. S. Bureau Plant Industry, 191 1; also consult American Forestry. Feb., Mch., Dec,
1916. In the December, 1916, number a map showing the distribution of the
disease is given. A conference was held at Washington in January, 1917, to
consider the establishment of stricter quarantine regulations of the methods of
checking the spread into the western states.
has been considered to be of such great importance, that strict quaran-

tine regulations were established in order to keep it out of the country,
but the result of a thorough exploration of the New England States
during the summer of 1916 has shown its general distribution through-
out them and even as far west as Minnesota. It appears to have been
Fig. 193. — White pine blister-rust, Cronarliutn ribicola. A, Diseased tree with
aecial blisters broken open from which spores are blown to currant or gooseberry-
leaves; B, D, teliosori on under leaf surface of currant, Ribes. {From Gager, after
Perley Spaidding.)
introduced into America on nursery stock from Holland, and all the
trees in these advanced posts of infection have been destroyed. In
igo6, there was an outbreak on currants at Geneva and measures were
taken to destroy the fungus in that vicinity. The aicidial stage, known
as Peridermium sirobi, appears on the pine tree and the uredinia and the
telia on species of the genus. Ribes, viz., R. aureum, R. nigrum, R.

rubrum with which intermediate hosts (it does little damage. The
susceptibiHty of different currants varies considerably (Fig. 193).
The attacked white pine trees are stunted, the tops show a bushy
growth and the part of the tree where the mycelium occurs is swollen.
The leaves of the currant infested by the fungus are thicker in texture
and assume a different color. The aecidia are erumpent from the bark
in the form of a bladder with an inflated peridium about one centi-
meter high and yellowish-white. The spores are roundish, or poly-
gonal, coarsely verrucose, orange in color and measure 22 to 29/i by 18 to
20/i. The urediniospores form orbicular groups surrounded by a deUcate
peridium which opens at the summit with a pore. They are ellipsoid
to obovoid in shape, echinulate, orange and their dimensions are
21 to 24/i by 14 to i8/i. The smooth teliospores are crowded along the
veins of the leaf. They are orange to brownish-yellow, 70/i long by
2i/i broad.
This serious disease may be controlled by the destruction of the
hosts, namely, the currant and gooseberry bushes especially in the wild
state. This disease threatens the extinction of all the species of five-
leaved pines including those of the Pacific States, such as sugar pine,
Pinus lambertiana.
Red-rot {Poly poms ponder osus, H. von Schrenk). —The red rot of
the western yellow pine {Pinus ponderosa) usually starts in the tops of
the "black-top" trees, i.e., trees which have been dead for two or more
years. At one or more points, one will find that the wood immediatelv
under the bark starts to rot and the rot proceeds inwardly to the wood
which becomes wet and soggy, and rapidly becomes brittle, so that it
crumbles into small pieces when rubbed. The color of the wood changes
to blue and later to red yellow. When the decay has gone on for
some time, bands and sheets of a white felty substance consisting of
masses of hyphae are found filling certain cracks which result, because
of shrinkage in the wood mass. The destruction of the wood continues
until the heartwood is reached.
Red-rot is caused by a higher fungus which enters the tree through
beetle holes made
into the dead cambium of the wood killed by the
"blue" fungus which precedes the red rot. When the wood has been
completely destroyed red-rot fungus forms its sporophores which begin
to grow out from the mycelium, as flesh-colored knobs, which rapidly in-
crease in size and turn reddish in color, assuming the form of a bracket,
or shelf. The lower surface is beset with pores, or tubes, on the

walls of which the spores are borne. This bracket fruit may grow
many years, and it adds a ring on the
outside when new growth com-
mences. The fruit bodies may occur
singly or in groups of two or three
together. They are rough on top
and appear to be covered with a waxy
substance, which has hardened and
cracked. It is and readily
brittle
soluble in alcohol and xylol. The
lower surface is smooth with regular
^
pores.
Plum {Primus americana, Marsh)
Black-knot (Plowrightia morbosa

(Schw.), Sacc). The black knot—
was at first mainly confined to the
New England states, but it now ex-
tends across the northern United
States to the Pacific coast with
areas free from the disease in the
middle west and southwest. Several
species of plums and cherries are sus-
ceptible.
The disease appears as wart-like
excrescences on the smaller and
larger branches of plum trees (Fig.
FiG. 194. — „, , , ,
Black-knot of plum,
Plowrightia morbosa, on cultivated
,
^ ....
104) which
pletcly killmg the
it com-
either surrounds
termmal part of
. ,
plum, Cold Spring Harbor, L. L, July the branch, or Only part way round
when the branch continues living
and fruit-bearing (Fig. 194). The common name is well given, because
Won Schrenk, Hermann: The "Bluing" and the Red Rot of the Western
Yellow Pine, with Special Reference to the Black Hills Forest Reserve. U. S.Bureau
of Plant Industry Bull. 36, 1903.
the hypertrophies are black in color. The knot begins as a slight

swelling of the branch, and as the swelling increases in size the bark
is cracked (Fig. 194).
The mycelium of the fungus occupies the cambium and bast areas
of the stem, extending throughout the cortex also. The knot consists
of dense areas of the fungus and tissue elements of the host. Bast
fibers, parenchyma cells and even vessels may be found in the gall
tissue. In the spring, small greenish areas may be noticed on the

surface of the knot, and later, the hyphae break through the bark in
all directions and form a pseudoparenchymatous layer. This stomatic
layer gives rise to the conidiospores, which are fiexuous and septate.
The conidiophores are 40 to 6o/i by 4 to 5)U and the conidiospores
abstricted off are light brown in color. Conidiospores are formed
from Spring to late midsummer. They are simple and light brown
in color. The fungous stromata is covered with papillae which locate
the opening of the perithecia which include the asci with eight asco-
spores, that ripen during midwinter, or later. Each ascus is 120/i in
length and the ascospores measure 16 to 20 fx by 8 to lo/j.. Between

the asci are paraphyses.
Since the conidial stage is produced during late Spring and early
Summer pruning out the developing knots is found an efficient remedy
in most cases against black knot.

Plum Pockets {Exoascus Pruni, Fckl.).^ —The plum pocket fungus is
widely distributed over the United States and Europe and its etiology
of the disease it produces is somewhat similar to that of the peach leaf
curl. The mycelium lives in the flower buds and causes remarkable
changes in the ovaries, as they develop into fruits. The hyphae are
found in the mesocarp, the cells of which are stimulated to form a
spongy growth, so that the plum fruit becomes swollen and somewhat
distorted. As a result of the fungus attack, the endocarp which nor-
mally would develop a putamen, or stone, fails to do so, and no stone,
or seed, is formed, but in their place a cavity appears which gives the
common name to the disease. The mycelium is probably perennial in
the twigs of the plum tree and is, therefore, in a position to grow out
into the young ovaries of the next succeeding crop of flowers. The
ascogenous cells develop beneath the cuticle of the well-formed fruits
and finally rupture the latter, appearing as a velvety layer. The asci
are 30 to 6o/i by 7 to 12^1, although Robinson notes a certain dimor-
phism of the asci where these figures vary. Each ascus contains eight
ascospores which measure 4 to 5/i (Fig. 42).
Potato {Solaniim tuberosum,!^.)
Late-bHght {Phytophthora infestans, deBy). — Historically, this is
one of the most interesting of fungi, for in 1845 the potato crops
of the British Isles, especially Ireland, were decimated by the late
blight to suchan extent as to cause a severe famine in Ireland. This
famine caused the emigration of hundreds of thousands of people from
the Emerald Isle to America and the British parHament in order to
alleviate the distress of the poor repealed the corn laws, and thus
began the free trade policy of that country.
Formerly, it was thought that the potato disease was distributed
widely in America, but it is now known to be most prevalent in New
England, in New York and the Canadian provinces, where the potato-
growing industry is an important one. It has a wide range in Europe
and is known throughout Great Britain and from France to Russia,
being especially favored, as it was in 1845, by warm damp weather in
the summer months.
The disease is characterized by leaf spots which first appear at the
margin, or apex of the leaf, and spread over its surface until the
leaf presents a dark somewhat water-soaked appearance. These spots
are brown in drier weather and in all cases a withering of the leaf fol-
lows the attack of the mycelium. The disease is known as dry-rot,
when it develops in the tubers, for the hyphae enter the cells, as haus-
toria kill the cells, and the condition of the tuber known as dry rot
is produced, which may be found especially in the stored tubers.
The hyphae fungus are unicellular and they spread
of the late-blight
through the intercellular spaces of the host sending filamentous haus-
toria into the cells of the leaves, or tubers. From this internal myce-
lium, long branched (dendritic) conidiophores grow out through the

stomata and the branches bear either laterally, or apically, egg-shaped
conidiospores, which measure 2 7 to 30/x by 1 5 to 20/i. The conidiospores
on germination form eight biciliate zoospores, which are motile for a
brief time perhaps not longer than an hour. If one of these swarm
spores finds its way to a leaf, germination speedily follows and the
hyphal germ tube enters the interior of the leaf either through a
stoma, or by boring a hole through the epidermis.
The germ tube of the swarm spores penetrate the tuber, as easily
as the leaf, if they happen to be washed down to the soil. Recently
G. P. Clinton^ has discovered the oogonia, antheridia and oospores of
Phytophthora injestans after they had been sought for by mycologists
since 1845, and thus an American mycologist has added one more
achievement to the list of important work accomplished by American
scientific men.
Spraying the foliage with Bordeaux mixture (5-5-50) has proved
an almost complete remedy against both the Phytophthora blight and
the rot, and also operates beneficially to the potato plant in other ways.
Burying the tubers to a sufficient depth (about 4 to 5 inches) has been
found beneficial, as also the disinfection of the tubers designed for seed
purposes by exposure to dry heat 40°C. (i04°F.) for four hours. Tuber
infection may be prevented by spraying the soil, even when the fungus
is allowed to develop unchecked on the foliage. When the tops are
attacked by late-blight, the harvesting of the tubers should be delayed
until a week or more after the death of the tops. Longer delay does no
harm, unless the season be wet and the soil exceptionally heavy. Dry
cool storage is of primary importance, the use of lime, or formalin, for
disinfection being valueless.^ It seems from investigations, that have
been made, that well-marked and fixed diflferences exist among potato
varieties in relative susceptibility to invasion by the late-blight fungus,
in other words, in disease resistance.
Powdering Dry-rot {Fusarium trichothecioides Wollenw.). —This
fungus kept in artificial culture has been used successfully in the artifi-
cial inoculation of potato tubers, as laboratory exercise with univer-

sity students in mycology. In every case, the rot has been secured and
the students have imbedded pieces of tuber and fungus in paraffin;
cut the same with a rotary microtome and mounted and stained the
sections for microscopic study.
Fusarium trichothecioides forms two kinds of conidiospores: (i) The
comma type, formed as a slightly curved comma ellipsoidally rounded
on both sides; and (2) the normal macroconidiospores. The plecten-
1Clinton, G. P.: Oospores of Potato Blight. Report of the Connecticut
Agricultural Experiment Station, 1909-1910: 753-774 with 3 plates.
J. and Lutman, B. F.:
2 Jones, L. R., Giddings, N. Investigations of the Potato
Fungus, Phytophthora infestans. Bull. 245 U. S. Bureau of Plant Industry,
1912, with full bibliography; Melhus, I. E., Hibernation of Phytophthora infestans
of the Irish Potato. Journ. Agric. Research V: 71-102.
chymatic mycelium and conidial masses are rosy white. The powdery
dry-rot with pink mycelium-Uned cavities is quite characteristic and
not easily confused with the other species of Fusarmm found on
potatoes.^
—
Scab {Actinomyces chromogenes). This scab disease is one well-
known throughout the United States and also in Europe, although
all the cases of scabby potatoes are probably not due to this fungus,
as a causal organism. Turnips, beets and mangels are susceptible
to the disease while carrots and parsnips are not. The first symptoms
of the disease are minute reddish-brown spots on the surface of the
tuber beginning usually at one of the lenticels of the tuber and spread-
ing rapidly to other tissues, assuming a deeper color and an abnormal
corky development over considerable areas. Thus arise the scab-like
crusts which have given the common name to the disease. The surface
of the tuber frequently becomes cracked to considerable depths. If
scabby potatoes are examined immediately after being gathered a fine

grayish, evanescent film will be found consisting of extremely delicate,
minute, refractive, branched filaments, which break up into bacteria-
like cells. Some branches are curved and structures suggesting true
spores are produced in certain cells. The writer has found the fungus
as minute white specks on horse manure. It has been found to
persist in the soil for several years.
The disease can be controlled by soil treatment, by the adoption of
a rational rotation of crops and by planting seed tubers only after they
have been treated for several hours with a solution of i ounce of
formalin to every 2 gallons of water, or by a solution of corrosive
sublimate in water.
Raspberry {Rubus occid entails, L.)
Anthracnose {Glceosporlum vcnetum, Speg.). —As this fungus pro-

duces injuries to the raspberry and blackberry canes, it was called by
Burrill, who published the account of the disease in 1882, the "rasp-
first
berry cane rust." It is known to occur in New Jersey, Illinois, Texas,

Wisconsin, Missouri and other states.
The fungus attacks both fruiting and non-fruiting canes, or suckers,
1 Carpenter, C. W.: Some Potato Tuber-rots caused by Species of Fusarium.

Journal Agricultural Research V: 183-209, Nov. i 1915.
—
producing small purple spots that are variously scattered along the
cane. The spots first formed rapidly increase in size, and as the
fungus develops the center of each becomes grayish-white in color sur-
rounded by a slightly raised, dark-purple border, separating the

healthy from the diseased tissues. The disease progresses in an up-
ward direction and as the advanced stage of the malady is reached,
the spots coalesce. The greatest injury is to the cambium, so that
the living tissues of the cane become sickly, the leaves do not attain
half their normal size, the fruit ripens prematurely, or dries up as
worthless. The petioles of the older leaves may be attacked and later
the veins of the leaves which show whitisii, blister-like spots. The
spots on the lamina are smaller than on the canes.
The mycelium lives in the intercellular spaces of the host, but is
supplied from the neighboring host cells with nutritive materials.

There is at first a slight discoloration of the cell contents, the cells
then lose their shape and finally collapse. The conidiophores are
formed beneath the epidermis of the host and later appear at the
surface bearing the conidiospores, which are surrounded by a gelatinous
substance. Pruning away the diseased canes and burning them in a
brush heap is the most important means of controlUng the raspberry
anthracnose. Spraying early in the season with Bordeaux mixture
(4-4-50) is useful, although not an absolute preventive.
Red Gum {Liquidamhar styraciflua, L.)
Sap-rot {Polystictus versicolor (L.), Fr.). Polystictus versicolor is
one of the most cosmopoHtan species of fungi known. known from It is

Europe, Africa, Australia, South America, Mexico, Japan, the West
Indies and throughout the United States. It grows on the sapwood
of every species of deciduous tree known. It is the most serious of all
the wood-rotting fungi, destroying probably 75 per cent, of the timber
used for railroad ties. A broad sheet of mycelium covers the entire
surface of the timber on which it grows, but it develops in the wood,
especially the sapwood, in which decay takes place with great rapidity.^
There is a rapid solution of the various parts of the woody structure
for the fungus has no preference for either the lignin, or the cellulose
1 Stevens, Neil E.: Polystictus Versicolor as a Wound Parasite of Catalpa.

Mycologia, vi; 263-270, Sept., 1912; see Ante p. 75.
35
parts of the cell wall, and the parts of the springwood fall apart readily,
because of their porous character. The fruiting bodies of this fungus
are extremely variable depending upon the kind of wood on which they
grow. The sessile sporophores may grow singly, or, more usually,
many of them together, forming a series of closely overlapping brackets.
They are readily recognized by the soft, hairy upper surface with bands
of white and yellow color, although these colors are variable. The
young sporophores are fleshy, but become leathery with age. Their
lower surface is white and the pores are minute and regular. Treat-
ment of the wood with chemic preservatives has been found efficacious
in preventing the attack of such fungi as Polystidus versicolor, and
most of our large railroads have machinery where the steeping of
the ties in chemic preservatives can be accomplished quickly and
inexpensively.
Rye (Secale cerale, L.)
—
Ergot (Claviceps purpurea, Tul.) (Figs. 56 and 57), The ergot fun-
gus is found on rye both in America and Europe, where during wet
warm weather it may be extremely prevalent. It gains entrance to the

host at the base of the young ovary penetrating the ovary wall and
gradually replacing the tissues of the rye ovary. This is accompanied
by an enlargement of the ovary which at its upper end presents a some-
what spongy character. This is due to the outgrowth of the mycelium
in the form of twisted strands, the marginal hyphas of which acting
as conidiophores abstrict off conidiospores. This early stage was
known as the Sphacelia stage. Later, as the time for the maturing of
the healthy grains arrives the diseased ovaries will be found to be re-
placed by bluish-black horn-like bodies which project conspicuously
from between the glumes of the rye spikelet. The rye ovary is re-
placed by a hard body with a blackish surface and white interior
known as the sclerotium. The ergot spurs, or sclerotia, perennate as
such until the following spring, when they send up one or several
outgrowths, or stroma, with a knob-like end of a yellowish-brown color.
In the hyphal tissue, which comprises the knob-like portion of the
stroma, fiask-shaped perithecia are formed with short necks and
slightly protruding ostioles. The asci contained in these perithecia
are elongated and contain eight needle-shaped ascospores, which
measure 60 to 70/i in length, and issue from the tip of the ascus by
a small opening. These ascospores bud off conidiospores, which are

capable of infecting the ovaries of rye plants, which have started their
growth toward maturity the following season.
The ergot spurs are used medicinally under police regulations, for
they are dangerous and poisonous. In the Baltic provinces of Germany
and Russia, the peasant class frequently eat bread made out of flour in
which ergot spurs have been ground. They suffer from gangrenous
affections of the extremities with a loss of the hair, teeth and finger-
nails. A nervous form of ergotism has also been prevalent. Cattle
eating ergoted grain show similar gangrenous and nervous symptoms,
the loss of hoofs, tails and horns.
Ergot can be controlled to some extent by the selection of the grain
seed and by removal of all ergoted masses, when detected in the
fields.
A closely related species, Claviceps microcephala (Wallr.), TuL, was

submitted to the writer by the late Dr. Leonard Pearson on red-top
hay, which had been responsible for gangrenous affection of a herd of
cattle in Pennsylvania.
Sweet Pea (Lathyrus odoratus, L.)
Streak {Bacillus lathyri, Manns & Taubenhaus). —This disease had

been noted by the growers of the sweet pea in England, and recently,
it has been detected in the United States.^ Like the bacteriosis of
beans, streak makes its appearance in the season of heavy dew. On
the sweet pea, the disease usually appears just as the plants begin to
blossom; it is manifested by light reddish-brown to dark brown spots
and streaks (the older almost purple) along the stems, having their
origin near the ground, indicating distribution by spattering
rain and
infection through the stomata. The
becomes quickly dis-
disease
tributed over the more mature stems until the cambium and deeper
tissues are destroyed in continuous areas, when the plant prematurely
dies. From the stems the disease spreads to the petiole, peduncles,
flowers and pods with symptoms similar to those on the stems. On the
leaves, however, the disease appears as small roundish spots, which
gradually coalesce, and eventually involve the entire leaf, which when
Taxjbenhaus, J. J.: The Diseases of the Sweet Pea.

^ Bull. 106. Delaware
Agricultural Experiment Station, Nov., 1914.
killed presents a dark-brownish appearance. If the causative organ-

ism, which is a small rod-shaped bacillus, is sprayed upon the sweet
pea plant, the disease makes its ap-
pearance from seven to ten days
after artificial infection and the
symptoms are similar to those pro-
duced in nature. The bacillus is
rarely found in chains and seldom

united in twos or fours. Its fiagella
are not easily demonstrated, as they
are shed so readily that not more
than two to five may be stained
and these are generally quite short.
If properly fixed and stained, very
long delicate flagella may be dem-

onstrated, 8 to 12 in number, and
peritrichous.
Sweet Potato {Ipomoea batatas),

Poir)
Black-rot {SphcEronema fimbriata

(Ell. & Hals.), Sacc.).^We owe our
past knowledge of this disease to
Halsted, who in 1890 described this,
as well, as other diseases of the sweet

potato. It is a seed-bed disease, a
field disease and a storage trouble.
It is characterized by irregular hard,
dark areas, or circular spots, varying
in size from that of a dime to that of
a silver dollar appearing on the skin
of sweet potatoes (Fig. 195). If the
root is injured, the fungus follows

Fig. 195. — Sweet-potato black the line of injury. The sprouts are
rot produced by a fungus, Sphar-
onema fimbriatum. (After Harler, dwarfed and the foliage turns yel-
L. L., U. S. Farmers' Bull. 714, low. The end of the hank is black-
March 11, 1916.)
ened and charred and this is asso-
ciated with a withering of the leaves which become black and crisp.
Frequently, the stems and petioles are affected and black areas appear
on them. In the field the appearance of black girdling lines between
two leaves is an indication of the disease. The part below the black
line remains healthy, while that above wilts and dies. Stem infection
is not always associated with root infections.
The
black-rot parasite lives skin deep on the roots extending only
to the cambial layer, while in infected stems, leaves and rootlets, it
invades all parts. The hyphae are septate and the cells are filled
with oil They are capable of breaking up into as many spores
globules.
as there are cells,and these spores are denominated chlamydospores.
Olive-brown conidiospores are also formed and these are cut off from
terminal, or lateral branches. The pycnidia are formed within the
diseased areas, and they can be had in artificial cultures. They are
flask-shaped with extremely long necks. The pycnospores are more or
less subglobose, or oblong, hyaline and measure 5yu to gn in length. The
mycelium, which has developed to a considerable extent on the root,
may develop sclerotia of a large size by which the fungus perennates,
and it may
also live over on stored roots and pieces of roots left in the
field. Pure cultures of the fungus are not difficult to obtain. It
grows well on any starchy medium, such as sweet and white potato
cylinders and on bean agar. As to the spread of the fungus, various
mites, as well, as watering the plants, help to distribute the pycnospores.
Roots attacked by the black rot fungus have a bitter taste.'
The disease can be controlled by the careful selection of seed
roots and by a judicial rotation of crops.
Sycamore (Platanus occidentaUs, L.)
Blight {Gnomonia venela (Sacc. & Speg.) Kleb.). — Within the last few
years in southeastern Pennsylvania, the sycamore, or plane trees have
been visited in the spring, when the young leaves are about half
developed, by attacks of this fungus, so that the young leaves appear
as if destroyed by early frosts. It is sometimes very disastrous, es-
1 Wilcox, E. Mead: Diseases of Sweet Potatoes in Alabama. Alabama Agric.

Exper. Stat. (Auburn) Bull. 135, June, 1906; Taubenhaus, J. J. and Manns,
Thos. F.: The Diseases of Sweet Potato and Their Control. Delaware Agric. Exper.
Stat. Bull. 109, May, 1915; Taubenhaus, J. J.: The Black Rots of the Sweet
Potato. Phytopathology III: 159-165.
5 so SPECIAL PLANT PATHOLOGY
pecially in low-lying country, as along stream banks, or in closed-in

valleys.Whole trees are practically attacked, the young leaves turn
brown and then they begin to wither and finally curl up into a brittle
mass. It also produces spots on the leaves of the white, black, and
scarlet oaks.
Until the life history of this fungus was fully known, it was con-
sidered as three distinct types of imperfect fungi by the older my-
cologists. The fungus known as Glceosporium nervisequum represents
the stage, which appears upon the leaves in the form of pustules, or
acervuli, especially localized upon the veins of both the upper and
lower leaf surfaces. Ovate conidiospores measuring lo to 15/x X 4 to 6/x
are formed upon small colorless conidiophores.
The acervuli measure 100 to 300/x in diameter and in moist weather
the numberless spores are ejected in creamy masses, or strings. The
same stage was known on the twigs by the generic name of Myxosporium.
The Sporonema stage is represented by the pycnidium, which develops
from the stroma of the fungus and the interior of the pycnidium is
lined by inwardly projecting conidiophores, which abstrict pycnospores.
The ascigeral stage is found on old leaves that have remained over
winter in the open, and it may appear in late winter or early in the
spring. The perithecia are not uniform in size, for we find them
measuring in diameter from 150 to 250/u with a beak 50 to loo/x long.
The broadly clavate asci are bent at right angles near the base. They
have a thickened apex, a terminal pore with a surrounding refractive
ringand bear invariably eight hyaline two-celled elliptic ascospores.
The two ascospore cells are unequal in size, the larger of the two giving
rise to a germ tube.
Application of the 5-5-50 Bordeaux mixture to young shade trees
and to nursery stock assists in controlling the disease.
Tobacco {Nicotiana tabacum, L.)
Root-rot {Thielavia basicola, Zopf). —

This fungus is found on a
great variety of host plants and growth on the roots of tobacco
its
may be taken as illustrative. It is found in the eastern United States

and in Europe from England to Italy. Roots attacked by this fungus
do not develop normally and the roots may be so injured, that if the
plant is pulled out of the soil everything will remain in the soil except
the broken stub of the main root system. Nature attempts to repair
the damage in the tobacco by the formation of a cluster of new roots,
so that affected plants may not be killed, but remain in the stunted
form (Figs. 196 and 197).
The intercellular mycelium is septate, hyaline at first and consists
of narrow hyphae. The fungus produces three kinds of spores, which
197. — Root-rot fungus {Thielavia basicola) in various stages. {After Gilbert,

W. W., Bull. 158, U. S. Bureau of Plant Industry, 1909-)
endospores measure 10 to 20^ by 4 to 5/i. (2) Another kind of spore is the

thick-walled chlamydospore which is cylindric in shape, borne in chains
and measures about 12/x in width. (3) The third kind of spore is the
ascospore, which borne in evanescent asci in simple perithecia. The
is
ascospores are unicellular and measure about 12/iby 5/i.

To check or control the disease sterilization of the soil has been
practised. All diseased roots about the place should be destroyed by
fire.
Timber
Decay (Stereum frustulosum (Pers.), Fr.). — The fruit bodies of this
fungus appear as slightly raised gray spots thickly placed on the

surface of wood and timber (Fig. 85). The fruiting bodies are 2
to 5 mm. in diameter. The action of this fungus on structural wood is
characteristic, as it forms pocket-like areas in the decaying wood,

causing changes in the wood fibers. The holes are more or less len-
ticular and are from each other by the sound wood. Layers of
isolated
white cellulose fiber line the margin of the hole.
Other decay producing fungi are punk fungus, Fomes igniarius
(Figs. ig8, 199, 200) and hedgehog fungus, Hydnum erinaceus (Fig.
201).
Dry-rot {Merulius lacrymans, Schum.). — The dry-rot fungus (Der
Hausschwamm) is one of the best-known and most destructive of wood-
destroying fungi. For many years, it was claimed, that it was purely
domestic found only in connection with the structural wood-work of
houses and buildings, but Hartig drew attention to the fact, that it
probably exists occasionally in a state of nature. Professor von

Tubeuf sums up the evidence of Hartig^ and a number of other observers
in this statement: "Hausschwamm ist bisher ganz auffallend selten,
direkt als botanische Raritat, im Walde gefunden worden. Die
wenigen Funde, welche bis jetzt bekannt wurden, sind nicht etwa in
urwaldahnlichen Forsten gemacht, sondern Nahe der mensch-
in der
lichen Kultur; in solchen Waldern, die in der Nahe grosser Stadte

liegen, oder an Orten in der Nahe von Waldhausem und von Wegen,
1Mez, Dr. Carl: Der Hausschwamm und die ubrigen holzzerstorenden Pilze
Wohnungen, Dresden, 1908, page 260; Moller, Dr. A.: Haus-
der menschlichen
schwamm forschungen im amtUchen Auftrage. Jena, Band 1907; Band ii, 1909; i,
Band iii, 1909.

Fig. 198.— Aspen tree with sevor:il spMi-Miih^ I's (if lutiurs igniarius. {After von
Schrenk, Hermann, Bull. 149, U. S. Bi ail of Plant Industry, 1909.)
zu deren Anlage bearbeitetes Holz verwendet wurde, kann die Mog-

lichkeit der Verschleppung des Hausschwamms den Wald., nicht
in
bestritten werden.'" To this wild form, the name

Meridius Silvester
of
has been given. The domestic form of the fungus Meridius lacry-
mans is an obligate saprophyte. The spores fall upon the exposed end
of a board, beam, joist, rafter, wooden column, or flooring, which may
be'in contact with, or resting on, a stone foundation, brick wall, or
—
Fig. 199. Cross-section of the trunk of a living silver maple rotted by Fames
igniarius. {After von Schrenk, Hermann, Bull. 149, U. S. Bureau of Plant Industry,
pi. a, 1909.)
earth, which is slightly damper, if not in dry weather, then during

rainy, than the more protected part of the same piece of structural
wood. Here the spore germinates and produces a mycelium, which
grows inside the wood from which it abstracts the proteins necessary
for its growth (Figs. 88 and 8g). At the same time, it dissolves the
coniferin and cellulose of the cell-walls, and leaves behind a brown
residue consisting of lignin, tannin and oxalate of lime (Fig. 88)
Fig. 200. —
Cross-section of a living aspen tree iulIlJ by Fames igniarius. {After
von Schrenk, Hermann, Bull. 149, U
S. Bureau of Plant Industry, pi. it, 1909.)
.
Fig 201. — Cioss-scction of a li\ ing wIiul wak tu'o dct.ued b> Hydnum eri-
naceus. {After von Schrenk, Hermann, Bull. 149, U S. Bureau of Plant Industry,
.
pi. vii, 1909.)

So long as sufficient moisture is present, these substances enable the

wood to retain its original volume, but whenever water is withdrawn
the wood becomes traversed by numerous fissures running at right
angles to each other, and frequently, it breaks up into regular cubes
which readily crumble away, if rubbed, or compressed, and a brown
punky substance is the result of the destructive attack of the myceUum.
When the opportunity is presented for the mycelium to develop
vigorously outside the nourishing substratum, it forms especially on the
side of the joist or board, which is facing a moister air-still chamber, as
under a porch floor, or the interior of some conduit (electric or other-
wise), a skin-Hke layer which often attains large proportions. In other

cases, it may fill cracks, or other cavities. If a microscopic examina-
tion is made of the hyphae of the dry-rot fungus, they will be found of
several kinds showing clamp-connections (Schnallenbildungen), the
formation of oidia and the anastomosis of hyphae that come in contact.
The hyphal cells are multi-nucleate. Three kinds of structural hyphae
are discernible, viz., the ordinary thin- walled hyphae, the water-con-
ducting hyphae of larger size and thicker walls, and the sclerenchyma-
like hyphae with very much thicker walls than the other two. The
function of the water-conducting hyphae will be explained, if we examine
the sheet-like mycelia, which cover at times the surface of structural
wood. Such a mycelium will be found covered with drops of extruded
water like tear drops (hence lacrymans > Lat. lacryma, a tear). This
water has been conveyed from the soil, or damp wall, in contact with
the joist, a beam, a distance sometimes of ten or twelve feet to the
drier parts of the wood*. This accounts for the rapid spread of the
mycelium and its abiHty to secure enough water for its insidious growth
through well-seasoned timbers. Sometimes in houses only a thin coat
of paint conceals the destructive work of the "house-fungus." Later the
fruit bodies appear as an extended thin superficial crust of a brownish-
smoke color covered with low anastomosing ridges and wrinkles, sug-
gesting the surface of tripe, over which the hymenial, or basidial, layer
is spread (Fig. 89). The basidiospores are deep yellowish-brown in
color and impart to the hymenium a yellowish-brown hue. Each
basidium terminates in four short sterigma which bear the basidio-
spores, which measure g/j. to i2yu in length by 5.5/x to 6.5^t in breadth.
Germination of the spores is readily obtained.
Kiln drying of structural wood is an excellent means of preventing
— —
the growth of the dry-rot fungus. Coating materials should be avoided

unless thewoods are absolutely dry and the well-seasoned wood should
be painted at once as neglect on this score may cause a lot of trouble.
The walls on which timbers are laid should be perfectly dry.
Sap-rot {Daedalea quercina (L.) Pers). —One of the most im-
portant enemies of structural oak, produces a soft, mushy decay of
the wood (Fig. 202, also page 76).
/mv
m^m.
Fig. 202. Dadalen quercina destroying a fence post, Nantucket, Aug. 23, 1915.
Xerophytic hoof-shaped fruit-body above, mesophytic bracket below in contact
with the grass.
Violet {Viola spp.)
Spot Disease {AUernaria violcB Gall. & Dorsett) (Fig. 203).

The wild violets in the yard of the author have been attacked by
the spot disease every year for the past six years. In some years, the
attack more virulent than in other years. It is also common on vio-
is
lets grown under glass, and in some districts, commercial violet growing
has been practically abandoned.The fungus attacks plants that are
making a rapid and vigorous growth. The first spots are circular,
greenish or yellowish white ones. They have a light colored central
portion surrounded by a narrow ring of discolored tissue, usually
black or very dark brown at first, but changing to a lighter shade, as the
spots grow older. The first diseased part of the leaf looks as if water-
logged, and in a few days, the diseased part of the leaf peripheral to the
central spot fades, or bleaches, to a yellow, or grayish-white. Here
the disease may stop and the plants recover, the diseased areas separate
from the healthy tissue and fall out leaving holes in the leaves. The
disease may spread, however, until the whole leaf is destroyed.
Fig. 203. —
Violet leaves affected with leaf-spot {Allernaria viola). (Photo, by
Heald, F. D. and Wolf, F. A., Bull. 135 (Sci. Ser. 14), Univ. of Te'x.. Nov. 15,
1909.)
The majority of the spots are free from fungous spores except under
conditions favorable to their development. Some spots produce spores
inabundance, especially upon the central, or older portions of the spots.
The spores are borne in chains on dark brownish hyphas that arise
from the diseased surface. The conidiospores are clavately flask-
shaped, muriform, strongly constricted at the septa, which are variable
in number, olivaceous, 10 to 17/i by 40 to 6ofj., exclusive of the isthmus,

which is 3 to 5/i by 3 to 25/^.^
To prevent the disease, only healthy vigorous stock of known par-
entage should be grown. These plants should be propagated at the
season most favorable to the growth of the violet. The frames, glass
houses and conservatories should be kept scrupulously clean.
Wheat {Triticum sativum Lam.)
Black-rust (Puccima graminis, Pers). — Before the rise of modern

scientific investigation in botany, the farmers of Germany believed that
there was some connection between the rusted condition of their wheat
plants and the barberry bushes in proximity to their fields. It re-
mained for de Bary in 1865 to give scientific demonstration of the life

cycle of the rust fungus by experimental methods. He found on the
branches and leaves of the wheat plant rust-red lines, which represent
cracks in the epidermis through which the summer spores known as
uredospores, or urediniospores, project. These together form the ure-
dinial sorus, or uredinium. The spores, as they rise from the inter-
cellularmycelium of the leaf, or stem, are ovate, yellowish-brown, spinu-
loseand measure 10 to iS/xby 20 to 35/i. They may be repeated, as long
as fresh blades and branches are provided for infection and spread
to new parts, but these spores are specialized, as they cannot infect any
other host plant like oat, rye, barley and so forth, but only wheat.
Later the rust-red sori are replaced by brownish-black sori, which repre-
sent the telium composed of teliospores, or teleotospores, which project.
The tehospores are spindle-shaped, two celled, thick-walled and deep
brown in color. They measure 35 to 60/i by 12 to 22^- Germination
consists in the formation of a four-celled promycelium,orbasidium,each
cell of a stalk gives rise to a single sporidium, or basidiospore. These if
blown to the barbery enter the barberry leaf by the formation of a germ
tube and the intercellular mycehum develops a flask-shaped pycnium
(spermogonium) with small, spore-hke bodies abstrictedoff from vertical
hyphae known as spermatia and aecia, or cluster cups on the under leaf
surface, which give rise to seciospores. These carried to the wheat
infect the wheat and the cycle is completed. The aeciospores germi-
nate irregularly and capriciously, the process being accelerated to some
1 DoRSETT, P. H. : Spot Disease of the Violet, Bull. 23, U. S. Division of Vegetable

Physiology and Pathology, 1900.
extent by chilly nights with alternating warm days. Cluster cups that
originate from spores produced on the wheat plant, develop aecio-
spores, which will infect only wheat plants. If it should happen that
these aeciospores are blown to rye, oats, barley and rye, no infection
takes place, so that the same specialization of spores form is noticeable
here as with the uredospores.
In America, the barberry shrubs are extremely rare and to account
for the completion of the life cycle on this side of the Atlantic Ocean,
Fig. 204. —
Germination of the chlamydospores of Tilletia falens several days
after being placed on moist plaster of Paris slabs, c' showing conjugating basidio-
,
spores. {After Bull. 57, Univ. III. Agric. Exper. Slat., March. 1909.)
recourse has been had to amphispores, which are thick-walled stalked

urediniospores produced in the western states under more or less arid
conditions, but Arthur thinks that the perennation of urediniospores
alone is sufficient to explain the recurrence of the disease on the wheat
plant in succeeding years.
It should be emphasized also that within the species of black rust,
there exist several specialized forms, more or less adopted to their own
^^6
host plants or plants. According to Eriksson, six forms can be dis-
tinguished in Sweden, namely, tritici (on wheat seldom on rye, barley

and oat), secalis (on rye, barley and couch grass), avencs on oat, orchard
grass, etc.), pocB (on the blue grasses) oir« or species of

, Aira and Agrostis
on Agrostis canina and A. stolonifera.
Fig. 205. —
Heads of wheat showing smut (Ustilago tritici), and to the right,
appearance of smutted stalks at harvest time. {After Jackson, F. S., Bull. 83, Del.
Coll. Agric. Exper. Slat., December, 1900.)
Stinking-smut {TiUetia fa'tcns (B. & C.) Schrt.). This is the com- —
monest smut on wheat in the United States. It occurs in the
wheat-growing regions of Canada^ and the Northwest, where it
1 Gussow, H. F.: Smut Diseases of Cultivated Plants. Bui. 73, Division of
Botany, Central Experimental Farm, Ottawa, Canada, March, 19 13.
DETAILED ACCOUNT OP SPECIFIC PLANT DISEASES 563
does considerable damage (Fig. 204). The fungus is confined to

the wheat plant, although nearly all the varieties of that cereal are
susceptible to it and under all climatic conditions. The production of
spores in the host is confined largely to the ovules, and as these begin to
grow, they become smutted. Such smutted grains cause a flaring of the
spikelets and diseased parts may be recognized by a slight difference in
color. With the formation of the spores, a penetrating and disagree-
able odor arises, the presence of which gives the common name to the
disease. The smut spores, or chlamydospores, are brown in color,
nearly spheroid in form and vary from 16 to 25/x in diameter. From
these chlamydospores on germination acicular or needle-shaped basidio-
spores (sporidia) arise, which are produced in the form of a crown on a
short basidium (promycelium). The spores may unite in pairs and
secondary basidiospores be formed.
This disease can be controlled by the use of formalin. The grain
of wheat should be sprayed with the solution (i pint to 30 gallons of
water).
Another wheat smut fungus is Ustilago tritici (Fig. 205).
CHAPTER XXXVI
NON-PARASITIC, OR PHYSIOLOGIC PLANT DISEASES
The non-parasitic diseases of plants traceable to the unfavorable

conditions of the slope, physical and chemical character of the soil in-
cluding the deficiency or excess of water content, as well as the unfavor-
able climatic influences, have been discussed at length by Sorauer in
his "Handbuch der Pflanzenkrankheiten" (3d Edition, assisted by

Lindau and Reh, 1908) and the English translation of the 3d edition
of this book by Frances Dorrance under title of " Manual of Plant
Diseases," issued in parts. Four parts have already appeared on Non-
parasitic Diseases. At length also are considered the poisonous in-
fluence of gases and other chemicals together with wound and gall
diseases. Gummosis and several other physiologic diseases have been
described by him. A general treatment of these diseases has been
made in Part II of this book and, therefore, such general considera-
tions need not be rehearsed here. A few specific cases will be given
by way of introducing the student to another phase of phytopatho-
logic work.^
It should be stated at the beginning that no sharp hne can be drawn
between parasitic and non-parasitic diseases. If they were controlled

by a single set of factors this might be done, but complications always
are involved.
The classification, however, is a convenient one and we can, there-
fore, use the terms physiologic and non-parasitic merely as conventional
designations for a certain class of diseases. A convenient bibliography
of non-parasitic diseases of plantsby Cyrus W. Lantz forms part of
Circular No. 183 Agricultural Experiment Station, University of Illinois,
Urbana, May, 191 5. The following are some of the names applied to
such diseases in the original papers listed in the above-mentioned
circular by Lantz: Anaheim, Bitter-pit, Brunissure, Brusone, Chloro-
1 Smith, R. E.: The Investigation of Physiological Plant Diseases. Phyto-

pathology, V, 83-93, Apr., 191 5.
564
NON-PARASITIC, OR PHYSIOLOGIC PLANT DISEASES 565
sis, Collar-blight, Coulure, Court-noue, Curly-top, Die-back, Exan-

thema, Foot-rot, Fruit-spot, Gummosis, Intumescence, Leaf-curl,
Leaf-scorch, Mai di gomma, Melanose, Mosaic, ffidema. Pithiness,
Pourriture, Roncet, Rosette, Scald, Stippen, Sunburn, Tipburn,
Tomosis, Tumor, Water-core, Yellows, Zopal.
The following diseases, selected because of their interest and im-
portance to plant growers, may be looked upon as belonging to this class.
Stag-head, or Top-dry. The disease so designated frequently re-
sults from lack of proper food in the soil. The gradual death of the
top of the tree is an indication of the malady, as well as the loss of
active growth in the lower part of the tree. It is found in forested
areas where by burning, or by denudation, the conditions have been
changed. Stag-head is frequently seen in park trees where the
natural undergrowth has been removed and where the covering of turf
prevents the access of rain to the roots of the trees, or where the stock
of humus has become depleted in the soil. The soil tends to dry out
in summer and in some of the parks in Philadelphia its surface for
several inches becomes baked hard. This is assisted by the constant
tramping of many
feet beneath the trees. The soil becomes impover-
ished, especially in nitrogen and starvation of the tree becomes evident
with the slow death of its terminal branches. As a preventive measure
a constant supply of food should be provided. Wherever practicable
the ground beneath the tree should not be sodded completely, but
should be planted to low-growing shade-enduring plants, and if pos-
sible, the soil should be top-worked and dressed each year with manure,
or other plant food. Along streets and walks this is rendered difficult
by the proximity of paving material, but as in Paris each tree should
have around its base an unpaved area through which the water can
seep into the soil and by which plant food can be added. An open
grating can be placed so as to protect the surface soil about the tree
from the tramping of passersby.
Root Asphyxiation (Suffocation). —The health of trees and other
plants depends on the proper aeration of the soil. This is conditioned
on the sizeand proximity of the soil particles or the amount of water
present, and on the proximity of pavements, fills or grading materials,
etc. The lack of airis of far-reaching importance. The organisms
of nitrification cannot carry on the process of nitrogen fixation in soils
poor in oxygen, and this is true of wet soils or those which are poorly
drained. Flooding of tree roots is frequently the cause of the death

of the tree. This is seen in low places underlaid by a hard pan, where
the groundwater comes close to the surface, or in stiff soils, which
become saturated and hold their water for a long time. Bad aeration
of the soil coupled with the presence of noxious gases is frequently the
cause of disease and death in street planted trees. As preventive meas-
ures the ground should be kept stirred about the bases of the trees, or
where the ground has been filled in around the tree, small patches of
bark should be removed to induce the formation of adventitious roots
from the wounded areas beneath the new soil surface.
Desiccation. —This phenomenon is noticeable in plants exposed to
bright sunlight following a spell of cold or cloudy moist weather. The
young leaves and tender shoots of such plants frequently wither and die
under such conditions. This is sometimes called sun-scald, but evi-
dently it is due to a too rapid loss of water, so that the tender parts
wither. The excessive loss of water is due to the fact that the leaves
produced in very moist air are not adapted to resist excessive transpira-
tion even where there is an abundant supply of water in the soil. In
other words, the leaves and tender shoots have not been sun hardened.
The writer has noticed such a state in the spring when a dry hot spell
of weather succeeds a moist cool spell. This disease is produced in the
West and Southwest by hot dry winds which sweep over the country,
or in South Florida by what are called dry hurricanes. The "Sirocco"
on the African coast of the Mediterranean Sea, in Malta and Italy is a
hot dry desiccating wind, and so is the " Khamsin," a hot wind from the
desert, which blows across Egypt. The leaves of plants are literally
cooked, or parched, with such dry winds. The cold dry winds of
winter may produce the same effects as the warm dry ones.^
Remedial measures under such climatic conditions would be difficult
to operate. Frequently in dry regions the formation of a dust mulch
by cultivating the soil surface is a method of conserving soil moisture, as
is also the application of litter of various kinds. Top pruning in dry
seasons will often check the excessive demand for water and thus pre-
vent injuries to the rest of the tree. Copious watering of the soil
under such dry conditions may save the destruction of the orchard
trees or cultivated plants. Winter blighting, or dry-out of coniferous
' Hartley, Carl and Merrill, T. C: Storm and Drouth Injury to Foliage
of Ornamental Trees. Phytopathology, V, 20-29, Feb., 1915.
trees may be prevented by proper shelter, or by liberal mulching.

Sometimes a light straw shelter, or wind-break, may be efficacious.
Water-logging. —Transpiration from the leaves of plants is much
reduced during periods of long-continued rains or fogs and as a result the
plant becomes gorged with water. Growth is stimulated, but the cells
are thin walled and easily dry up, or are the easy prey of fungi and in-
sects. Such excess of water may result in the formation of little warts
and swellings. These may be formed on leaves or stems. Sometimes
the leaves become diseased by being water-logged in spots which are
translucent in appearance. Galloway and Woods^^ describe the in-
fluence of the excess of water during the season of 1896 in Washington,
D. C. " In early spring vegetation was at first a little retarded by cool
weather, but this was suddenly followed by good growing weather,
during which the leaves of most trees and shrubs especially those of
Norway maples pushed out with great rapidity. This latter period was
followed by one quite dry and warm, during which red spiders increased
to unusual numbers, particularly on the lower and more protected leaves
of the crown. After this came a period of several days of rainy weather,
and many of the spiders were washed off, but the leaves where they had
been working became water-logged. The Norway maples and horse-
chestnuts suffered most, the leaves of these trees in many cases appear-
ing to have been scorched with fire."
Such injuries as water-logging resulting from an excess of moisture
in the air cannot be prevented readily. Proper planting may render
trees less liable to such trouble especially if care is exercised in feeding
them after they are planted. Susceptible trees such as horse-chestnut
and Norway maple require special care and if the conditions under
which these trees can be grown open the way to serious water-logging
they should be discarded and other trees planted in their stead.
—
Qidema of Manihot. The blister-like pustular outgrowths on plants
variously designated as oedemata or intumescences have been the subject
of careful investigation by a number of plant pathologists. The disease
is also known as dropsy^ and has been observed both in greenhouses
and out-of-doors (Fig. 206). The diseased condition known as oedema

or dropsy occurs on stems, leaves and fruits. It has been found recently
^ Galloway, B. T. and Woods, Albert F.: Diseases of Shade and Ornamental
Trees. Yearbook, U. S. Dept. Agric, 1896: 245.
^ SoRAUER, Paul, Lindau, G. and Reh, L.: Manual of Plant Diseases, trans.
by Frances Dorrance, i: 335.
Fig. 206.— CEdema on Manihot (Ceara). A, Normal arrangement of leaf tis-

sues; B, division and enlargement of palisade cells in oedematous leaf; C, division of
cells in the spongy parenchyma which become giant cells; D, early stages of disease
in which all of the cells except lower epidermal ones are oedematous; E, division and
enlargement of cells in lower epidermis; F, cedeinatous leaf tissue double that of
normal leaf; C, shrinking and collapse of cells in oedematous leaf. (After Wolf and
Lloyd, Phytopathology, 2: 134, pi. xi.)
by Wolf and Lloyd affecting the leaves of rubber-producing plants be-

longing to the genus Manihot of which M. glaziovii, M. heptaphylla
and M. pianhyensis are known as ceara. The leaves of the ceara
plants growing in the greenhouses of the Agricultural Experiment
Station, Auburn, Alabama, were found with numerous, glistening,
prominently projecting elevations on either surface of the leaf. When
the elevations or swellings occur on the upper surface there are corre-
sponding depressions or concavities on the lower reaching as much as
three millimeters in diameter and protruding a millimeter above the
surface. The bUsters are circular in outline and mostly isolated, but
if they exceed 300 to 500 they become more or less confluent. At first
there is no change in the color of the leaves, but as the disease
progresses the oedematous tissue turns brown and finally dries and
collapses. The anatomic details of healthy as contrasted with the
diseased oedematous cells are shown in the accompanying details of
Figure 206.
A number of explanations have been given for the origin of oedema,
or dropsy in plants. Giant cells have been found in dropsical tissues
similar to those found in insect galls. Woods found that thin walled
oedematous cells were found puncture by
in carnations as a result of the
aphids, and in such the possible acid conditions must be considered.
Sorauer and also von Schrenk have shown that intumescences may be
caused by spraying leaves with copper salts. Several other plant
pathologists hold to the general view that the disease is due to impaired
transpiration. Sorauer was the first to attribute the cause to abnormal
elevation of temperature, together with excessive water supply. He
finds that weak light or semi-darkness favors the accumulation of water
in the tissues, in that reduced illumination lowers assimilatory activity,
and swollen tissue results. Viala and Pacollet believe that brilliant
light is a prepotent cause, while Fisher argues that oedema is due to
the increased affinity of the colloids of the tissues for water. This may
be due to the accumulation of acids and Wolf -and Lloyd^ believe that
the oedematous tissue of ceara seems to afford some evidence for the
truth of this contention.
Frost Necrosis of Potato Tubers. — Jones and Bailey'' have called at ten-
' Wolf, Frederick, A. and Lloyd, Francis E.: (Edema on Manihot, Phy-
topathology 2: 131-134, pi. I, 191 2.
-Jones, L. R. ant) Bailey, Ernest: Frost Necrosis of Potato Tubers, Phyto-

pathology 7: 71-72, Feb., 191 7.
57© SPECIAL PLANT PATHOLOGY
tion to a type of non-inheritable "net necrosis" of potato tubers which

has developed under conditions which suggest frost injury and this
hypothesis has been confirmed by chilling experiments. Tubers
"frozen solid" are totally killed and collapse when thawed, and if the
chilling stops with incipient ice crystalUzation, such interior tissues as
are most sensitive may be Such frozen tubers are normal in
killed.
external appearance but when cut open they show

that the most
sensitive internal vascular tissues are discolored and are killed. There-
fore, moderate exposure to freezing temperature may produce either
"ring" or "net" necrosis, the blackened vascular tracts penetrating the
fundamental tissue cells filled with starch. Tubers vary individually
in their sensitiveness but in general the best types of "net necrosis"
have been secured by about two hours exposure to + 5°C. with similar
results on exposing them to — i°C. for eight and one-half hours to — 9°
C. for one hour. Slightly more severe treatments, or unequal exposures,
may give frozen spots with corresponding dark blotches involving the
general parenchyma. The stem end of the tuber is always more
sensitive than the other end.
Apple Fruit Spots. —This disease of the fruit of the apple is also
known as Baldwin-spot, bitter-pit, fruit-pit, pointe bruns de la chair
and stippen. It is cosmopolitan in its distribution, being found wher-
ever apples are grown. It has recently received the attention of a
number of mycologists and a number of explanations as to its cause have
been given. The most recent study seems to indicate its non-parasitic
character. The observed spots are dark in color, circular or some-
what angular from one-eighth inch or less to one-fourth inch
in outline,
in diameter. Although distributed over the surface of the pome they
appear most commonly on the blush, or sun-exposed side. A lenticel
forms the center of the sHghtly depressed areas or "pocks," which con-
sist of necrotic tissue. The injury is superficial extending only sHghtly
into the pulp. Pathologists appear to have agreed that the disease is
due to extreme variations in the water-supply of the apple tree during

the growing season.
McAlpine,^ an Australian mycologist, has published four quarto
1 Eastham,
J. W.: Bitter Pit Investigation, Phytopath. 4: 121-123, 1914
Brooks, Charles: Bitter Pit Investigations, Piiytopath. 6: 295-298, 1916
Crabill, C. H. and Thomas, H. E.: Stippen and Spray Injury, Phytopath. 6
51-54, 1916.
1
volumes with plates and which he presents the evidence

illustrations in
in favor of the hypothesis that the stippen is due to irregularities in the
factor influencing the balance between transpiration and water supply

and not to poisoning of cells, e.g., by arsenical sprays as supported by
abundant experimental proofs. He beheves that the principal contrib-
uting factors are:
1. Intermittent weather conditions when the fruit is at a critical
period of growth.
2. Amount and rapidity of transpiration.
3. Sudden checking of the transpiration at night when the roots are
still active owing to the heat of the soil.
4. Failures of supplies at the periphery of the fruit followed by
spasmodic and irregular recovery.
5. Irregularity of growth, so that the vascular network controlling
the distribution of nutritive material is not formed regularly.

6. Fluctuations in temperature when fruit is in store.
7. Nature of the variety.

Water-core of Apple.^ —The diseased fruits are characterized by
hard watery areas in the flesh, usually in the core and extending out-
ward. marked by scattered small spots with
Occasionally the flesh is
extensive watery areas near the surface. The abnormal areas are
usually associated with the vascular tissues. The seed cavities contain
liquid and the hard partition membranes become cracked and covered
with the hair-like out-growth known as tufted carpels. Norton states
that the intercellular spaces so conspicuous in the normal apple
flesh are filled with fluid in the diseased tissue so that the white opaque
appearance of the normal flesh is lacking. "The occurrence of the
disease under conditions favoring excessive sap pressure or cell turgor,
on vigorous growing trees, or trees with the foliage reduced by blight,
and especially in late summer when the air is cold at night and the soil
warm, the cracks in the carpels, the occurrence along the vascular tissue,
the liquid filling the intercellular spaces, lead me to the conclusion that
the trouble due to sap forced into the seed cavities and intercellular
is
spaces by excessive sap pressure under conditions of reduced transpira-

tion. The air being excluded from the inner cells by the liquid filling
the intercellular spaces, anaerobic respiration may be increased and
^ Norton, J. B. S.: Water Core of Apple. Phytopathology i: 126-128, Aug.,
1911.
account for the alcoholic flavor, if not lead to the decrease in acid
and the sweeter taste.
Die-hack or Exanthema of Citrus Fruits.^ — Exanthema is a disease
of the orange groves of the United States occurring in California and
Florida. It affects all varieties of the genus Citrus, both young and
old trees being susceptible. The malady is worse in trees which grow in
poorly drained soils underlaid by an impermeable ferruginous sandstone
but it occurs in hammocks as well. Exanthema attacks the small
branches and. shoots, though the fruit shows symptoms of diagnostic
value. The disease is diagnosed more surely when the shoots become
more or less stained sub-epidermally by a yellowish-brown material
and begin to die back. The fruit may become similarly stained and
its epidermis so dry that it cracks and splits by the pressure of the
developing pulp cells. The disease may be held in abeyance for a

number of years, but if it progresses, the shoots swell at the nodes,
infrequently along the internodes and as they mature, Hnear, erumpent
pustules break out on the internodes. On the older branches the
pustules may be extremely numerous and a small amount of gum may
be observed in them. Proliferation of young buds takes place and these
may develop into short branches with chlorotic foliage producing a
pseudo witches' broom.
Exanthema is induced, like gummosis, by the concurrence of active
growth and active tissues. "The soils in which exanthema occur are
typically dry soils, which when saturated by irrigation water or rains,
promptly become dry once more when the weather clears or irrigation
is discontinued. The rings of growth, which, as we have seen, are very
marked in diseased shoots and branches of trees affected by exanthema,
could not be caused except by a more or less rapid succession of maxima
and minima of growth." Obviously as climatic conditions cannot be
said to be causative, we must look to changes in the water relations of
the "plants which causes a marked development of the rings of growth.
Webber and Swingle have observed that cultivation increases the sus-
ceptibility of the Citrus trees to exanthema, and even causes a
more virulent outbreak of the disease in the affected trees. Any
method of cultivation which tends to promote regular instead of fluctu-
1 Butler, Ormand: A Study on Gummosis of Prunus and Citrus with Obser-
vations on Squamosis and Exanthema of Citrus. Annals of Botany 25: 107-153,
1911.
ating growth may be regarded as a preventive or remedial measure.

Drainage may prove to be remedial to exanthema which is only of one
kind while there may be several kinds of die-back.
Mottle-leaf. — Mottle-leaf of Citrus trees is marked by the loss of
chlorophyll from parts of the leaf, the portions farthest removed from
the midrib and larger veins being first affected. As the disturbance
progresses, the yellowish spots increase in size until the remaining
chlorophyll is found in narrow areas along the midrib and larger veins.
The advanced stages are distinguished by a marked decrease in the
size, quality and yield of fruit. No organism has yet been proved
to be associated with mottle-leaf which is common in the groves
of southern California. Orchards fertilized with organic materials,
such as stable manure, usually showed less mottling than groves the
soils of which were treated with commercial fertilizers. The results
of soil analyses show in the case of oranges a marked inverse correla-

tion between the humous content of the soil and the percentage of
mottling, the latter tending to diminish as the humous content increases
and experiments show that this humus should be well decomposed.
It would seem, therefore, that the mottling of orange leaves in the areas
studied is definitely correlated with the low humous content of the
soil, the mottling diminishing as the humus increases.^
Curly-top of Sugar Beets. '^
—The curly-top of sugar beets seems to
have attracted the attention of growers in California about 1898. It
is distinguished by the following symptoms. An inward curHng of the
leaves, a distortion of the veins of the affected leaves, having roots and
checked growth. It has caused great financial loss in the beet dis-
tricts of the western United States. Experimental study of the disease
shows that the leaves of the curly-top plants have an oxidase content
two or three times as great as the healthy and normally developed
ones. It appears that an abnormal retardation of growth in sugar
beet plants is accompanied by an increase in the concentration of
oxidases in the leaves or a change in the juice of the latter by which
the pyrogallol oxidizing oxidase becomes more active.
Peach Yellows. —This disease which according to the early records
1 Briggs, Lymax J., Jensen, C. A. and McLane, J. W.: Mottle-leaf of Citrus

Trees in Relation to Soil Conditions. Journ. Agric. Res. 6: 721-739, pis. 3, 1916.
BuNZEL, Herbert H.: A Biochemical Study
2 of the Curly-top of Sugar Beets,
Bull. 277, U. S. Bureau of Plant Industry, 1913.
seems to have spread from the region around Philadelphia as a center

has been known about one hundred years. It is a contagious disease of
unknown origin. Erwin F. Smith^ in 1894 gave the first complete
scientific account of yellows founded upon experimental data. He
describes the symptoms as follows: "Prematurely ripe, red-spotted
fruits, and premature unfolding of the leaf buds into slender, pale
shoots, or into branched, broom-like growths. The time of ripen-
ing of premature fruit varies within wide limits; sometimes it pre-
cedes the normal ripening by only a few days, and at other times by
several weeks. The red spots occur in the flesh as well as on the skin,
making the peach more highly colored than is natural. The taste of
of the fruit is generally inferior and often insipid, mawkish, or bitter.
Often this premature ripening is the first symptom of yellows. Often
during the first year of the disease this kind of fruit is restricted to cer-
tain limbs, or even to single twigs, which, however, do not differ in
appearance from other limbs of the tree. The following year, a larger
part of the tree becomes affected and finally the whole of it, the parts
first now showing additional symptoms, if they have not
attacked
already done so. These symptoms are the development of the winter
buds out of their proper season. The buds may rush into shoots only
a few days in advance of the proper time in the spring, or may begin to
grow in early summer, soon after they are formed, and while the leaves
on the parent stem are still bright green. This is a very common and
characteristic symptom, and is especially noticeable in autumn when the
normal foHage has fallen. Usually under the influence of this disease
feeble shoots also appear in considerable numbers on the trunk and main
limbs. These arise from old resting buds, which are buried deep in the
bark and wood and remain dormant in healthy trees. Such shoots are
sometimes unbranched, and nearly colorless, but the majority are green
and repeatedly branched, making a sort of broomlike, erect, pale green,
slender growth, fiUing the interior of the tree."
Yellows can be well controlled by destroying the diseased trees as
soon as they show premature fruit, or shoots with the narrow yellow
leaves. The best treatment is to pull out or grub out and burn the dis-
eased trees, and remove the stumps at a more convenient time. This,
however, does not remove all source of infection as the disease may pos-
sibly spread from the stumps or yellowed shoots arising from them.
1 Smith, E. F.: U. S. Farmers' Bulletin No. 17, 1894.
The next year young trees may be set in the vacant places, care being
taken to obtain trees for resetting that are free from yellows.
Tip-burn of Potato. —This disease is also called leaf burn or scald. It
occurs in many parts of the country and is often confused with early
blight. The and edges of the leaves turn brown and these dis-
tips
colored areas soon become hard and brittle. The burning or scalding
may occur at any time and as a rule is the result of unfavorable con-
ditions surrounding the plant. Long continued cloudy and damp
weather followed by several hot bright days are very apt to result in the
burning of the foliage. This is especially the case on soils carrying a
comparatively small percentage of moisture. When the weather is
cloudy and damp the tissues of the potato become gorged with water and
this has a tendency to weaken them. If the sun appears bright and hot
when the leaves are in this condition there is a rapid evaporation of the
moisture stored up in their cells. The evaporation may be more rapid
than the supply absorbed by the roots, and if this continues for any
length of time the weaker and more tender parts first collapse, then
die, and finally turn brown and dry up. Tip burn may also occur as the
result of protracted dry weather.^
Little of a specific nature can be said as to the treatment of this
trouble. The plants should be kept as vigorous as possible by good
cultivation, with plenty of available food.
Leaf-casting. —The fall of leaves at the end of the growing season, at
the approach of winter, or periodically in the tropics is a normal result
of the formation of an abscission layer. The premature dropping of
leaves, the leaf-fall in house plants, the dropping of flowers and twig
abscission are all manifestation of abnormal, even diseased conditions.
The premature dropping owing to the sudden weakening of
of leaves
functional activities concerns the plant pathologistand is known as
"leaf-casting." The dropping of pine needles is only one phase of the
general phenomenon. I may be allowed to quote here from the English
translation of the third edition of Sorauer's "Manual of Plant Diseases"

(1:349) by Frances Dorrance, concerning the leaf-fall in house plants.
"Among the most delicate of the house plants belong the Azaleas,
because, as a rule, they suddenly drop their leaves in summer, or in the
autumn; the broom-like little tree then at best develops only a few piti-
1 Galloway, B. T.: Potato Diseases and their Treatment. U. S. Farmers'
Bulletin 91, 1899.
f ul flowers. Here too are concerned sharp contrasts occurring suddenly.

Either the plants (usually set in peat soil) in summer are left too dry, and
later watered very abundantly, or they are brought too suddenly into
the warm house
in the autum. In both cases the leaves are weak func-
tionally and then their functioning is increasingly stimulated by the
increased upward pressure of the water. If the transition is brought
about gradually, the inactive leaf Surfaces would have time to resume
their normal action by a general slow increase in their turgidity and
there would be no resultant injury. But, with the sudden upward
pressure of the water, the basal region alone is stimulated, thus causing
the development of the cleavage layer." Here are briefly a few of the
observations of the writer on two plants of Ftuhsia brought into the
house from out of doors and placed in a window with a bright southern
exposure. Soon removal to the house although abundantly
after
watered the leaves began to drop until the window sill was covered with
the litter. New leaves were constantly formed, but these in turn
dropped off and this phenomenon continued through the winter until
the plants were transplanted the following summer to garden soil when
the dropping of the leaves ceased and the plants again became apparently
normal. The general concensus of opinion among plant pathologists is
that the disturbance in the equilibrium of the turgor distribution is the

cause of all premature dropping of the leaves. "For house plants it
may*be recommended as a fundamental principle that the plants should
be subjected gradually to other vegetative conditions, and the dormant
period, upon which every vegetative part enters, should not be inter-
rupted by an increase in the supply of heat and moisture."
—
Curly-dwarf of Potato. This is a peculiar disorder characterized by
a dwarfed development of the potato plant accompanied by a curling
and wrinkling of the foliage, so that it resembles the foliage of the va-
rieties of cabbage known as Scotch Kale and Savoy Cabbage. The
Germans call it Krausel Krankheit. The disease is manifest in the
shortening of the leaf petioles, midribs and veins of the leaves and es-
pecially in the nodes, so that the foliage is clustered thickly. The

diminished growth of the veins in proportion to the cells of the funda-
mental tissue results in a wrinkled leaf surface, often curled downward.
There seems also a tendency for the formation of a greater number of
secondary branches, associated with brittle stems. The color of the
foliage is not altered as it remains a normal green except in very severe
cases, when it becomes a lighter green sometimes with brown or reddish
flecks, where the tissues are dying. This malady is distinguished from
leaf-roll by the bullate, downward curling of the leaves, the persistence
of the normal leaf green and the general firmness of the leaves. It
results in the reduction in the yield of tubers, and in several cases no
tubers have been found.
The nature and cause of this disease remain inexplicable. That it is
an hereditary trouble has been attested by German plant pathologists.
The tubers from diseased hills all develop into curly-dwarfs, while those
from healthy hills remain normal. The disease which is found in
Europe and in this country plays a large role in the deterioration of
potatoes. It seems from our knowledge of the disease that it is a
physiologic disorder resulting in a permanent deterioration of the
potato stock. It may develop at any time under the influence of
conditions not yet fully understood, and the vigor of the strain is reduced
apparently without any chance of its restoration. Perhaps it is concerned
with the senescence of the particular race of potatoes attacked or in
other words a varietal decline.
The disease can be controlled to some extent by selecting tubers
from healthy hills,and if it is prevalent in a field of potatoes, it would
be better not to use any of the tubers from such a field for seeding
purposes.^
—
Bean Mosaic.- Hundreds of acres of pea beans Phaseolus vulgaris in
New York showed the mosaic disease in 191 6 and in some fields prac-
tically every plant was affected and these plants rarely form pods. The
malady is not confined exclusively to the pea beans, but affects varieties
of dry and snap beans and perhaps is the same disease described by Mc-
Clintock as attacking pole and bush lima beans. The leaves of the
plants attacked by mosaic show irregular crinkled areas, somewhat
deeper green than the surrounding yellowish-green tissue. The dis-
ease is transmitted through the seed for diseased seedlings develop

from bean seeds taken from mosaic parents. The disorder has been
induced experimentally by rubbing healthy seedlings with crushed
leaves from diseased plants, the reaction taking place four weeks later.
The first signs of the disease are seen about the time of blossoming.
1 Orton, W. a.: Potato Wilt, Leaf Roll and Related Diseases, Bull. U. S.
Dept. Agric. 64, 19 14.
^ Stewart, U. B. and Reddick, Donald: Bean Mosaic, Phytopathology 61.
7:
37
Experimental treatment indicates that high temperature and humidity

at the time of inoculation favor infection.
—
Mosaic Disease of Tobacco} This disease is one of the most serious
which attacks the tobacco plant. It is known locally as "calico,"
"gray-top," "mottled-top," "mottling" and "foxy" tobacco. The
term "frenching" is used in southern tobacco sections to designate
abnormal, sickly plants with stringy, very thick and leathery leaves
which may be mottled, or not. It is not known whether this disease is
distinct from mosaic. Chlorosis has also been used for mosaic, as well
as the terms" brindle" or "mongrel." Allard states that the mosaic dis-
ease of tobacco is attended with various physiologic and morphologic
changes in the leaves, branches and sometimes flowers of all affected
plants. The character and the intensity of these symptoms vary
greatly, dependingupon the age, habits of growth, species of plants
and external conditions. Allard classifies the characteristic
affected
symptoms of mosaic, as follows:
1. Partial or complete chlorosis.
2. Curling of the leaves.
3. Dwarfing and distortion of the leaves.
4. Blistered or "savoyed " appearance of the leaves.
5. Mottling of the leaves with different shades of green.
6. Dwarfing of the entire plant.
7. Dwarfing and distortion of the blossoms.

8. Blotched or bleached corollas (in Nicotiana tahacum only).
Mosaic sucker growths.
9.
Death of tissues (sometimes very marked in Nicotiana rustica).

10.
The first visible symptom of mosaic in very young plants appears as a
slight downward curling and distortion of the smallest innermost leaves,
which at the same time become more or less chlorotic. Small abnor-
mally dark-green spots and areas appear as these leaves increase in size
and if the plants are not crowded these spots develop rapidly into large,
irregular, crumpled swellings or blisters of a "savoyed" appearance.
The leaves of these young plants may grow to a disproportionate size,
1 F.: Observations on the Mosaic Disease of Tobacco, Bull. 18,

Woods, Albert
U. Bureau of Plant Industry, 1902; Chapman, G. H. Mosaic and Allied Diseases,
S. :
Report of Botanist in 25th Annual Report Massachu êtts Agricultural Experiment

Station, 1913; Allard, H. A.: The Mosaic Disease of Tobacco, Bull. U. S. Depart-
ment Agriculture, 40, 1914.
in some cases becoming long and sinuous. As the plants approach

maturity and become infected they develop into the characteristic
"gray-top" or "mottled-top." The incubation period of 10 or 15 days
is followed especially in the hot sun by a very noticeable wilting of the
upper leaves which become finely mottled. The motthng is due to the
distribution of the dark-green shades along the fine anastomosing veins,
while the lighter shades occupy the small inclosed areas. The roots of
mosaic plants appear superficially quite normal but it is probable they
are impaired, in form and function. It is however in the leaves that the
disease most manifest, which become blotched and mottled accom-
is
panied by distortions which produce at times fantastic leaf forms. The

lamina is suppressed at times so that the leaf is reduced to a twisted
midrib. Sometimes long sinuous ribbon-like leaf blades are found.
The flowers of diseased plants are characterized by the presence of
the normal pink color in lines, specks, or conspicuous blotches, usually
of very irregular distribution. A rather striking and symmetric color
character is the occurrence of the pink color as a fine line in the sinus of
each corolla lobe. Some blossoms are entirely devoid of color and have
a blanched appearance.
Various solanaceous plants are susceptible to the mosaic. Such are
many tomato varieties. Petunia, two distinct garden
species of tobacco,
varieties of Physalis, Datura, Hyoscyamus, Solanum (2 species), and in
several varieties of Capsicum. It is probably distinct from the mosaic
of pokeweed.
The incubation period of the mosaic disease is variable, depending
upon conditions favorable or unfavorable to the growth of the plants.
Eight days is the shortest period recorded. The mosaic virus permeates
all parts of the plant, including the roots and corollas as well as the
foliage, but it does not infect the embryos of seed produced by mosaic
mother plants, and, therefore, such seeds produce healthy plants. The
sap of mosaic plants after passing through a filter still retains its infec-
tious properties and mosaic material ground and dried retained its
virulence one and a half years. The virus preserved by ether, toluene
and glycerin was virulent four months later, as was also the original
juice, which had been allowed to undergo natural fermentation during
that time. Certain species of aphides are active dissemmators of the
mosaic disease.
"Various theories have been advanced to explain the primary origin
of the mosaic disease of tobacco. The view most generally accepted

defines the disease as a disturbance of the enzymatic equilibrium in-
duced by unfavorable conditions of growth. An enzymatic disease is
physiological in its nature, has its origin within the protoplasmic com-
plex, and and sometimes permanent impairment of
results in a serious
the assimilative functions." Although it has been shown by previous
workers that the oxidase and peroxidase content of mosaic leaves is
higher than in normal healthy plants, this fact alone does not warrant,
Allard thinks, its being considered the initial cause of the disease, for it
might well be an effect rather than a cause. It is true that physiologic

symptoms attend the mosaic disease such as chlorosis and various mor-
phologic changes in the leaves, and hence we have placed it among the
physiologic diseases, but notwithstanding, Allard thinks, that parasi-
tism accounts for the primary origin of the disease more consistently
than the enzymatic hypothesis.^
BIBLIOGRAPHY OF NON-PARASITIC DISEASES
A complete bibliography of non-parasitic diseases up to May, 1915, will be found

in Circular 183 Agricultural Experiment Station, University of Illinois by Cyrus W.
Lantz, 81-111.
Âdditional papers on mosaic are, as follows: Gilbert, W. W. Cucumber :
Mosaic Disease, Phytopath. 6: 143-144 with i plate; Doolittle, S. P.: A new

Infectious Mosaic Disease of Cucumber, Phytopath. 6: 145-147; Jaggee, I. C:
Experiments with Cucumber Mosaic Disease, Phytopath. 6: 148-151, iqi6.
PART IV
LABORATORY EXERCISES IN CULTURAL

STUDY OF FUNGI
CHAPTER XXXVII
LABORATORY AND TEACHING METHODS
Introductory Remarks. —-The fourth part of this book is designed
principally to give directions for laboratory exercises in mycology,'
plant pathology and the determination of fungi. The teacher will find
perhaps more than can be covered conveniently in a year's work, unless
the number of hours to be devoted to the study is greater than usual in
college or university work. The instructor will be compelled therefore
to make a selection. There is provided in the fourth part laboratory
exercises in the making of culture media and stains, the methods of
study of bacteria and fungi, the manufacture and use of spray materials
and keys for the identification of different kinds of fungi for use as class
exercises in learning how to identify fungi and in becoming acquainted
with the terms used in systematic mycology. The teacher system-
atically inclined can emphasize the taxonomic exercises provided in the
lessons and appendices. The professor, who wishes to emphasize the
important phases of plant pathology, will find in the fourth part
exercises in the description and study of plant diseases and the
pathogenic organisms concerned in disease production.
The teacher interested in technique will find many lessons which
deal with that subject, as also the apparatus used in the scientific study
of the fungi. The endeavor has been to appeal to a larger circle of
students than those engaged in purely pathologic study. The inquirer,
who wishes to lay a foundation in technical mycology, will find much
along this line in Part IV and the preceding parts of the book. The
teacher, who wishes to acquaint himself with the pedagogic methods,
will find suggestions on this important phase of mycology in the last
part of the text. The mycophagist, who desires to grow mushrooms, will
581
582 LABORATORY EXERCISES
find in detail a method for doing so, and lastly, the practical grower will
find formulse and methods for combating the various fungous and insect
foes which prey upon his crops and which must be subdued or held in
subjection.
LESSON 1
Micrometry.- —The unit of length used in microscopic measurement is the micron

(im) which is the one-thousandth part of a millimeter (o.ooi mm.). There are
four kinds of micrometers in use: the stage, the eyepiece, the step, the filar, or cob-
web, micrometer, and where in modern types, the cobweb is replaced by a finely
spun platinum wire.
—
Method with Stage Micrometer.- The stage micrometer is a slide with a scale
engraved on it divided to hundredths of a millimeter (o.oi mm.) every tenth line
being made longer than the intervening ones, to facilitate counting.

I. Attach a camera lucida to the eyepiece of the microscope.
• 2. Adjust the micrometer on the stage of the microscope and accurately focus
the divisions.
3. Project the scale of the stage micrometer on to a piece of paper and with pen,
or pencil, sketch in the magnified image, each division of which corresponds to lo/x.
Mark on the paper the optic combination (ocular objective and tube length) em-
ployed to produce this particular magnification. Do this for each of the possible
combinations of oculars and objectives, and keep the scales that you have made
for future work in measurement, which is accomplished by projecting the image
of the object on the scale corresponding to the optic combination at use in the
study.
—
Method with Eyepiece Micrometer. The eyepiece micrometer is a circle of glass
with a scale etched on the surface and suitable for insertion inside of the ocular
used during the operation of measurement. The scale is divided to tenths of a
millimeter (o.i mm.) or the entire surface of the glass may be etched with squares
(o.i mm.), the net micrometer.
The value one division of the micrometer scale must be ascertained for each
of
optic combination by the aid of the stage micrometer, thus:
1. Insert the eyepiece micrometer within the tube of the ocular by placing it
on the diaphragm of the ocular, and adjust the stage micrometer by placing it on
the stage of the microscope.
2. Focus the scale of the stage micrometer accurately; the lines of the two
micrometers will appear in the same plane. Make the lines on the two micrometers
to parallel each other.
3. Make two of the lines on the ocular micrometer to coincide with those bound-
ing one division of the stage micrometer; this is effected by increasing or diminish-
ing the tube length; and note the number of included divisions.
;u4. Calculate the value of each division of the eyepiece micrometer in terms of
by means of the following formula: x = icy.
"Where x = the number of included divisions of the eyepiece micrometer.
y = the number of included divisions of the stage micrometer.
LABORATOKY AND TEACHING METHODS 583
Nole the optic combinations used and keep a record of them with the calcu-
5.
lated micrometer value. Repeat for each of the other combinations. To meas-
ure an object by this method, read off the number of divisions of the eyepiece
micrometer it occupies and express the result in microns by looking up the standard
value for the optic combination used.
—
Example. Determine how many of the stage micrometer divisions correspond
with the eyepiece micrometer divisions. Divide the first by the last, the quotient
will be the true value of the ocular micrometer divisions in units of the objective
micrometer. If 20 divisions of the ocular micrometer cover 87 divisions of the
stage micrometer then ^%o = 43-5 = 0.0435 mm.
Method uith Filar Micrometer (Fig. 207). —This consists of an ocular having a
fixed wire stretching horizontally across the field with a vertical reference wire
Fig. 207. — Screw micrometer eyepiece (Filar micrometer).
adjusted at right angles to the first and a fine wire, parallel to the reference wire,
which can be moved across the field by the action of the micrometer screw. The
trap head is di^^ded into 100 parts, which pass successively a fixed index as the head
is turned. A fixed comb with the intervals between its teeth corresponding to one
complete revolution of the screw head is found in the field. As in the previous
method, the value of each division of the comb scale must be found for each optic
combination.
1. Place the filar micrometer and the stage micrometer in their respective
positions.
2. Rotate the screw of the filar micrometer until the movable wire coincides with
the fi.xed one, and the index marks zero on the screw head.
3. Focus the scale of each micrometer accurately and the lines in them parallel.
4. Turnthe micrometer screw until the movable line has traversed one division
of the stage micrometer note the number of complete revolutions (by means of the
recording comb) and the fractions of a revolution (by means of scale on the head
of the micrometer screw) which are required to measure the o.oi mm.
5. Make several estimations and average the results.
6. Note the optic combination employed in this experiment and record it care-
fully, together with the micrometer value in terms of /i.
7. Repeat this process for each of the different optic combinations and record
the results.
To measure an object by this method, simply note the number of revolutions and
fractions of a revolution of the screw, and express the result as microns by reference
to the recorded values for that particular optic combination.
Table of Micrometer Values
Designation of
objective
LABORATORY AND TEACHING METHODS 58s
Table of Micrometer Values. — {Contiuued)

100 intervals of the step
micrometer covers as
Designation of
objective
j
Focal length,
mm.
A/r^,i
th^'e
^t „,\^;û
dtw\ute'has -*—
—/micrometer
ject
^^ °f ^ê ob-
as men-
1
Vto u^ a-.^Zî-Ja
be adjusted I
tioned below, (i interval

equals Moo mm.)
4.2
3-2
30
2.6
2.2
Oil
immersion
K2ff
Oil
immersion
Vie
16 mm.
—
used. The step micrometer has loo intervals distinctly indicated in the middle.
It is necessary to find the number of intervals of the object micrometer covered
by loo intervals of the step micrometer, viz., with objective 3 (16 mm.), at a tube
length of 141 mm., 100 intervals of the step micrometer cover 100 intervals of the
object micrometer, equal to i mm.
One interval of the step micrometer is as i : 100 = o.oi or 10 micra. Micrometer
value = 10. •
With objective 6 (4 mm.) at a tube length of 160 mm. 100 intervals of the
step micrometer cover 20 intervals of the object micrometer =
0.2 mm. One interval of the step micrometer therefore 0.2 = 100
= 0.002 or 2 micra. Micrometer value 2.
This new micrometer eliminates the time-consuming measure-

ment with three or more figures after the old method and is still
more accurate.
—
Comment.- M. Nobert of Griefswald in Prussia engraved lines
more than 100,000 to the space of an inch.
Laboratory Work. —
Compute the various micrometric values
according to the three methods outlined above. After determin-
ing these values for the various combinations of which your
microscope is capable measure the following objects:
Spores of black mould, spores of slime moulds studied, various
diatoms, etc. Practice these methods until you have perfected
yourself in them.
REFERENCES
Beale, Lionel S.: How to Work with the Microscope, 1868 (4th
Edition), pp. 35-38.
Behrens, Julius W., trans, by Rev. A. B. Hervey: The
Microscope in Botany. A Guide for the Microscopical
Investigation of Vegetable Substances, Boston, 1885, pp.
120-133.
DoLLEY, Charles S.: Notes on the Methods Employed in Biolog-
ical Studies, 1889, pp. 18-20.

Fig. 208.
Scale of step Gage, Simon Henry: The Microscope. An Investigation of Micro-
micrometer. scopic Methods and of Histology, 1899, pp. 100-108.
LESSON 2
Directions for Plugging Test-tubes and Flasks. —Before sterilization all test-tubes
and flasks must be carefully plugged with cotton-wool, and for this purpose best
absorbent cotton-wool (preferably that put up in cylindric one-pound cartons and
interleaved with tissue paper) can be used (Fig. 209).
I. For a test-tube or a small flask, tear off a piece of cotton-wool some 10 cm.
ong by 2 cm. wide from the roll.
,
L/VEOJÂTORY AND TEACHING METHODS 587
2. Turn in the ends neatly and rull the strip of wool lightly between the thumb
and fingers of both hands to form a long cjdinder.
3. Double this at the center and introduce the now rounded end into the mouth
of the tube or flask.
4. Now, while supporting the wool between the thumb and fingers of the right
hand, rotate the test-tube between those of the left, and gradually
screw the plug of wool intoits mouth for a distance of about the
r?='*^=^
"'
same length of wool projecting. *
'
The plug must be firm and fit the tube or flask, but not so
tightly that it cannot be removed by a screwing motion when
grasped between the fourth, or third, and fourth fingers and the
palm of the hand.
Rough Method of Cultivating Bacteria and Fungi.— 1. Make
decoctions of split peas, cabbage, lettuce, hay, lima beans, broad
beans and water lily leaves by boiling in water. Expose decoc-
tions to air by placing in an open vessel. This gives the
organisms introduced from the air.
2. Boil a similar lot of material in a glass flask over a water

bath. After material is thoroughly steamed, close opening of the
flask with a cotton plug. Note result.
3. Place untreated in distilled water previously

material
boiled. Plug the flask with cotton. This wiU serve as a control.
This gives the organisms introduced on the material.
Desiderata. —
Flasks, cotton, water bath and Bunsen burner for
these experiments will be found in the Culture Room. Perform
all experiments there.
Other Materials. — Procure a loaf of dry bread, cut it into

slices and place slices on a dinner plate. Wet
bread until well
soaked with water, cover -with a bell jar provided with wet filter
paper. Fig. 209.-
Similarly take horse manure, wet it and place under a bell °^ plugged
^Jl with
jar. Place jars in a dark place Inoculate the following culture potato slant rest-
media with the spores of the various fungi that grow on the bread ing on a bit of
and manure. For this purpose, use a platinum needle sterilized ^lass rod to keep
the potato out of
in the Bunsen flame.
the water in the
Culture of Slime Moulds.- -Compare: The Culture of Did- bottom of the
ymitim xanlhopus (Ditmas) Fr. in Synthetic Media, Science, new tube {After
XL: 791, Nov. 27, 1914. Williams, in
Schneider, Phar-
maceutical Bac-
LESSON 3 teriology, p. 54.)
Microscopic Study of Culture Material. A study is to be made—

of the organisms raised in the culture media prepared as directed in Lesson 2.
—
Hanging-drop Preparation. i. Smear a layer of vaseline (sterile) on the upper
surface of the ring cell of a hanging-drop slide by means of the glass rod provided
with the vaseline bottle, and place slide on a piece of filter paper.
2. Flame a cover-slip and place it on the filter paper on which rests the hanging-
drop slide.
3. Place a drop of water on the center of the cover-glass by means of the platinum
loop.
4. Remove some of the material in the culture flasks by means of a platinum loop
and mix itwith the drop of water on the cover-slip.
5. Raise the cover-glass with the points of a forceps and rapidly invert it on to
the ring cell of the hanging-drop slide, so that the drop of fluid occupies the center
of the ring. (In exact investigation, carefully avoid contact between the drops of
fluid and either the ring cell or the ring of vaseline. Should this happen, the in-
fected hanging-drop slide and its cover-slip must be dropped into lysol solution and
a new preparation made.)
6. Press the cover-slip firmly down into the vaseline on to the top of the ring cell.
This spreads out the vaseline into a thin layer, and besides ensures the adhesion
of the cover-slip seals the cell and almost prevents evaporation.
7. Examine microscopically (vide infra).
Microscopic Examination of the Unstained Material. — i. Place the tube of the
microscope in a vertical position.
2. .Arrange the hanging-drop slide on the microscope stage so that the drop of
fluid is in the optical axis of the instrument, and secure it in the position by means
of the spring clips.
3. Use one-sixth inch objective, rack down the body tube until the front lens of
the objective is almost in contact with the cover-slip.

4. Apply the eye to the eyepiece and adjust the plane mirror to the position
which secures the best illumination.

5. Rack the condenser down slightly and cut down the aperture of the iris
diaphragm so that the light, although even, is dim.

6. Rack up the body tube by means of the coarse adjustment until the organisms
come into view; then focus exactly by means of the fine adjustment.
Some diflficulty is experienced at in finding the hanging-drop, and if the first
first
attempt is unsuccessful, the student must not on any account, while still applying
his eye to the eyepiece, rack the body tube down, for by doing so there is every chance
of breaking the cover-glass and contaminating the objective.
The examination of fresh material in a hanging-drop is directed to the
determination of:
1. The nature of the bacteria and other organisms present.

2. The purity of the culture.
3. The presence or absence of motility.
"When the examination is completed and the specimen finished the slide with
cover slip should in the study of contagious material be dropped into the lysol pot.
Cf. KissKALT, K.: Prakticum der Bakteriologie und Protozoologie, Zweite

Auflage, Erster Teil (Bakteriologie), pp. 10-12 (1909).
Mounting and Staining. —The mounting and staining of bacteria, protozoa and
other microorganisms may
be accomplished as follows:
I. Take the square, or round cover-slip, which has been previously cleaned out
of the alcohol pot, dry it between filter paper.

LABORATORY AND TEACHING METHODS 589
2. Hold it in the bacteriologic forceps which are so constructed that a spring

holds the cover-slip firmly, while an enlargement of the wire handle permits the
placing of the forceps on the table while the culture ma^terial is obtained.
3. Place several drops of distilled water on the cover-slip and add a loopful of
the organisms secured from the culture media as described in this lesson and from
the pure culture in a test-tube as follows:
4. Remove the cotton plug by the third and fourth fingers of the left hand.
Hold the open test-tube between the thumb and forefinger of the left hand.
5.
6. By means of a previously flamed platinum needle remove a little of the
culture from the surface of the culture media.

'/. Replace the cotton plug.
8. Add the culture material to the drop of distilled water on the cover-slip
and distribute this material by stirring.
9. Evaporate the water on the cover-slip to dryness by holding it some distance

above the Bunsen flame and slowly enough to prevent convection circles being formed
by the material affixed to the cover.
10. Pass the cover-glass three times rapidly through the Bunsen flame.
11. Apply the stain, which should remain long enough to stain the objects.
The stains to be used are described in detail below.

12. Wash off the stain with distilled water either from a wash bottle, or from
a bottle suspended some distance above the laboratory table.

13. Dry between filter paper.
14. Apply a drop of balsam, turn the cover-slip over and drop it into the center
of a glass slide previously provided and cleaned for the purpose.
Stains. —
One of the most useful bacteriologic stains is:
Ziehl's Carbol Fuchsin, prepared as follows:
Fuchsin (basic) i
Absolute alcohol 10
Carbolic acid (5 per cent, solution in water) 100
The fuchsin should be dissolved first in the alcohol and then the two fluids
mi.xed.
Loeffler's Alkaline Melhylcne Blue. —
Alcoholic solution of methj-lene blue (saturated) 3c

Caustic potash i \ 100
Distilled water 10,000 /
This fluid retains its valuable properties for a considerable time and is an
excellent stain.
Ehrlich's AniUn-watcr Gentian Violet.
Alcoholic solution of gentian violet (saturated) 5

Anilin water 100
This should be used as soon as prepared. It does not keep well.
1
Rhdich-Weigcrl Anil in Methyl ViolcL
Alcoholic solution of methyl violet (saturated) 1
Absolute alcohol ic
Anilin water i oo
This preparation does not keep well.
Gram's Stain. — This a method of differential bleaching after a stain. The

is
cover-glass preparations, or sections, are passed from absolute alcohol into Ehrlich's
anilin gentian violet, or into a water> solution of methyl violet, where they remain
one to three minutes, except tubercle bacilli preparations, which remain commonly
twelve to twenty-four hours (Gram). They are then placed for one to three minutes
(occasionally five minutes) in iodine potassium iodide water (iodine crystals, potassic
iodide 2 gr., water 300 c.c), with or without first washing lightly in alcohol. In
this way they remain one to three minutes. They are then placed in absolute alcohol
which they are cleared in clove oil and mounted
until sufliciently bleached, after
in Canada balsam. By this method the stain is removed from some kinds of bacteria
and not from others. Too much confidence must not be placed in this method, since
in some cases the removal, or non-removal of the stain from the organism depends
on the length of exposure to iodine water. It would be better, therefore, to expose
all for the same period, e.g., two minutes.
—
DelafieWs Hematoxylin. To 100 c.c. of a saturated solution of ammonia alum
add, drop by drop, a solution of i gram of haematoxylin dissolved in 6 c.c. of absolute
alcohol. Expose to air and light for one week. FUter.
Add 25 c.c. of glycerin and 25 c.c. of methyl alcohol. Allow to stand uhtU the
color is sufficiently dark. Filter, and keep in a tightly stoppered bottle. The
addition of the glycerin and methyl alcohol will precipitate some of the ammonia
alum in the form of small crystals. The last filtering should take place four or five
hours after the addition of the glycerin and methyl alcohol.
The solution should stand for at least two months before it is ready for using.
This "ripening" is brought about by the oxidation of the haematoxylin into haematin,
a reaction which may be secured in a few minutes by a judicious application of per-
oxide of hydrogen (see Chamberlain, Methods in Plant Histology, p. 34).
—
Safranin Gentian Violet. Stain two to three days in safranin (dissolve 0.5 gram
safranin in 50 c.c. absolute alcohol, and after four days add 10 c.c. distilled water);
rinse quickly in water; stain one to three hours in a 2 per cent, aqueous solution
of gentian violet, wash quickly in water. Transfer from stain to absolute alcohol,
clear in clove oil and mount in balsam.
Other useful stains in mycologic work are Fuchsin and Methyl Green, Fuchsin
and Methylene Blue, Eosin Water, Erythrosin and Acid Fuchsin. For the prepara-
tion of these and directions for using consult Chamberlain, Methods in Plant His-
tology, and other books on microscopic technique.
Neisser's -Stain. —
To differentiate between diphtheiia bacilli and pseudo-
diphtheria bacilli.
1. Cultivate the organisms on fresh Loeffier's blood-serum at 34° to 35°C. for

ten to twenty hours.
2. Stain with acid methylene blue three seconds.
I
3. Wash.
4. Stain with Aq. Vesuvin five seconds.
5. Wash.
6. Mount.
Diphtheria bacillus should show the polar granules stained blue and the body
brown. Pseudo-diphtheria show no polar granules.
AuerhacK's Stain. —Auerbach, Leopold: Untersuchungen iiber die Spermato-
genese von Paludina vivipara. Jenaische Zeitschrift fur Naturwissenschaft, 3c:
405-554-
B. Acid fuchsin and Methyl green
Ba. Simultaneous.
I part methyl green \ I.
1000 parts of water

I part acid fucLsin
1000 parts of water.
To so grams of the red solution add i drop of 10 per cent, glacial acetic acid.
Solution I: 3 parts \ ^^.^^

Solution II: acid 2 parts
If necessary to filter, use a filter paper moistened with solution i, as the paper
absorbs the methyl green. Take slides from alcohol and stain slides five to fifteen
minutes, having dried the glass leaving only the sections moist before immersion.
20° to 25° is best temperature; more heat hastens the absorption of methyl green,
cold retards it. Place in absolute alcohol and destain five to fifteen minutes, or
even an hour.
Polychrome Methylene Blue. — See McFarland, Joseph: Pathogenic Bacteria
and Protozoa, 191 2, p. 197.
Toa 0.5 per cent, aqueous solution of sodium bicarbonate add methylene
blue (B X
or "medicinally pure") in the proportion of i gram of the dye to 100 c.c.
of the solution. Heat the mixture in a steam sterilizer at ioo°C. for one full hour
counting the time after the sterilizer has become thoroughly heated. The mixture
is to be contained in a flask of such size and shape, that it forms a layer not more
than 6 cm. deep. After heating, the mixture allowed to cool, placing the flask
is
in cold water, if desired, and is then filtered to remove the precipitate which has
formed in it. It should, when cold, have a deep purple-red color, when viewed in
either layer by transmitting a yellowish artificial light. It does not show this
color, while it is warm. To each 100 c.c. of the filtered mixture, add 500 c.c. of a
cox per aqueous solution of yellowish water soluble eosin and mix thoroughly.
cent,
Collect the abundant precipitate which immediately appears on a filter. When the
precipitant is dry, dissolve it in methylic alcohol (Merck's reagent) in the proportion
of 0.1 grain to 60 c.c. of alcohol. In order to facilitate the-solution, the precipitate
is to be rubbed up with methyl alcohol in a porcelain dish, or mortar with a metal
spatula, or pestle.
This alcoholic solution of the precipitate is the staining fluid. It should be kept
: o
in a well-stoppered bottle, because of the volatility of the alcohol. If it becomes too

concentrated by evaporation, and thus stains too deeply, or forms a precipitate on
the blood smear, the addition of a suitable quantity of methylic alcohol will correct
quickly such fault. It does not undergo any other spontaneous change e.xcept that
of concentration by evapoiation.
Differential Staining tf Fitngcits and Host Cells. — Another useful method is set
forth in the following:
Vaughan, R. E. : A Method for the Differential Staining of Fungous and Host
Cells. Ann. Mo. Bot. Gard., i: 241, 242.
LESSON 4
Liquid Nulrienl Solutions. — Synthetic culture media (see Smith: Bacteria in

Relation to Plant Diseases, i: 197):
Tasteitr's Culture Fluid (Yeasts)
Ammonium tartrate 10 gr.

Ashes of yeast 10
Rock candy 100
Distilled water 1000 c.c.
Dissolve cold.
Naegeli's Nutrient Solution.
Calcium chloride o. i gr.

Magnesium sulphate 0.2
Dipotassium phosphate i .0
Ammonium tartrate 10.
Distilled water 1000. o c.c.
Cohn's Nutrient Solution.
Acid potassium phosphate 5 . o gr.

Magnesium sulphate 50
Neutral ammonium tartrate 10.0
Potassium chloride 0.5
(DeBary, p. 86, Vorles. liber Bakterien, 2 Auflage).
Raulin's Culture Fluid.
Distilled water Magnesium carbonate. . 0.40 gr.

Ammonium sulphate. . . o. 25
Zinc sulphate 0.07
Ferrous sulphate o. 07
Potassium silicate 0.07
. o
5
Ptazmcwski's Culture Fluid.
Dipotassium phosphate 5 . o gr.

Magnesium sulphate 5.0
Ammonium carbonate 5.0
Calcium chloride 0.5
Dissolve cold. Any desired sugar may be added as carbon food.

Adolf Mayer's Culture Fluid (Unters ii-d. ale. Gahr., 1870).
Magnesium sulphate 10. o gr.
Ammonium nitrate 15.0

Tri-basic calcium phosphate o. i
Potassium phosphate 10.

Distilled water , 1000. o c.c.
Dissolve cold and add sugar. Add NaCl (3 per cent.), if it is to be used for
luminous bacteria, and an excess of pure carbonate of hme, if acid-forming bacteria
are to be grown.
Uschinsky's Sclution.
Distilled water 1000 c.c.
Glycerin 30-40 gr
Sodium chloride 5-7
Calcium chloride o. i
Magnesium sulphate 0.3 to 0.4

Dipotassium phosphate 2 o to
. 2 .
Ammonium lactate 6-7

Sodium asparaginate 3-4
Modified Uschinsky's Sclution. — The modified Uschinsky's recommended by

Smith for use with starch jelly is made as follows:
Distilled water 1000.00 c.c.
Ammonium lactate 5 . 00 gr.

Sodium asparaginate ,
2 50.
Sodium sulphate 2.50

Sodium chloride 2 . 50
Dipotassium phosphate 250 .
Calcium chloride o.oi

Magnesium sulphate o.oi
Fraenkel and Voges' Solution.
Water 1000 c.c.

Sodium chloride 5 gr-
Dipotassium phosphate 3
Ammonium lactate 6
Sodium asparaginate 4
38
o
Hygienische Rundschau, Bd. iv, 1894, p. 769.

Fermi's Culture Fluid.

Magnesium sulphate o 2 gr.
Acid potassium phosphate i . o

Ammonium phosphate 10.
Glycerin 450
This may be added to agar in place of peptonized beef-broth (De jSchweinitz)
or to silicate jelly in which case the volume of water must be reduced.
Knop's Solution.
Calcium nitrate (Ca(No3)2, gram i .00 gr.

Calcium chloride (KCl), gram o. 25
Magnesium sulphate (MgS04), gram o. 25
Acid potassium phosphate (KH2PO4), gram o 25
Distilled water, c.c 1000.00 c.c.
Mdisdi's Culture Medium {for luminous bacteria).
Water 1000.00 c.c.
Gelatin 100.00 gr.

Sugar 20 00.
Pepton 10.00
Dipotassium phosphate o. 25
Magnesium sulphate o. 25
Enough sodium hydroxide added to render the medium fully alkaline. On

is
this substratum, the bacteria grow feebly and are not luminous until sodium
chloride, or some equivalent substance, is added (usually 3^ per cent.). Then they
grow well and become luminous.
Leherle-Will Culture Medium {for Yeasts). — See KtJSTER, Ernst: Kultur der
Mikroorganismen, p. 143.
CaHPOi, gram o. 50
K2HPO4, grams 4-55
MgSOi, grams 2 . 10
Pepton, grams 20 00.
Water, liter i . 00
Hansen's Culture Media jor Yeasts.

Per cent. Per cent.
Pepton I Pepton i
Dextrose 5 Maltose 5
Potassium phosphate 0.3 Potassium phosphate 0.3
Magnesium sulphate 0.2 Magnesium sulphate 0.5
Claiibsen's Culture Medium for Pyronema conflucns. — See Kuster, Ernst:

Kultur der Mikroorganismen, p. 152. Claussen places in a Petri dish a small glass
vessel and fills this to the rim with agar of the following formula:
Per cent.
Agar 2 .
000
Inulin puriss 2 . 000
KH2PO4 0.050
NH4NO3 0.050
MgS04 • o 020
Fe3(P04)2 o.ooi
HoO ; 95 000
.
The is filled with inulin-free agar to a similar

outer free margin of the Petri dish
height as in the inner glass dish. In the middle one, spores of Pyronema are sown.
After a few days the fungus will fruit on the inulin-free substratum.
TubeuJ's Culture Medium for Dry-rot Fungus. —
See Kuster, Ernst: Kultur
der Mikroorganismen, p. 154.
Grams
Ammonium nitrate 10
Potassium phosphate 5
Magnesium sulphate i
Lactic acid 2 •
Water 1000 c.c.
Laboratory Work. — Each member of the class should make up at least three of
the above culture media. In order to save material, if the class consists of four to
six students, the full amount of materials can be used and the final amount of liquid
divided into four to six parts for the experiments of each member of the class with
the media made according to the above formulas.
all of Where the class is smaller
than four students, then one-half, or one-fourth of the materials should be used,
as some of them are expensive chemicals.
Inoculate all of the culture solutions with yeast obtained from a cake of Fleish-
man's compressed yeast. and add some of the yeast on the end
Sterilize the needle
of the sterile needle. Study and note the growth of the yeast in the several culture
media inoculated. Bacteria can also be used.
—
Fermenting Power of Different Yeasts. Take a series of fermentation tubes and
fill to the tops of the upright long branch with any of the liquid culture media used
especially for yeasts. Inoculate one with dried yeast, one with brewer's yeast,
one with compressed yeast, one \vith baker's yeast and others with several of the
yeasts kept in pure culture, and plug the open end with cotton. Compare the de-
pression of the upright column of liquid in the different fermentation tubes in order
to determine the relative amount of gas formed.
59^ LABORATORY EXERCISES
Rani his Medium for Moidds.

Grams
Cane sugar 70.00
Tartaric acid 4 00.
Ammonium phosphate o. 600

Magnesium carbonate 0.400
Ammonium sulphate o 250
Zinc sulphate 0.750
Ferrous sulphate 0.075
Potassium silicate 0.070
Water 1500.00 c.'c.
Too complicated to be of much value.
LESSON 5
—
Medium. Whole white potatoes are taken and washed with corrosive
Potatoes as
sublimate i 1000. They are then wrapped in filter paper and steamed in the
:
sterilizer about thirty minutes, the next day twenty minutes, the third fifteen
minutes. The potatoes are then cut in two by a knife heated in a Bunsen flame.
The cut pieces are laid in a large fiat glass dish on a circular piece of filter paper, the
glass dishes having been sterilized by corrosive sublimate. Inoculations are then
made on the surface of the potatoes. This method is especially useful for the
growth of glanders, and chromogenic bacteria.
Potato Juice.
Grated potato, grams 100

Water, c.c 300
Mix and put in ice chest over night; strain off 300 c.c. through a cloth. Cook
for one hour in water bath, filter and add 4 per cent, glycerin. Sterilize. Do not
neutralize as best growth of tubercle bacillus is obtained when the juice is acid.
Growth is rapid and luxuriant, but non-virulent (Archiv fiir Hygiene, XVI). For
culture in tubes with potatoes. Use knife designed by Ravenel, which is used in
the same manner as a cork punch (Fig. 210). The semi-tubular pieces of potato,
punched out, are beveled by a slant cut and placed in a test-tube which is laid
flat with flat side of the potato down to prevent warping; the whole is then sterilized
by the intermittent German process. After sterilization, it is sometimes advisable
to add glycerin soaked in a cotton plug, to the test-tube in order to prevent drying.
A specially designed test-tube (Fig. 211) is used so that the cut piece of potato
can be introduced at the top and the glycerin in the enlarged bottom.
Glycerinated Potato. — i. Prepare ordinary potato wedges.
2. Soak the wedges in a 25 per cent, solution of glycerin for fifteen minutes.
3. Moisten the cotton-wool plugs at the bottom of the potato tubes with a 25
per cent, solution of glycerin instead of plain water.
4. Insert a wedge of potato in each tube and replug the tubes.
5. Sterilize in the steamer at ioo°C. for twenty minutes on each of five consecutive
days.
LABOEATORY AND TEACHING METHODS
597
Glycerin Potato Broth.~i. Take i kilo of potatoes, wash thoroughly in H2O,
peel and grate finely on a bread grater.
2. Weigh the potato gratings, place them in a 2-liter flask, and add distilled
water in the proportion of i c.c. for every gram weight of potato. Allow the flask
to stand in the ice chest for twelve hours.
O^
Fig. 210. —Knife punch designed to Fig. 211. —Culture tube with bulb
cut cylinder of potatoes and other vegeta- to hold glycerine and water below
bles for insertion as slant cylinders in slant of vegetable.
test-tubes as culture media.
3. Strain the mixture through cheese cloth and filter into a graduated cylinder.
Note the amount of the filtrate.
4. Place the filtrate in a flask, add an equal quantity of distilled water, and
heat in a steam sterilizer for an hour.
5. Add glycerin, 4 per cent., mix thoroughly and again filter.
0. Tube and sterilize in the steamer at ioo°C. for twenty minutes on each of
three consecutive days.
—
LESSON (i
Solid Vegetable Subslance (Fig. 210).- —These should consist of slant cylinders
(Fig. 211) in cotton-plugged test-tubes with some distilled water and steamed twenty
minutes at ioo°C. on each of three consecutive days or at the same temperature for
over an hour. Discontinuous sterilization is best. The following are some of the
vegetable substances recommended:
1. Potato 7. Salsify 13. Peanuts

2. Sweet potato 8. Parsnip 14. Brazil nuts
3. Carrot 9. Onion 15. Apple
4. Sugar beet 10. Tulip bulb 16. Pear, or quince
5 Turnip 11. Banana 17. Pineapple
6. Radish 12. Coconut 18. Macaroni
This list may be extended almost indefinitely. The method of preparation of

these solid vegetable substances for the test-tubes is fully described in Lesson 5.
Oat Meal. — Put 10 grams of oatmeal in looo-c.c. Erlenmeyer flask. Add 200
c.c. of distilled water. Stir until thoroughly mixed and autoclave for twenty-five
minutes at i20°C.
Corn Meal.
10 grams + 10 c.c. of water.
10 grams + iS c.c. of water.
10 grams + 20 c.c. of water.
LESSON 7
Plant Juices (With and without the addition of water). Hay Infusion.
1. Weigh out dried hay, 10 grams, chop it up into fine particles and place in a
flask.
2. Add 1000 c.c. distilled water, heated to 7o°C. Close the flask with a solid
rubber stopper.
3. Macerate in a water bath at 6o°C. for three hours.
4. Replace the stopper by a cotton plug, and heat in the Arnold sterilizer at
ioo°C. for an hour.

5". Filter through filter paper.
6. Tube and sterilize in the Arnold sterilizer at ioo°C. for one hour on each of
three consecutive days.
Orange Juice.
1. With a wooden, or metal lemon-squeezer remove the juice from one or several
oranges according to requirements.

2. Filter through ordinary filter paper.
3. Add to the test-tubes provided for the purpose.
4. Plug the test-tubes with cotton.
5. Sterilize on three consecutive days.
Prime Juice.
1. Take a dozen or two of prunes and boil them in water until the water is decid-
edly colored with the prune extract.
2. Add prune juice to test-tubes and plug.
this
3. on three consecutive days.

Sterilize
—
Coconut Water. This is removed directly from the nut to sterile test-tubes by
means of sterile pipettes, which are useful in many ways. The pipettes should be
dry-heated and kept from contamination, or in long, narrow, covered tin boxes.
Wheat Broth (After Eyre and Gasperini).
1. Weigh out and mix wheat flour, 150 grams; magnesium sulphate, 0.5 gram;
potassium nitrate, i gram; glucose, 5 grams.

2. Dissolve the mixture in 1000 c.c. of water heated to ioo°C.
3. Filter through filter paper.

4. Fill test-tubes and sterilize on three consecutive days.
Plant Decoctions, or Infusions in General (After Heald). —Liquid media contain-
ing the soluble nutrients derived from various plant structures are of special value
in dealing with fungi and may be used with bacteria, although they are not so
important for these organisms. By the selection of parts of a host plant for making
a medium for the growth it will be provided with food nearer
of the attacking fungus,
to its immediate needs than from the standard nutrient media. Plant decoctions
may be used as liquid media, or they may serve in combination with other media
solidified by gelatin, or agar.
Some of the most valuable plant decoctions are obtained from fruits, seeds, root
parts and other plant organs. Decoctions may be made from fresh plant parts as
sweet potatoes, beets, turnips, carrots, celery, bean pods, plums, apples, etc., or
dried plants such as dried apples, dates, beans, leaves, etc.
In preparing either decoctions, or infusions, it is well to h'ave the parts employed
in a finely divided state. The parts may be run through a food chopper or ground
finely by a small coffee mill. The pharmaceutic standard should be selected for
decoctions and infusions, i.e. 1000 c.c. should contain the soluble constituents of
50 grams of dry weight of the product employed. To secure uniformity of compo-
can be used in determining the weight of the fresh product
sition the following table
to be employed.
Table to Determine Amount or Dry Substance to be Used
Water
Name of plant organ content,
Potato !
Sugar beet
Carrot
Celery
Leaves (young
Leaves (mature)
Bark (fresh)
Bark (air dry)
6oO LABOKATORY EXERCISES
Directions for Making Plant Infusion.

1. Add looo c.c. of boiling distilled water to 50 grams dry weight of the sub-
stance of the equivalent, chopped or ground fine.
2. Macerate in a closed vessel for thirty minutes.
3. Strain through cheese cloth or filter as for other media and pass distilled water
through the filter to make 1000 c.c. If a clear medium is desired the white of an
egg may be added:

Directions for Making a Plant Decoction.
1. Add 1000 c.c. of cold distilled water to 50 grams dry weight of the substance,
or the equivalent, chopped or ground fine.
2. Heat in a cooker over a gas burner and boil for fifteen minutes, stirring suffi-
ciently to prevent burning.

3. Filter as for infusion and clear, if desirable. Decoctions are preferable to
infusions since there will be a somewhat more complete extraction of the nutrients.
Laboratory Study.— In the use of the culture fluids observe the rapidity, density
and persistency of the growth. Record the formation of acids, alkalis, odors, gas
bubbles, stains, etc.
LESSON 8
Milk. —Nearly all bacteria Ordinary cow's milk is used. The

grow in milk.
cream is separated off Ordinary milk as sold is contami-

and the skim milk used.
nated with fecal bacteria, those found in cow's dung and around stables. Conse-
quently the milk before it is used must be thoroughly sterilized. It may be used
in this form, or a tincture of blue litmus is added until a pale blue color is obtained.
Different organisms react differently with this milk; some render the litmus more
deeply blue, others are indifferent, some give an acid reaction.
The milk should not be acid to taste and should not contain formaldehyd, or
other antiseptic substance which milk dealers sometimes add to milk to improve its
keeping qualities. It should be steamed in wire-crates fifteen minutes at ioo°C.
on each of four consecutive days (loo-c.c. portions in test-tubes) and should not be
used until at least a week after the last steaming. Such milk should be kept under
observation at least si.x or eight weeks.
Litmus milk prepared from fresh milk which has been passed through a separa-
is
tor (centrifuge) or from milk which has stood eighteen or twenty hours at 2o°C. and
has had the cream removed by skimming. To each 100 c.c. of this milk is added
2 c.c. of a saturated solution of high-grade lime-free blue litmus (litmus i gram, water
15 c.c). This gives a lavender color of just the right degree, which reddens distinctly
under the action of acids and blues with the development of alkalis. After adding
the litmus water, the milk should be pipetted in lo-c.c. portions into cotton-plugged
test-tubes and heated as directed above. This is a very useful medium.
Litmus Whey (After Eyre).
1. Curdle fresh milk by adding rennet (or by acidifying with hydrochloric acid).
2. Filter off the whey into a sterile flask.
3. Heat in the Arnold sterilizer for one hour.

4. Filter into a sterile flask.
LABORATORY AND TEACHING METHODS 6oi
5. Tint the whey with litmus solution to a deep purple red.

6. Tube, and sterilize as for milk.
Laboratory Study. —
Milk offered for sale in cities is frequently more than forty-
eight hours old and often contains 3,000,000 to 6,000,000 bacteria per cubic centi-
meter. Such milk is not fit for laboratory use.
Observe in particular:
(o) The separation of the casein without the development of any acid, indicating
the presence of lab, or rennet, ferment. The milk
usually becomes more alkaline.
{b) Saponification of the fat. The becomes transparent without any pre-
fluid
cipitation of the casein; but the caseinogen may be thrown down subsequently by
acidifying the clear liquid.
(f) Ropiness. The liquid becomes viscid and strings when touched.
(</) Formation of acids.
(f) Resolution of precipitated casein (trypsin ferment); formation of crystals,
tyrosin, leucin, etc.
{{) Milk alkaline.
Gelatinization of old cultures.
{g)Changes in smell, color, taste.
Beerwort. —
Beerwort obtained from the brewery is put in test-tubes with cotton
plugs. These test-tubes are then sterilized by discontinuous sterilization and then
inoculated. It is a useful medium for the culture of yeasts.
Beerwort may be added to agar, or ia the cultivation of moulds for class study
it may be used to soak bread or other material on which the moulds are to be
cultivated.
LESSON 9
Bouillon. — Bouillon forms the nutrient basis for culture media. It is made up
in the following proportions, a certain amount of water being used: i per cent, pep-
tone, .5 per cent. NaCl and .5 per cent, beef extract are added and the liquid
boiled. Thus for r liter of H.O
10 grams peptone
]
5 grams salt \ are added.

5 grams beef extract J
This solution has a slight acid reaction and is neutralized by 10 per cent. NaOH
until it is no longer acid but is still acid to phenolphtalein. Bouillon

to blue litmus,
is used either alone or with other media in combination.
Fresh Bouillon. —
Prepared by digesting fresh veal (3 pounds) in water over night.
This mass is then pressed until the water and dissolved juice are separated from the
meat fiber. After filtration, the liquid is brought to a boil and a coagulation of the
albuminoids present takes place. The liquid is again filtered and is found to be
decidedly acid. In one case 3 pounds of veal made 2800 c.c. of
it was found that
liquid beef tea, or meat
which consists essentially of the salts of the meat.
extract,
To make the regulation bouillon, to this liquid must be added salt and peptone
according to the following proportion. Cllycerin may be added for the growth of the
tubercle bacillus.
,
6o2 LABORATORY EXERCISES
2800 c.c. Water extract beef

38 grams Peptone (Bouillon alone)
14 grams Salt
[40 c.c. Glycerin
After boiling, to this be added enough of 10 per cent. NaOH to neutralize

is to
the acidity of the meat extract. It must be neutralized until it is alkaline to blue
litmus and acid to phenolpthalein. It is again filtered and is ready for use. Tu-
bercle bacilli grow exceptionally well in this second solution.
Fresh Bouillon (Another formula). —
Standard peptonized beef bouillon is made
as follows: To 500 grams of finely minced lean beef add 1000 c.c. of distilled water.
-Diagram illustrating construction and action of Arnold steam sterilizer.

(Fig. 15, p. 39, Schneider, Pharmaceutical Bacteriology, 1912.)
The soluble parts may

be removed from the meat by allowing the water to stand
on it one hour in the water bath at 55°C.
for twenty-four hours in the ice chest or for
The second method is perhaps preferable. Then boil for sixty minutes either in the
steamer, or in a covered dish. Filter through clean cloth, using pressure (meat press)
cool, and remove fat by filtering through filter paper; make up to 1000 c.c. by addi-
tion of more water; then add i per cent. Witte's peptonum siccum and 0.5 per cent,
c.p. sodium chloride. Steam one-half hour, filter, cool, titrate, add required alkali,
steam again for one-half hour, filter pipette into test-tubes or flasks, and autoclave or
heat for a minimum time in the Arnold
sterilizer (Fig. 212). Plugs should be well
made and fit be scrupulously clean. For some purposes
tightly; glassware should
both the peptone and the salt should be omitted. A greenish bouillon indicates
insufficient boiling, and will usually throw down some additional vexatious pre-
—
cipitatewhen heated in the test-tubes. Other meats may be substituted for beef,
and other peptones for Witte.
Glycerin Bouillon.
1. Measure out nutrient bouillon, looo c.c.
2. Measure out glycerin, 60 c.c. ( = 6 per cent.) and add to the bouillon.
3. Tube and sterilize as for bouillon.
Sugar Bouillon.
Measure out nutrient bouillon, 1000 c.c.
1.
Weigh out glucose, 20 grams ( = 2 per cent.) and dissolve in the fluid.
2.
3. Tube and sterilize as for bouillon. Ordinary commercial glucose serves the
purpose equally well, but it is not recommended, as during the process of steriliza-
tion the medium gradually deepens in color. In certain cases a corresponding per-
centage of lactose, maltose, or saccharose, is substituted for glucose.
LESSON 10
—
Egg Albumen. Absolutely fresh eggs should be used. The end of the egg from
which the albumen is poured must be thoroughly flamed before it is broken, and care
must be used in the transfer to test-tubes, so as to exclude air-borne germs; other-
wise, the sterilization will be diiificult. By being placed in a steam sterilizer and
sterilized by intermittent sterilization for three days, care being taken to leave off the
cover of the sterilizer (Fig. 211), the albumen will be found to be white, quite hard
and ready for use. If the lid of the sterilizer is kept on and the heat becomes too
great, bubbles will form in the albumen and thus spoil its usefulness. The albumen
of eggs may be cut with sterile scissors.
Egg Albumen (After Eyre, The Elements of Bacteriological Technique, 1902:
160).
1. Break several fresh eggs (hens', ducks', or turkeys' eggs) and collect the
"whites" in a graduated cylinder, taking care to avoid admi.xture with 3'olks.

2. Add 40 per cent, distilled water, and incorporate the mixture thoroughly by
aid of an egg whisk.

3. Weigh out 0.15 per cent, sodium hydrate and dissolve it in the fluid (or add
the amount of decanormal caustic soda solution (see infra) calculated to yield the
required percentage of soda in the total bulk of the fluid i.e., 0.375 c.c. of deca-
normal NaOH solution per 100 c.c. of the mixture.
3a. Glucose to the extent of i or 2 per cent, may now be aaded, if desired.
4. Strain the mixture through butter muslin and filter through a porcelain
filter candle into a sterile filter flask.
5. Tube, and stiffen at ioo°C. in the serum inspissator, or in the steam sterilizer
with the lid off.
•
6. Incubate at 37°C. for forty-eight hours and eliminate any contaminated tubes;
store the remainder for future use.
Egg Yolk. —This is poured into test-tubes and solidified in a slanting position bj'
8o°C. heat, or the egg may be boiled hard and the yolk cut with a sharp knife and
transferred to sterile petri dishes. If desired the yolk and white may be mixed
before solidifying, i.e. by shaking the egg vigorously before breaking the shell.
Solidified Blood-scrum. — As this medium is rather difficult to obtain and prepare,

being one of the most difficult to make in culture work, the plant mycologist must
in general obtain blood-serum from the animal bacteriologist. Near Philadelphia it
can be purchased from the laboratory of H. K. Mulford & Co., Glenolden, Pa. The
solidified serum may be used either plain or with the addition of grape sugar.
Fresh Serum How Obtained. — Procure blood by a sterile method from a horse,
or a cow, and stand it aside in a cool place, breaking the clot from the side of the jar
until the amber-colored serum rises to the surface, when it is to be drawn off. It
is then filtered and measured off. To 3 parts of this serum, i part of bouillon, pre-
pared in the ordinary way, is to be added. The mixture of bouillon and serum is
then to be filled into the sterile test-tubes, care being taken to slant the tubes. By
being placed in a steam sterilizer and sterilized by intermittent sterilization for three
days, care being taken to leave off the cover of the sterilizer, the serum will be found
to be white and quite hard and ready for use. If the lid of the sterilizer is kept on
and the heat becomes too great, bubbles will form in the serum and thus spoil the
usefulness of the hardened serum.
Before filling the tubes, care must be taken that the mixed serum and bouillon
are thoroughly neutralized by NaOH. As blood-serum is rarely used in mycologic
work, the above notes are given merely for reference. The teacher will probably
find it convenient to omit this part of Lesson 10 entirely.
LESSON 11
Nutrient Gelatin.- —To 1000 c.c. add 100 grams

of sterile peptonized beef-bouillon
of best quality gelatin. Soak two hours at room temperature, then steam five
minutes, cool, titrate, add the necessary alkali, steam thirty minutes, filter through
filter paper, wash with sterile boiling hot water, tube at once, and heat in the steamer
on three successive days fifteen minutes, ten minutes and five minutes respectively
at ioo°C. Do not autoclave, and carefully avoid long heating in the steamer.
Have all glassware sterile, the fluids sterile and sufficiently boiled to begin with.
The very best English, French or German gelatins should be used. +10 or +15 is
a good degree of alkalinity for many purposes.
Sugar Gelatin.
Water, c.c 600

Peptone, grams 6
Salt, grams 3
Beef extract, grams 3
Glucose, grams 6
Gelatin, grams 60
The added as the mixture in water is brought to a boil. The mixture

gelatin is
is cooled down below the coagulating point of albumen and the white of two
to 60°
eggs for every ioqo c.c. of water added. It is then brought to a boil, the albumen
coagulates and clarifies the medium. The fluid is then filtered through filter paper
previously wetted with boiling water. A funnel with wire support for filter paper is
to be preferred for ease in filtering.

—
Sugar Gelatin (Another formula). Prepare nutrient gelatin and weigh out
glucose 20 grams ( = 2 per cent.) and dissolve in the hot gelatin. Filter, tube and
sterilize as for nutrient gelatin. In certain cases, lactose, maltose or saccharose in
similar percentages is substituted for glucose.
Litmus Gelatin. — Prepare nutrient gelatin, add sterile litmus solution, sufficient
to tint the medium a deep lavender color, tube and sterilize as for nutrient gelatin.
LESSON 12
Agar-agar. —To make i liter of agar-agar take
A. Dried peptone (i per cent.), grams. . 10

Common salt (0.5 per cent.), grams. .
5
Liebig e.xtract (0.5 per cent.), grams. 5
Water, c.c 500

Boil for three minutes and neutralize.
B. '
Agar-agar ( 1 . 2 percent.), (in shreds, or as
flour) grams 12
Water, c.c 500
Chop the agar and put into autoclave (Fig.

213). Run autoclave up to two atmospheres of
pressure, giving i2i.4°C. of heat. As soon as
this pressure is reached, turn out the flame, and
allow the autoclave to cool until below ioo°C. be-
fore opening. The two solutions A and B are
then mixed, cooled to 6o°C., the whites of two
eggs beaten in 50 c.c. of water added, well stirred
and the whole then boiled, the solidified albumen
in,
and precipitate skimmed ofT and the residue filtered

through paper. Fig. 213. — Usual form of
The whole process requires only an hour and laboratory autoclave for sterili-
a quarter to an hour and a half, and the result zation with steam under pres-
sure. (Fig. 16, p. 40, Schneider,
is a most excellent jelly. Instead of .the white of
Pharmaceutical Bacteriology,
egg, blood-serum may be used, which seems to 1912.)
add also to the nutritive value of the medium.
Agar made with meat extract will often form a precipitate during the sterilization,
which is objectionable if one wishes to use it in the pouring of Petri dishes, or the
making of Esmarch's roll-tubes.
Agar with Fresh Meat.— To make an absolutely and permanently clear agar,
fresh meat should be used as follows:
To make i liter' take:
A. Chopped meat, grams 500

Water, c.c 5^*°
Mix and place in cool place over night, then strain through towel.
B. Agar-agar (1.2 per cent.), grams 12
Water, c.c 5°°
Put run up to two atmospheres of pressure, put out flame, and allow
in autoclave,
to cool until below ioo°C. before opening (Fig. 213). Let the solution of agar cool
still further to about 7S°C., and then mix A and B, add (i per cent.) 10 grams dried
peptone and (0.5 per cent.) 5 grams common salt, bring to a boil for about three
minutes, neutralize and filter. The product is an absolutely clear jelly, which
never forms any precipitate. The whole process, with the exception of the time
the meat is about one hour and a half. In both the above
steeping, requires only
methods of making is very quick from ten to twelve minutes
agar, the filtration —
for the liter. It is not necessary to use a hot-water funnel, but wet the filter paper
with boiling water immediately before pouring in the agar. In the process with
fresh meat the clarification is effected by the coagulation of the albumen in the
meat water, hence solution B must not be added to A until cool enough to avoid
coagulation. In general the fresh meat is to be recommended, and the process is
easier than with the meat extract, though the latter has the advantage of cheap-
ness and convenience, since the meat extract can always be kept on hand, and the
time lost in soaking the fresh meat is saved.
Methods of Inoculations of Agar-agar. —Agar is stored in test-tubes in one of two
waj's, viz.: as a straight, or cylindric medium; or, as an oblique, or slanted medium.
1. Oblique or slanted medium. Here the medium has been allowed to solidify
with the tube in an inclined position, thus forming a flat surface extending nearly
to the mouth of the tube. Such slanted agar is used for "streak" (Strich cultur),
or "smear" cultivations.
2. Straight, or cylindric, medium. Here the medium forms a cylindric mass in
the lower part of the test-tube and the upper surface is at right angles to the long
axis of the tube. Such a cylindric medium is suitable for stab culture when the
platinum needle is thrust deeply into the substance of the medium with the needle
held vertically.
LESSON 13
Various Nutrient A gars. — In addition to beef bouillon, or in place of it, various

substances organic and inorganic may be added to the agar with advantage.
Litmus lactose agar is made out of ordinary nutrient agar by adding milk sugar
and enough pure litmus to give the tests. To 1000 c.c. of ordinary agar, preferably
that made from bouillon free from muscle sugar, add 10 grams of c.p. lactose and
20 c.c. of a saturated (water) solution of c.p. (lime-free) blue litmus.
Glycerin agar, maltose agar may be made with any amount of the substance
desired, generally i or 2 per cent. (1000 c.c. agar plus 50 c.c. Schering's c.p. glyc-erin).
Beerivort agar is conveniently made in i or 2 per cent, combinations of beerwort
and ordinary agar. Take a measured quantity of agar by volume and after it is
liquefied in the steam sterilizer add enough beerwort by volume to make a i or 2 per
cent, quantity of that liquid.
Glucose agar is a useful culture medium. Take i or 2 per cent, of glucose by
weight (i gram = i c.c. by volume) and add to a measured volume of agar in the
liquid form.
Dextrose Agar.
Dextrose, grams 10
Agar, grams 15
Water, c.c 500
Nutrient solution (same as for cellulose agar) c.c 500
Hesse and Niedners Nutrient Agar for Water Bacteria (Smith, p. 196).
Distilled water, c.c 980.

Nahrstoff Heyden, an albumose, grams 7.5
Agar-agar, grams 12.5
This agar is said to be the most suitable medium for the bacteriologic examina-
tion of water. It gives a much larger number of colonies than ordinary agar. It
requires no neutralizing. The poured plates are counted according to Dr. Robin
on the ninth and tenth day. Chromogens are brilliantly colored (Zeitschr. fur
Hygiene, Bd. XXIX: 454-462; see also Amer. Journ. Pharm., LXXVI: 112).
:4. — Manner of holding test-tubes in making subcultures. {After Williams

in Schneider, Pharmaceutical Bacteriology, p. 54.)
Prune Agar (C. S. Shear and N. E. Stevens, Cultural Character of the Chest-
nut Blight Fungus and Its Near Relatives. Circular No. 131, U. S. Bureau of
Plant Industry). —Take four ordinary prunes and add i liter of water. Boil over
an open flame for one hour, being careful not to break the skin of the prunes. Strain
through gauze, make up to the original amount with distilled water and add 2 per
cent, of agar. Steam for three-quarters of an hour, filter and tube. Autoclave for
fifteen minutes at ii5°C. (Fig. 213).
Media for Mine Fungi (Dr. Caroline Rumbold).

1. Pure gelatin 10 per cent., 20 per cent. Bausch and Lomb imported seal gelatin.
2. 6 per cent, gelatin, 2.5 per cent. Liebig's extract, i per cent, citric acid. Cox's
gelatin can also be used. This was more successful than the golden seal gelatin.
This with i per cent, citric acid solidified.
Laboratory Work. —Inoculate any or all of the several nutrient agars with several
of the stock cultures of fungi. Note the rate of growth and dififerential character of
the growth on the different media (Fig 214).
LESSON 14
General Directions for Making Plant A gars. — Plant agars of various kinds may
be made by substituting the desired decoction (made as directed later) for the
bouillon and each looo c.c. of agar should contain the soluble nutrients from 50
grams of dry weight of the plant structure used.
Decoctions (F. D. Heald) are made by adding 1000 c.c. of cold distilled water to
50 grams dry weight of the substance. Heat in a steam sterilizer and boil for
fifteen minutes. The following data are applicable in this connection.
Table of Dry Contents
Water Dry Approximate weight

content, substance, giving so grams of dry
per cent. per cent. substance, grams
Potato 25
Sugar beet
Carrot
Celery
Corn meal.
LESSON 15
Potato Juice Agar (150 + 10 + 500). —Take 150 grams of peeled potato, slice
it thin, soak it in tepid water and allow it to simmer for half an hour. The juice
is used from this in place of bouillon in making the agar-agar.
Potato Agar. —
Put clean pared potatoes rapidly through a grater and immedi-
ately throw into the required quantity of distilled water, which should be used in
ratio of 2 c.c. of water to i gram of the potato. Then put in the Arnold sterilizer.
Soak the agar in water (i gram of agar to 100 c.c. of water), add to the potato and
mix thoroughly (Washington formula).
Fig. 215. — Square form ut Arnold steam st , showing two front doors as
Pharma-
recommended by the Boston Board of Health. (Fi^ 17, p. 42, Schneider,
cetitical Bacteriology, 191 2.)
Mel — Cook says 500 grams 500

T. Cook's Formula. water, 10 grams
in c.c. of of
agar in 500 c.c. of water.

Dr. Caroline Rumbold's Formula. — The freshly grated potato, 500 grams 500 in
c.c. of water, is put Arnold steam sterilizer and heated up to 90°C. Part of
in the
the pulp is strained through cheese cloth. 7.5 grams of agar are soaked in 500 c.c.
of distilled water and before the agar has dissolved, it is put into the potato, and
the whole thorughly mixed. It is then steamed by discontinuous sterilization
(Fig. 215).
McBetk and Scales Formula (McBeth, I. G. and Scales, F. M.: The Destruction
39
'6 10 LABORATORY EXERCISES
of Cellulose by Bacteria and Fungi. Bull. 266, Bureau of Plant Industry, 1913: 28).
— Pare, steam and mash a quantity of potatoes. To 100 grams of mashed potato
add 800 c.c. of tap water ar.d steam for one-half hour; filter through cotton.
Potato solution, c.c 500

Agar, grrims 15
Nutrient solu tion, c.c 500
Potato Agar (AnCither formula). —

Put clean pared potatoes through a meat
gB-inder. To
1000 gn ims'of the potato pulp add an equal quantity of distilled water.
Stir thoroughly aJid let stand in an ice box for an hour, with occasional stirring.
Strain through gauz€ of medium mesh. Make up to three times the weight of the
original pulp with di stilled water. Strain for one hour, filter through cotton and
paper and make up to 3000 c.c. with distilled water. Add iî, per cent, of agar
flour, steam for one hour, filter through cotton and paper, tube and autoclave for
fifteen minutes at 1 1 s^C. As this potato agar varies widely in acidity, to reduce this
variation a large cjuan tity of potato juice made from a uniform lot of Burbank po-
tatoes is used. This if placed in looo-c.c. flasks tightly plugged and kept in a refrig-
,
erator. The juice is "then made up in agar tubes as needed. It was found that this
agar varied less than i per cent, in acidity, changing from -[-7 to -|-6 during five
months.
LESSON 16
Starch Agar— -TfefSjoo c.c. of boiling water add 10 grams of potato starch sus-
pended in a little eoitfi water.. Concentrate by boiling to 500 c.c. This breaks up
the starch grains endJ it should give a nearly transparent starch solution.
Starch solu ifiiiam, c.c Soo

Nutrient S( Jtetion (same as for cellulose agar), c.c 500
Agar, gran is... 10
Cellulose Agar (Ai "^cBeth and Scales: Bull. 266, Bureau of Plant Industry,
p. 27). — Prepare a^ Mt< ^r of dilute ammonium hydroxide solution by adding 3 parts
of water to 10 parts o; f ammonium hydroxide, sp. gr. 0.90. Add a slight excess of
copper carbonate ands. hake, allow to stand over night and then siphon off the super-
natant solution. AcM 10 grams of unwashed sheet filter paper and shake occasion-
ally until thepaper isi; 'ssolved. Dilute to 10 liters and add slowly a i to 5 solution
of HCl, with vigorous ,
shaking until the precipitation of the cellulose is complete.
Dilute to 20 liters, fjilib? w the cellulose to settle and decant the supernatant liquid.
Wash by repeated clha iges of water, adding HCl each time until the copper color
disappears; then wa^ with water alone until the solution is free from chlorine.
Allow it to settle severnil days and decant off as much of the clear solution as possible.
If the percentage of: fflt llulose is still too low, a portion of the solution is centri-
fugalized to bring i-^ ifc ellulose content up to i per cent.
LABORATORY AND TEACHING METHODS Oil
Cellulose solution, c.c 500

Agar, grams 10
Nutrient solution, as follows:
Potassium phosphate (dibasic), gram i
Magnesium sulphate, gram i
Sodium chloride, gram i
Ammonium grams
sulphate, 2
Calcium carbonate, grams 2
Tap water, c.c 1000
Chestnut Twig Agar. —To 275 grams of one- or two-year-old chestnut branches
add 500 c.c. of distilled water and boil over an open flame for one-half hour. Filter
the juice and make up to 550 c.c. with distilled water. To 50 parts of this infusion
add 100 parts of distilled water and 2 per cent, of agar flour. Steam for one-half
hour, filter, tube and autoclave for fifteen minutes at ii5°C.
LESSON 17
Culture Medium. — Winogradsky employed for culturing upon solid media a

mineral gelatin. A solution of from 3 to 4 per cent, of silicic acid in distilled water
is placed in flasks. By addition of the following salts to such a solution gelatiniza-
tion occurs.
(a) Ammonium sulphate, gram 04

Magnesium sulphate, gram 0.05
Calcic chloride a trace
{b) Potassium phosphate, gram o. i
Sodium carbonate, gram 0.6,0.9

Distilled water, c.c 100 o .
The sulphates and chloride are mixed in 50 c.c. of distilled water, and the latter
substance in the remaining 50 c.c. in separate flasks. After sterilizing and cooling
these are all mixed and added in small quantities to the silicic acid. Upon this
medium, it is possible to subculture a pure growth from the film at the bottom of
the flasks in which the nitrous organism is first isolated (c/. Newman, George:
Bacteria, pp. 154-157).
Isolation of the Nitric Organisms. — Nitrobacter develops freely in solutions to
which no organic matter has been added; indeed, much organic matter will prevent
its growth. Winogradsky used the following medium to isolate it:
Water, c.c 1000 o

Potassium phosphate, gram i . o
Magnesium sulphate, gram 0.5
Calcium chloride a trace
Sodium chloride, grams 2.0
About 20 c.c. of this solution is placed in a flat-bottom flask and a little freshly
washed magnesium carbonate is added. The flask is closed with cotton-wool, and
^
6l2 LABORATORY EXERCISES
the whole is sterilized. To each flask 2 c.c. of a 2 per cent, solution of ammonium
sulphate is subsequently added. The temperature for incubation is 3o°C (Fig.
216), This organism can be successfully grown on silicate jelly. As silicate jelly
is difficult to make it is optional for the students to attempt its manufacture. For
reference the method is given.
Pot Experiments with Nitrogen Fixation.-— ?>\nct the experiments of Hellriegel
and Wilfarth and other experimenters, it has been known that certain bacteria
Fig. 216. —
Double-walled copper incubator constructed with non-conducting
materials, with water gauge and openings for insertion of thermometer and thermo-
stat. Padded outer door of copper, inner door of glass. (Fig. 22, p. 46, Schneider',
Pharmaceutical Bacteriology, 191 2.)
{Bacillus radicicola, etc.) have the power of fixing free atmospheric nitrogen, when
they enter the roots of leguminous plants with the formation of root nodules. The
formation of these nodules can be followed in a series of experiments.
^ It is optional of course for the teacher to omit these rather difficult exercises
entirely. If followed by the student or class, a useful work to consult in connection
with Lesson 17 is Smith, Erwin F.: Bacteria in Relation to Plant Diseases, I 36-39. :
2 An important paper on the culture and isolation of Bacillus radicicola is by

Harrison, and Barlow, B.: The Nodule Organism of the Leguminosae— Its
F. C.
Isolation, Cultivationand Commercial Application. Centralblatt fiir Bakteriologie,
Parasitenkunde und Infektionskrankheiten, 19, Abt. 2, 1907: 264-272, 426-440,
pis. 9. Consult for other details Lipman, J. G. and Brown, P. E.: A Laboratory
Guide in Soil Bacteriology, 191 1.
LABORATORY AND TEACHING METHOr)S 613
Take three pots A, B, C, which have been thoroughly sterilized by dry heat in
a sterilizing oven. Place in pot A ordinary rich garden soil. Fill pot B and C with
sand and thoroughly sterilize Plant in pots, A, B
both pot and sand with dry heat.
and C seeds of pea, bean, clover or those of other leguminous plants and water
pots A, and B only with distilled water previously carefully sterilized. Pot C with
sand, is watered with distilled water which has been allowed to percolate through
rich garden earth and which removes the bacterial life which such rich soil con-
tains. Pot C watered with such water, therefore, becomes microbe-seeded. After
the first watering, all subsequent applications of water should be made with
thoroughly sterilized distilled water.
Note daily the growth of the plants in each of the pots and explain the difference
in the rate and character of the growth, if any.
In order to be able to study microscopically the entrance of the organisms from
the soil into the root of the leguminous plants a larger series of pots should be used
than three. By doing this successive stages in the development of the nodules can
be obtained and made ready for microscopic study by the paraffin method described
in a subsequent lesson (Page 656).
LESSON 18
Media (F. D. Heald). Bacteria and fungi are in-

Standardization of Culture —
fluenced in their development by the degree of acidity or alkalinity of the medium
in which they are growing. Since this is true, it is important to employ media of
known reaction. In order to secure results which may be compared, the adoption
of auniform method of standardization is necessary and the reaction of a culture
medium should be indicated always when cultural or morphologic characters are
described. The standardization of culture media requires the following solutions:
N
c^r—r— = a normal solution of sodium hydroxide.
'N
— NaOH = twentieth normal solution of sodium hydroxide.
N
ff™ = normal hydrochloric acid.
N N
The — NaOH used for the titration
.
is of culture media and the ^^-7^^ for their

20 NaOH
N
neutralization, vj^ is used for acidifying media. A normal solution contains
I gram of basic H, or the equivalent to each 1000 c.c. Since the above normal solu-
tions are required in every pathologic laboratory, directions are here given for their
preparation.
—
Preparation of Normal Solutions. Normal solutions of NaOH or HCl cannot be
made by weighing. NaOH readily absorbs CO2 and water from the air and so can-
not be weighed accurately enough for making standard solutions. HCl is liquid
and of varying strength. It is necessary, then, to start with an acid or alkali that
is in solid crystalline form and is not altered on exposure to the air. Oxalic acid
presents the requisite characteristics.
6 14 LABORATORY EXERCISES
N
— Oxalic Acid Solution. — Weigh out exactly 6.3 grams of chemically pure oxalic
. . . .
acid (H2C2O4 plus 2H2O) and add distilled water in looo-c.c. volumetric flask.
After the crystals of acid have dissolved, dilute the solution until it measures exactly
1000 c.c.
N
Tj—7;tf or Normal Sodium Hydroxide. —This solution should contain 40 grams of
NaOH in i liter. It can be made by titrating against the standard oxalic solution
already prepared. Weigh out 90 grams of NaOH and dissolve in 2 liters of dis-
tilled water. This solution is now too strong and the amount necessary to dilute it
N
must be determined. Place exactly 50 c.c. of the oxalic acid in a beaker and add
10
a few drops of phenolphthalein solution to serve as an indicator and then add to this
drop by drop from a burette some of the NaOH solution, stirring with a glass rod
and continue until the solution is turned a faint, but permanent pink color. Read off
from the burette the amount of NaOH solution used to neutralize 50 c.c. ot
—N
10
oxalic acid, which contained as much acid as 5 c.c. of normal acid. Now calculate the
amount of dilution necessary. Supposing 4.5 c.c. of NaOH be the amount used and
1950 c.c. the amount of NaOH to be diluted, the proportion would be as follows:
4.5 : 5 :: 1950 : x where x = 2167, and this means that 2167 c.c. of water must be
used. After the dilution, repeat the titration and adjust if necessary.
N —This may be prepared by making an acid
TTpj or Normal Hydrochloric Acid.
solution which is a little over strength, and determining the amount of dilution
N N
necessary by titrating with the „ ^ ^^ -
i c.c. of c^t^ should exactly neutralize
N .
^
ic.c.ofjj^l-
Expressing the Reaction of Media. — Fuller's scale has been generally adopted for
expressing the reaction of culture media. The plus sign (+) indicates that the
medium is acid to phenolphthalein, while the minus sign ( — ) indicates that the me-
dium is alkaline to phenolphthalein, the figure following the sign indicating the
N
degree of acidity, or alkalinity. For example, a -f 10 medium contains 10 c.c. of tt;^,
for 1000 c.c. beyond the neutral point for phenolphthalein paper. A— 10 medium
N
is alkaline and would require 10 c.c. of t77^\ for 1000 c.c. to bring it back to the
neutral point. Media may then have the reaction + 5, + 10, + 15, etc., or — 5,
— 10, —15, etc. The neutral point for litmus is not the same as the neutral point
for phenolphthalein and this fact should be kept in mind when working with culture
media.
25 of Fuller's scale gives approximately the neutral point for litmus, so that any
medium with a reaction less than + 25 is still alkaline to litmus.
—
The Optimum Reaction. For every organism there is a definite optimum reaction.
It lies near -f 5 for most animal pathogens, about -fio to +15 for most water and
putrefactive bacteria and +10 to +25 or even higher for fungi. There are some
bacterial organisms which prefer distrnctly alkaline media (Fuller's scale), while
others prefer acid media. A good general practice to follow in the preparation of the
basic culture media to be kept in stock iis to standardize to +10 of Fuller's scale and
vary the reaction according to the preierence of the organisms under cultivation.
When other acids than HCl are used for acidifying the media, the kind should be:
definitely specified, when the reaction is expressed.
Titration of Media. — In outlining the m ethod of preparation of bouillon for routine
work, directions were given for neutralization of the medium and the addition of the
requisite amount of acid. In accurate work, or in the prosecution of research, a
more careful method of standardizati.on is employed. The medium should be
neutralized by the titration method. iTie process is as follows:
1. Add exactly 5 c.c. of the medium to 45 c.c. of distilled water in an evaporating
dish (use a 5-c.c. Mohr pipette), boil for three minutes to drive off the CO2 and add
I c.c. of phenolphthal(;in solution.
N
—
2. Add NaOH drop by drop from a burette, stirring constantly until the
20
solution turns a faint, but permanent pink. Repeat the titration for two more 5-c.c.
samples, and determine the average of the three readings.
N
3. Calculate the amount oi
^ ^.tV necessary to neutralize the medium (10 to
15 CO.), add the amount determined to the medium, te^t reaction and if neutral,,
proceed with preparation of the medium; if not, repeat the titration on neutralization.
LESSON 19
Germination Studies. —The examination of spore germination of various fungi can

be studied best by the hanging-drop method. Take a hanging-drop slide and sterilize
thoroughly in the hot-a ir oven at iio°C. after it has been wrapped in a crepe napkin
or piece of tissue paper. After sterilization plunge it into a beaker of absolute alcohol
(or such sterilized slides may be kept in stock in absolute alcohol) and then drain
off the greater part of the spirit, grasping the slide in a pair of sterile forceps. Burn
off the remainder of the alcohol in the flames.
Place the hanging-drop slide on a piece of blotting paper moistened with a 2
per cent, lysol solution and cover it with a small bell glass that has been rinsed with
the same solution and not dried.
Raise the bell glass slightly and smear sterile vaseline around the rim of the cell
by means platinum wire. Remove a clean cover-slip

of a steriHe spatula of stout
from the alcohol pot with sterile forceps and burn off the alcohol; again raise the
bell glass and place the sterile cover-slip on the blotting paper by the side of the
hanging-drop slide.
Remove a drop of the culture medium selected for use (see below) and place the
drop on the center of ihe cover-slip. Sterilize the loop.
Raise the bell glass; sufficiently to allow of the cover-slip being grasped with the
sterile forceps, invert it and place over the cell of the hanging-drop slide. Remove
the bell glass altogether and press the cover-slip firmly on the cell.
6l6 LABORATORY EXERCISES
Germination on Solid Media. — Observing precisely similar technique a few drops

of liquefied gelatine or agar may be run over the surface of the cover-slip and a
hanging-drop plate cultivation thereby prepared. After sealing down the prepara-
tion it may be set aside and the growth watched at definite intervals under the
microscope.
Dilution Method to Obtain Material for Inoculating Hanging-drop Media. — In
the case of yeast this problem was solved by Hansen, who developed the method to
such a degree of perfection as to create, in fact, an exact method (1881). He
employed dilution with water. The yeast developed in the flask is diluted with an
arbitrary amount of sterilized water, and after vigorous shaking, the number of
cells in a small drop of liquid is determined. The counting, in this case, is effected
in a very simple manner by transferring a drop which
to a cover-glass, in the center of
some small squares are engraved and this is then connected with a moist chamber;
the drop must not be allowed to extend beyond the limits of the square. The cells
present in the drop are then counted. Suppose, for instance, that ten cells are
found: a drop of similar size is transferred from the liquid, which must first be
shaken vigorously, to a flask containing a known volume of water, e.g. 20 c.c. of
sterilized water. This flask, then, will in all likelihood contain about ten cells. If
it is then vigorously shaken for some time until the cells are equally distributed in
the water, and then i c.c. of the liquid introduced into each oftwenty flasks contain-
ing nutritive liquid, it is probable that half of these twenty flasks have received one
cell each. But, here again, as in Lister's experiments, it is entirely a calculation
of probabilities. If the flasks are allowed to stand for further development of micro-
organisms, there will be a chance of getting a pure culture in some of them. Hansen
succeeded, however, in adding a new factor, which first gave certainty to this experi-
ment. Thus, if the freshly inoculated flasks are vigorously shaken, and then left in
repose, the individual cells will sink to the bottom and be deposited on the walls of
the flask. It is self-evident that if a flask contains, for instance, three cells, these
cells will always, or at least in the great majority of cases, be deposited in three
distinct places on the bottom. After some days, if the flask is raised carefully, it
will be observed that one or more white specks have formed on the bottom of the
flask. If only one such speck be found, we have a pure culture by the dilution
method.
—
Method of Preparing Squared Cover-glasses.- Since such cover-glasses are some-
what expensive and can be easily etched, the method of their preparation is de-
scribed below. A little paraffin or wax is melted in a saucer and the cover-glass
dipped into it, being held at one corner by a forceps; it is taken out quickly and as
much as possible of the melted parafiin is allowed to run off, leaving on either side a
thin cover of parafiin which is allowed to harden. By a very fine needle and a small
ruler the required lines are then scratched on the wax, and the cover-glass immersed
for a moment in hydrofluoric acid which should be poured into a platinum crucible
or dish. The paraffin can now be dissolved off in xylol, leaving the surface etched
with the squares used in making bacterial, or fungous spore counts (Fig. 217).
These squared covers may be raised above the slide, while the count is being made,
either on four pillars of paraffin, or in a moist chamber.
LESSON 20
Counting of Yeast Cells, —

Fungous Spores and Bacteria. In many cases the cells
are in a liquid which is inclined to form froth when shaken, hence the liquid can be
treated with dilute sulphuric acid (i part concentrated sulphuric acid and 10 parts
water). This prevents aggregations of the cells and also furnishes in addition a
liquid in which cells do not sink to the bottom too quickly, an important point, when
single drops are taken out for counting purposes.
In counting, the counting chamber is employed. Thoma's ha;matimeter consists
of a glass slip on which a cover-glass is fastened which has a circular hole "in the
2
6i8 LABORATORY EXERCISES
As soon as the cover-glass has been put into position the chamber is laid under
the microscope, and if a haematimeter is being used as a counting chamber the "net
eyepiece" is required. It is not advisable to use a greater magnification than is
necessary. After waiting a short time, the counting is proceeded with when all the
cells in the preparation have sunk to the bottom. The "net eyepiece" consists of
a large square divided into sixteen or twenty-five smaller squares, the latter being
used as aids in counting. The cells inside the large square are counted; it does
not matter how the cells lying on the side lines of the square are counted, if the
same rule is always followed. Many squares in each haematimeter may be counted
by di^placihg the haematimeter. It is to be recommended always to count a certain
Tief-e
0.1 OO mmi
1 A 1
400 ^ 25
q mm.
Sample i
Square
When It is intended to sow a definite number of cells,i water is usually added to

the yeast to be used as sowing material, the cells being thus more easily separated
from one another on shaking; also, no appreciable increase of the cells takes place,
especially if the flask is subjected to a low temperature after the sample has been
withdrawn.
Fig. 2i8A. —
Blood counter case, a, Slide with counting chamber; b, rubber cork
covering tip of white pipette; c, soft rubber tubing; d, red pipette provided with
rubber cork; e, cutting needle in 95 percent, alcohol; g, Hayem's' solution; h, .5 per
cent, acetic acid. {After McJunkin.)
The yeast is, therefore, shaken up vigorously and continuously with sterile water,
and an average sample removed. There are three different cases to be considered
now viz.: (i) When we wish to know only how many cells are present in a certain
portion of the water-yeast mixture; (2) when it is intended to inoculate a previously
determined number of cells into the liquid to be dealt with; and (3) when it is desired
to sow so many cells, that after the seeding the definite number of cells desired may
be present in an arbitrary space unit, e.g., when making comparisons of the multiply-
1 Klocker, Alb.: Fermentation Organisms, 1903.
I
ing powers of two species. In the first two cases, it is required to determine the
actual number of cells which are to be seeded, and no attention is paid to the quantity
of liquid indtulated; in the last case, it is required only to know the relative number
of cells,but regard must be had to the quantity of liquid seeded. Finally, the follow-
ing must be remembered: If there is to be a definite volume in the flask after seed-
ing, then, in the case where the seeding is not to be made in water, or where the con-
centration of the liquid is of some account, no water must be used in shaking up the
yeast. In this case the same culture liquid must be employed. The same quantity
of culture liquid is then removed from the flask before seeding, as will be added when
seeding takes place.
The procedure in the above three cases is as follows: (i) After shaking, a drop of
the water is placed in the haematimeter, or in the Thoma chamber, and the number
of cells is determined in the usual manner. On seeding a measured portion of the
water mixture is taken, and we thus know how many cells have been sown.
2. As above. In counting we learn, for example, that a cells are present in a
certain volume. It is know the quantity of culture liquid in the
here necessary to
flask to be inoculated; assume the amount to be p c.c. If it is desired to sefed so
many cells that there will be ai cells per unit of volume, the number of cubic centi-
meters X of the water-yeast mixture, which must be added in order to arrive at this,
isfound from the following equation:

a
=
p -\- X
—
or the number of cells in the ; —
water mixture (the seeding liquid) has the same proportion to the cells after seed-
ing as the whole amount of liquid after seeding has to the amount of seeding liquid.
The quantity of liquid in the flask after seeding has taken place is thus p -\- x.
From the given equation, x = Example: It is found that the seeding

a — 02
.
liquid contains 75 cells per unit of volume and the flask to be infected contains 70
c.c. of wort, and it is further desired to have 5 cells per unit of volume after inocula-
tion. Accordingly, x = = 5 c.c. to be withdrawn from the seeding liquid.

75 ~ 5
The result may be checked by another counting after seeding. If the result is in-
correct, either more liquid or more cells must be added. -But in exact work this
contingency does not arise.
Suppose it is wished to sow ai cells of a yeast species /I, and bi cells of a species B
in a flask containing p c.c. of culture liquid, from two seeding liquids containing a
and b cells per unit of volume respectively. The number of cubic centimeters x
and y, to be sown from A and B respectively, is found from the following equations.
a^
^ p +x+ y , ^* ^ P +x+ y
ai X bi y
the quantity of liquid after infection being p + x + y; from this we find:
_ aibp _ abip
ab — Oib — aibi ab — Oib — aibi
Combinations of the above three cases may of course occur but from the explana-
tions given here it will not be difficult to solve them.
.
LESSON 21
Cidtivalion of Yeasts on Gypsum Blocks.- —Spore Cultivation. — Blocks of gypsum

are used generally for the cultivation of the spores of the yeasts. The block is in
the form of a truncated cone, and the cover of the vessel fits quite loosely. The
dishes used in the Carlsberg laboratory are the so-called bird troughs (Vogelnapfe)
Fig. 219. — Method of pouring gelatin into Petri dishes. {After Lohnis.)
A suitable size for these, taking outside measurements, is as follows: height 4.5 to
S cm.; diameter of the bottom about 7 cm. The gypsum block is 3 cm. high; the
diameter of the lower surface is upper surface 3.8 cm. To make
5.3 cm., that of the
a gypsum block, 2 parts of powdered gypsum are mixed with %
part of water and the
mixture poured into a tin mould. The block should be hard, and the mould must
not be rubbed with fat, oil or such material. A culture on a gypsum block in such
Fig. -Petri dish. {After Williams in Schneider, Pharmaceutical Bacteriology,

p. S9-)
a vessel cannot, as a rule, be kept free from bacterial infection, for the cover must
not be closed down but should allow free access of the air. The dishes with
tightly,
gypsum blocks are sterilized for one to one and a half hours at 110° to ii5°C., the
dishes first being wrapped in a crepe napkin or in filter paper. The gypsum blocks
are sterilized in a moist condition before planting the yeast on their upper surface.
The gypsum blocks can be used several times.
Method of Pouring Plates (Fig. 219). — Place three sterile Petri dishes (Fig. 220)
in a row after previously sterilizing them wrapped in a crepe napkin in the hot-air
oven.
Take numbered i, 2 and 3 and fill with the liquefied
three sterile test tubes
nutrient to be used. Plug each tube with cotton and flame the plugs, which should
be removed readily from the mouths of the tubes.
Add one loopful of inoculum to tube No. i. After replugging, rotate the tube
between the palms of the hands with an even circular movement to diffuse the in-
oculum throughout the medium; avoid jerky movements as these cause bubbles
of air to form in the medium.
platinum loop and add two loopfuls of diluted inoculum to tube
Sterilize the
No. 2 and mix as before. In a similar manner transfer three loopfuls of liquefied
medium from tube No. 2 to tube No. 3 and mix thoroughly.
Flame the plug of tube No. i, remove it, then flame the lips of the tube; slightly
raise the cover of Petri dish No. i, introduce the mouth of the tube; then elevate
the bottom of the tube, pour the liquefied medium into the Petri dish to form a thin
layer. Remove the mouth of the tube and close the "plate." If the medium has
failed to flow evenly over the bottom of the plate, raise the plate and tilt it to rectify
the fault.
Pour plates No. 2 and No. manner from tubes Nos. 2 and 3. Label
3 in a similar
the plates with the distinctive name or number of the inoculum, the number of the
dilution, also the date.
In this way colonies may be obtained quite pure and separate from each other.
They may be described as such, and may then be transferred as pure cultures to
other media in other test-tubes.
In plate No. i probably the colonies will be so numerous and crowded, and there-
fore so small, as to render it useless. In plate No. 2 they will be more widely sepa-
rated, but usually No. 3 is the plate reserved for careful examination, as in this the
colonies are usually widely separated, few in number and large in size.
Agar plates are poured in a similar manner, but the agar must be melted in boil-
ing water and then allowed to cool to 42°C. or 45°C. in a carefully regulated water
bath before being inoculated and the entire process must be carried out very rapidly
otherwise the agar will have solidified before the operation is completed. After
the agar has hardened it is incubated at 37°C. and the plates are inverted as this
prevents flooding of the agar surface by the squeezing out of the water of condensa-
tion as the agar hardens. Gelatin plates are not inverted.
Streak Method. —The isolation of pure cultures of organisms by the streak
method differs from the plate method in that the medium (gelatin, agar, blood
serum) is not inoculated in the fluid state but the necessary dflution to secure iso-
lated colonies is secured by drawing a glass rod with its end bent into a triangle,
as recommended by Bergey, several times across the surface of the sterile medium
in Petri dishesby lifting the cover while so doing. The <^ glass rod
has been previously infected with the material to be studied qualitatively. It is
preferable, according to Bergey, to place a small quantity of the mixed culture

in the center of the first plate of a series, and thence distribute the material
over three or more plates in succession with the glass spreader. Eventually a
degree of dilution is reached where distinct colonies are in evidence.
LESSON 22
Isolation of a Leaf Wilt Fungus in Pure Culture. —

Given a fungus causing leaf
wilt, to obtain a pure culture by excluding the non-pathogenic forms.
I. Look for the fruiting stage of the suspected fungus, or fungi. Transfer some
of the spores with a sterile needle into a tube of 5 c.c. of sterile water. (If pycnidia
or perithecia are present, transfer a whole pycnidium or perithecium into sterile
water, and crush the fruit body to cause the escape of the spores). Then with a
sterile needle transfer some of the water with the spores into a tube of agar-agar
which is liquid by putting in a vessel of hot water and then allowed to cool.
made
This tube marked A. Then from tube A transfer a drop of agar with a sterile needle
is
to another similar test-tube with liquid agar designated as B (Fig. 221). Then
perform the same sort of transfer to a third tube C. Distilled water or nutrient
bouillon can be used for these dilutions instead of agar.
Fig. 221. Fig. 222.
Fig. 221. —
Method of holding test-tubes in transfer of fungi from one test-tube
to another. {After Lohnis.)
Fig. 222. —
Cylindric form of wire basket for holding test-tubes during steriliza-
tion and other operations. {After Schneider, Pharmaceutical Bacteriology, p. 37.)
^, J5 and C are thoroughly shaken and each is transferred to Petri dishes marked
A, B and C. If water is used to must be mixed with the material
dilute, or bouillon, it
poured into the Petri dishes. These are observed for any growth that may take place
on the surface of the agar-agar. Transfers are made from the single colonies into
agar slants in test-tubes.
If no spore forms are present, cut out piecesof the affected leaf and place in a
tube containing per cent, mercuric chloride diluted in equal amounts in 50 per cent,
i
alcohol. Shake the tube so that the material is bathed in the disinfectant. Do
this for half a second to two minutes according to the thickness of the leaf.
Pour
off the disinfectant and wash the material three times in sterile water, care
being
taken to keep out foreign infection. Then with a sterile forceps, take each piece
of the materialand crush it thoroughly at the mouth of a tube containing melted
—
;ind cooled agar. When the materuU it is well shaken up with agar and
is crushed,
the whole poured into a Petri dish. growth of one fungus appears, it means
If the
that we have the parasite in captivity, or pure culture. If more than one fungus
is obtained, they must all be transferred separately into agar slants in test-tubes
and tested by inoculation for their pathogenicity. The true pathogen is of course
the one which will reproduce all of the symptoms of the disease.
Note. —
To keep out bacterial infection put one drop of a 5 per cent, lactic acid
in each of the agar tubes used in making the cultures.
Differential Methods of Isolation
Pasteurization and Sterilization. —In order to compare the effect of these two
operations on organic material, take some milk and pasteurize part of it and sterilize
the other part by one sterilization. Conduct both operations in previously sterilized
flasks plugged with cotton after the milk is introduced (Fig. 223).
Milk is pasteurizedby heating it up to a temperature of 8s°C. followed by a
rapid cooling. Milk is sterilized by heating up to ioo°C. for five minutes. Set
the flasks aside and compare. Note any changes that may take place.
Differential Media. — (a) Selective. — Some media are specially suitable for cer-
tain species of bacteria and enable them to overgrow and finally choke out other
varieties.
{b) Deterrent. —
The converse of the above also. Certain media possess the
power of inhibiting the growth of a greater or less number of species. For instance,
media containing carbolic acid to the amount of i per cent, will inhibit the growth
of practically everything but the Bacillus coli communis.
Differential Sterilisation. —
(a) Non-sporing Bacteria. —
Similarly, advantage may
be taken of the varying thermal death points of bacteria. From a mixture of two
organisms whose thermal death points differ by, say, 4°C. e.g., Bacillus pyocyanens,
thermal death point 5S°C., and Bacillus mese7itericus vulgatus, thermal death point
6o°C. —
a pure cultivation of the latter may be obtained by heating the mixture in
a water bath to 58°C. and keeping it at that point for ten minutes. The mixture
is then planted on to fresh media and incubated, w^hen the resulting growth will be
found to consist entirely of B. mescntericus.

{b) Sporing Bacteria. —
This method is found to be ofeven greater practical value
when applied to the differentiation of a spore-bearing organism from one which
does not form spores. In this case the mixture is heated in a water bath at 8o°C.
twenty minutes. At the end of this time the non-sporing bacteria are
for fifteen to
dead, and cultivations made from the mixture will yield only a growth resulting
from the germination of the spores only.
differential atmosphere cultivation
—
For the separation of bacteria, it is possible to draw the
Aerobic and Anaerobic.
linebetween those that need oxygen for growth (aerobic) and those that will grow
without ox>'gen (anaerobic). By excluding oxygen, anaerobic forms alone develop.
Inoculation into various animals or plants may be used as a means of separation.
40
LESSON 23
Walcr Analysis.
I. Collect water from tap in a sterile Erlenmeyer flask, allowing H2O to run for
ten minutes before collecting.
II. Melt two tubes of gelatin at 42°C.
III. Add to tube No. A o.i c.c. and tube No.
2 0.2 c.c. from the flask. Shake
to mix H2O with gelatin.
IV. Pour in Petri dishes No. A and B and place in locker.
V. Count colonies which develop at end of twenty-four and forty-eight hours.
VI. Estimate the number of colonies which would have developed in i c.c. of
water.
Example.
Twenty-four hours
50 colonieshave developed on plate No. A — 50 X 10 = 500 in i c.c.
96 colonies have developed on plate No. B —96 X 5 = 480 in i c.c.
2)980
490 in I c.c.
Forty-eight hours
62 colonies have developed on plate No. A — 62 X 10 = 620 in i c.c.
102 colonies have developed on plate No. B — 102 X = 510

5 in i c.c.
2)1130
565 in I c.c.
LESSON 24
METHODS OF IDENTIFICATION
Descriptive Terms. — For complete details consult Eyre, J. W. H. The Elements
:
of Bacteriological Technique, 1902: 208.
Types of Colonies
A. —The
Size. the
size ofand the spores at various
cells ages.
— Punctiform, round,
B. Shape. fusiform, cochleate, amoeboid,
elliptic, irregular,
mycelioid, filamentous, conglomerate,

floccose, rhizoid, toruloid, rosulate.
C. Surface Elevation. — convex, capitate, umbonate,
Flat, pulvinate, effused,
umbilicate, raised.
D. Character of Surface.- — Smooth, alveolate, punctate, bullate, vesicular,
verrucose, squamose, echinate, papillate, rugose, corrugated, contoured, rimose.
E. Internal Structure of Colony (Microscopic). — Refraction weak, refraction
strong, amorphous, hyaline, homogeneous, homochromous, finely granular, coarsely
granular.
F. Optic Characters. —Transparent, vitreous, oleaginous, resinous, translucent,

porcelaneous, opalescent, nacreous, sebaceous, butyrous, cetaceous, opaque, creta-
ceous, dull, glistening, fluorescent, iridescent, color of colonies.
G. Edges of Colonies. —Entire, undulate, repand, erose, lobulate, auriculate,
lacerate, fimbriate, ciliate.
Pig. 223. —
îy
Types of growth in stab cultures. A, Non-liquefyinp; i, Filiform
{Bacidiis coli); 2, beaded {Streptococcus pyogenes); 3, echinate {Bacterium acidi

lactici); 4, villous {Bacterium murisepticiim); 5, arborescent {Bacillus mycoides).
B, Gelatin liquefying. 6, Crateriform {Bacillus vulgare, 24 hr.) 7, napiform {Bacillus
;
sublilis, 48 hr.); 8, infundibuliform {Bacillus prodigiosus) ; 9, saccate {Microspira

Finkleri); 10, stratiform {Pseudomonas flavescens). {From McFarland after Frost in
Schneider, Albert: Bacteriological Methods in Food and Drug Laboratories, 1915: 87.)
TYPES OF STAB CULTURES
A. Surface Growth. — Filiform, beaded, echinate villous, arborescent.

—
B. Character of Liquefied Gelatin. Pellicle on surface, uniformly turbid, granular,
mainly clear but containing flocculi, deposit at apex of liquefied portion, production
of gas bubbles.
C. Area of Liquefaction (if present). — Crateriform, saccate, infundibuliform,
napiform, fusiform, stratiform (Fig. 223).
LESSON 25
Plate Counter. —The most accurate method of counting the colonies on each of
the plates is by means of the counting disk. These disks consist of a piece of paper,
upon which is printed a dead black disk, subdivided by concentric circles and radii
painted in white. In Jeffer's counter each subdivision has an area of i sq. cm.: in
Pake's counter, radii divide the circle into sixteen equal sectors, and counting is
facilitated by equidistant concentric circles. (For disks see Eyre, p. 322.)

(a) In the final counting of each plate, place the Petri dish over the counting
disk, and center it, if possible, making its periphery coincide with one or other of the
concentric circles.
{h) By means of a hand lens count the colonies appearing in each sector in turn.
Make a note of the number present in each.
(c) If the colonies present are fewer than 500 the entire plate should be counted.
If, however, they exceed this number, enumerate one-half, or one-quarter of the
plate, or count a sector here and there, and from these figures estimate the number
of colonies present on the entire plate.
counting plate^ (Fig. 224) consists of concentric zones which are divided
Jeffers'
into small sections, each having an area of i sq. cm. To determine the position of
the circles marked 10, 20, the position of the circles marked 10, 20, 40, 60, 100 and
140 in the diagram, whose areas equal 10, 20, 40, 60, 100 and 140 sq. cm. respectively,
the formula, wr"^ = was used. In order to show the application of the formula,
area,
the radius of the circle whose area is equal to 10 sq. cm., will be found from the formula
as follows:
TT = 3.I416.
irr- = 10 or r^ = 10 -i- tt.
10 -
^ 3-1416 = 3.18309 or r-.
\/3-i8309 = 1.78-f or r.
1.78 + cm. = the radius of a circle whose area is 10 sq. cm. Dividing the circle
into ten equal sectors, each sector has an area equal to same i sq. cm. By the
method we whose area equals 20 sq. cm. thus making each
find the radius of a circle
of the ten spaces between circles 10 and 20 and bounded laterally by the ten radii
equal to i sq. cm. We next construct a circle whose area equals 40 sq. cm. and divide
each sector as far as circle 20, making twenty equal areas between circles 20 and 40,
each equal to i sq. cm. In like manner we construct circles 60, 100 and 140 divid-
ing the sectors in the zone lying between circles 60 and 140 to produce areas equal
to I sq. cm. each. If a plate whose area is greater than 140 sq. cm. is used, a circle
whose area is 180 sq. cm. can be drawn and the radiating lines extended out to the
circle (Fig. 224).
The Petri dish can be centered upon this apparatus by the circles and the area
read from the line its edges approach. To facilitate the reading of the area of the
plate the circles 80 and 120, whose areas are equal to 80 and 120 sq. cm., respectively,
1
Jeffers, H. W. An Apparatus
: to Facilitate the Counting of Colonies of
Bacteria on Circular Plates. Journ. Applied Micros., I: 53-54, March, 1898.

were drawn as dotted circles, thus making the areas marked "a" and "6" equal to
0.5 sq. cm. The colonies in several areas can be counted, an average taken, and the
result multiplied by the number of square centimeters in each plate.
A apparatus could be made by covering a plate of glass with a uniform layer
fine
of wax and with a sharp instrument cut the figure in the wax and subject it to hydro-
fluoric acid for a few minutes which would etch the glass where exposed. Cleaning
Fig. 224. —
Jeffer's circular counting plate for Petri dish cultures. The entire
area (100 sq. cm.) is marked off into the equal sectors of ten sq. cm. each. {After
Schneider, Pharmaceutical Bact. p. 90.)
off the wax and placing the glass plate over black velvet, the colonies could easily
be counted.
—
Marking and Counting Apparatus for Bacterial Colonics. The apparatus
Neisser's
is for counting bacterial colonies and for marking off their position.
employed
When in nsC the apparatus is mounted on the lid of the box with which it is
supplied, thus the latter serves at the same time as a, base.
For this is screwed on to the inside of the lid, which

purpose a metal guide plate
latter is reversed when arranged for use and the marking apparatus
the instrument is
is placed on this plate. This apparatus consists of a vertical pillar with square base
plate and a metal frame which is vertically adjustable by means of a rack and pinion.
The horizontal movement is obtained by moving the entire dish carrier along the
guide plate which is screwed on to the box lid.
The Petri dish is secured in the frame by means of two milled heads which are
fixedon the right-hand side and at the bottom.
Immediately behind the Petri dish is mounted a glass screen divided into squares,
which as a further aid to localization, are subdivided and numbered.
A second pillar is screwed into the lid in front of the dish holder and carries the
lens. The lens is vertically adjustable and is threaded for focusing purposes.
Below the lens carrier is fitted a horizontal bar which serves as a hand rest when
marking ofT the colonies.
A special counting screen is provided with fifteen square openings arranged in a
V-shape (echelon) by means of which the number of colonies at four places in sixty
squares may be determined.
At the upper edge of the counting screen lines are ruled which serve as scales for
the Petri dish; the numbers on the one side indicate the diameters in millimeters
corresponding to each scale line, while the numbers on the other side indicate how
many times the area of the sixty squares is contained in the area of the whole Petri
dish. Thus in order to ascertain the total number of colonies in the dish, it is only
necessary to count the number of colonies in the sixty squares and to multiply the
figure thus obtained by the proportional number required by the diameter of the dish.
LESSON 26
LABORATORY WORK IN SYSTEMATIC BACTERIOLOGY
As important for students in mycology to be able to identify the various

it is
species of bacteria, which they may meet in their investigation of the fungi, the fol-
lowing suggestions are made as to the systematic study of the forms of bacterial
life. Ordinarily, where the other groups of fungi are to be considered, time will not
permit a detailed systematic study of the bacteria where cultural methods are re-
quired in the identification of the specific forms. Yet much can be done in the class-
room with the microscope in the study of the morphology of selected species. The
following exercises are presented as suggestions to the teacher and student of
mycology.
First Exercise.— The teacher can distribute to each member of the class a selected
number of bacteria in culture tubes. Each tube should be numbered, so that the
student, after determining the generic character of the different organisms handed
to him, can attach the number to his specific determinations, so that the teacher
can check off the results of each student's work by the numbered list of species
kept for such classroom work. The bacteria from each of the culture tubes should be
mounted in balsam after staining with carbol fuchsin, or some other approved stain,
and kept for future reference and study.

1
Second Exercise. —The members of the class can raise material for such morpho-
logic study after tlie first exercise has been completed by partially filling test-tubes
with such materials as chopped hay, prunes, lima beans, split peas, cracked oats and
cabbage leaves, adding water, and treating, as follows:
One set of tubes should be plugged and thoroughly sterilized by differential
sterilization. This experiment, after examination of the material under the micro-
scope, demonstrates that bacterial growth in the tubes does not take place.
A second set of test-tubes can be left open to the air after the water and the
culture material have been completely sterilized. This gives the organisms that
come from the air.
A third set of tubes can be partially filled with water, plugged and then sterilized,
and after sterilization unsterilized material can be added. This gives the organisms
that enter through the vegetable substance.
A fourth set of tubes can be filled with the culture material, plugged and steril-
ized. Unsterilized water can be then added to each of these tubes. This gives the
microbes that come in through the water. These are rough methods adapted to
general class work, and in each case the organisms which appear should be mounted
and systematically studied to determine the different generic forms which are present,
as far as that can be done by staining methods and the microscope.
—
Third Exercise. The teacher can distribute material of diseased plants in which
the disease is directly traceable to some bacterial organism. For this exercise, the
professor should have a stock of at least a half dozen diseased plants properly fixed
and preserved in 50 per cent, alcohol. The material, which has been distributed,
should be cut free-hand by the student and the sections mounted as directed, or the
student can imbed the material in celloidin, or in paraflan, to secure thinner serial
sections by the use of a sliding, or rotary microtome. To carry on this exercise, the
student should have an acquaintance with celloidin and paraffin technique.
—
Fourth Exercise. Where the student has plenty of time and expects to specialize
in the study of the bacterial diseases of plants, then he, or she, should follow the
following scheme suggested by Chester in his "Manual of Determinative Bacterio-
logy," the descriptions and keys of which can be used in a detailed systematic
study of bacterial organisms. This exercise can be pursued only after the student
has learned cultural and isolation methods and not at the beginning of a course in
mycology and its technique.
LESSON 27
—
Scheme for the Study of Bacteria. The Society of American Bacteriologstis has
adopted a numeric system of recording the salient characters of an organism (group
number).
100 Endospores produced.

200 Endospores not produced.
10 Aerobic (strict).
20 Facultative anaerobic.
30 Anaerobic (strict).
I Gelatin liquefied.
2 , Gelatin not liquefied.
0.1 Acid and gas from dextrose.
0.2 Acid without gas from dextrose.
0.3 No acid from dextrose.
0.4 No growth with dextrose.
o.oi Acid and gas from lactose.
o. 02 Acid without gas from lactose.
o. 03 No acid from lactose.
0.04 No growth with lactose.
o. 001 Acid and gas from saccharose.
o. 002 Acid without gas from saccharose.
o. 003 No acid from saccharose.
0.004 No growth with saccharose.
o.oooi Nitrates reduced with evolution of gas.
o. 0002 Nitrates not reduced.
0,0003 Nitrates reduced without gas formation.
o ooooi . Fluorescent.
o. 00002 Violet chromogens.
o 00003 .
Blue chromogens.
o 00004 . Green chromogens.
0.00005 Yellow chromogens.
0.00006 Orange chromogens.
o. 00007 Red chromogens.
0.00008 Brown chromogens.
o 00009
. Pink chromogens.
o 00000
. Non-chromogens.
o. oooooi Diastatic action on potato starch (strong).
o. 000002 Diastatic action on potato starch (feeble).
o. 000003 Diastatic action on potato starch (absent).
o. ooooooi Acid and gas from glycerin.
0.0000002 Acid without gas from glycerin.
o 0000003
. No acid from glycerin.
o 0000004
. No growth with glycerin.
The genus, according to the system of Migula, is given its proper symbol which
precedes the member thus: According to the above the symbol of Bacillus coli
would be B. 222.111102 and of Pseudomonas campeslris Ps. 211.333151. This will
be found useful as a quick method of showing close relationships inside the genus,
but is not a sufficient characterization of any organism. The descriptive chart of
the Society of American Bacteriologists of which the above decimal system forms
a part will be found useful in the detailed systematic study of the bacteria. It was
prepared by F. D. Chester, F. P. Gorham and Erwin F. Smith, appointed as a
committee on methods of identification of bacterial species. Their report was
endorsed by the society at the annual meeting, December, 1907.
LESSON 28
The and other fungous organisms, as out-

detailed investigation of the bacteria
lined below, can be undertaken only after the student has become acquainted with
the cultural methods given in another section of this handbook, but the table adopted
by the Society of American Bacteriologists is given below, because it fits into the
general discussion and study of the classification previously given.
I. MORPHOLOGY.
1. —
Vegetative Cells. Medium used
temp age , , days
Form, round, short rods, long rods, short chains, long chains, filaments, commas,
short spirals, long spirals, Clostridium, cuncate, clavate, curved.
Limits of size
Size of majority
Ends, rounded, truncate, concave.
I
Orientation (grouping)
Agar I Chains (number of elements)
hanging block I Short chains, long chains.
[ Orientation of chains, parallel, irregular.
2. Sporangia. — Medium used
temp
Form, elliptic, short rods, spindled, clavate, drum-sticks.
Limits of size
Size of majority
Location of endospores, central, polar.
3. Endospores. — Form, round, elliptic, elongated.
Limits of size
Size of majority
Wall, thick, chin.
Sporangium wall, adherent, non-adherent.
Germination, equatorial, oblique, polar, bipolar.
4. Flagella. —No Attachment, bipolar polar, perilrichiate.
How stained
5. Capsules. — Present on
6. ZOOGLCEA, PSEUDOZOOGLCEA.
7. Involution Forms. — On days at
in .°C.
8. Staining Reactions. — 10 watery fuchsin, gentian
i : carbol violet, fuchsin
Loeffler's alkaline methylene-blue.
Special stains
Gram Glycogen
Fat Acid-fast
Neisser.
IL CULTURAL FEATURES
I, 2, 3. Agar Stroke, Potato, Loeffler's Blood-serum. —
Growth, invisible, scanty, moderate, abundant.
Form of growth, filiform, cchinulatc, beaded, spreading, plumose, arbores-

cent, rhizoid (Fig. 225).
Elevation of growth, jial, effuse, raised, convex.

Luster, glistening, dull, cretaceous.
Topography, smooth, contoured, rugose, verrucose.
Optic characters, opaque, translucent, opalescent, iridescent.
Chromogenesis,
Odor, absent, decided, resembling
.Consistency, slimy, butyrous, viscid, membranous, coriaceous, brittle.
Medium, grayed, browned, reddened, blued, greened.

Liquefaction (Loefifler's biood-serum) begins in days, complete
in days.
Agar Stab, Gelatin Stab. — Growth, uniform, best at top, best at bottom,
surface growth scanty, abundant; restricted, widespread.
Fig. 225. —
Types of streak culture, i, Filiform {Bacillus colt); 2, echinulate
(Bacterium acidi lactici); 3, beaded {Streptococcus pyogenes); 4, effuse {B. vulgaris);
5, arborescent {Bacillus mycoides). {From McFarland, after Frost in Schneider,
Albert: Bacteriological Methods in Food and Drug Laboratories, 1915: 89.)
Line of puncture, filiform, beaded, papillate, villous, plumose, arborescent.

Liquefaction, crateriform, napiform, infundibidiform, saccate, stratiform,
begins in days, complete in days.
Medium, fluorescent, browned.]
6. Nutrient Broth.-— Surface growth, ring, pellicle, flocculent, membranous, none.
Clouding, slight, moderate, strong; transient, persistent; none, fluid turbid.
Odor, absent, decided, resembling
Sediment, compact, flocculent, granular, flaky, viscid on agitation, abundant,
scant.
7. Milk. — Clearing, without coagulation.

Coagulation, prompt, delayed, absent.
Extrusion of whey, begins in days.
Coagulum, slowly peptonized, rapidly peptonized.
Peptonization, begins on days, complete on days
.
Reaction, i day , 2 days , 4 days , 10 days

20 days
Consistency, slimy, viscid, unchanged.
Medium, browned, reddened, blued, greened.
Lab. ferment, present, absent.
8. Litmus Milk. — Acid, alkaline, acid then alkaline, no change. Prompt reduction,
no reduction, partial slow reduction.
9, 10. Gelatin Colonies. Agar Colonies. — Growth, slow, rapid.
(Temperature )
Form, punctiform, round, irregular, amoehoid, mycelioid, filamentous, rhizoid.

Surface, smooth, rough, concentrically ringed, radiate, striate.
Elevation, flat, effuse, raised, convex puhinate, umbonate, crateriform
(liquefying).
Edge, entire, undulate, lobate, erase, lacerate, fimbriate, floccose, curled.
Internal structure, amorphous, finely, coarsely granular, grumose, filamen-

tous, floccose, curled.
Liquefaction, cup, saucer, spreading.
11. Starch Jelly. — Growth, scanty, copious.
Diastatic action, absent, feeble, profound.
Medium stained
12. Silicate Jelly (Fermis' Solution). — Growth, copious, scanty, absent.
Medium stained
13. Corn's Solution.— Growth, copious, scanty, absent.
Medium, fluorescent, non-fluorescent.
14. Uschinsky's Solution. — Growth, copious, scanty, absent.
Fluid, viscid, non-viscid.
15. Sodium Chloride in Bouillon.— Per inhibiting growth

cent,
16. Growth in Bouillon over Chloroform. — Unrestrained, feeble, absent.
17. Nitrogen. — Obtained from peptone, asparagin, glycocol, urea, ammonia
salts, nitrogen.
18. Best media for long-continued growth
19. Quick tests for differential purposes

III. PHYSICAL AND BIOCHEMIC FEATURES
I. Fermentation Tubes Containing Peptone

Water or Sugar-free Bouillon, and
8.
—
Vitality on Culture Media. Brief, moderate, long.
9. —
Temperature Relations. Thermal death point (ten minutes exposure in
nutrient broth when this is adapted to growth of organism) °C.

10. Killed readily by drying, resistant to drying.
11. Per cent, killed by freezing (salt and crushed ice or liquid air).
12. Sunlight. — Exposure on ice in thinly sown agar plates; one-half plate cov-
ered (time fifteen minutes), sensitive, non-sensitive.
Per cent, killed
13. Acids produced
14. Alkalis produced
15. Alcohols
16. Ferments. —Pepsin, trypsin, diastase, invertase, pectasc, cytase, tyrosinase,
oxidase, peroxidase, lipase, catalase, glucase, galactase, lab, etc.

17. Crystals formed
18. Effects of Germicides
Substance .
The Society of American Bacteriologists has endorsed a brief characterization

as a part of the descriptive chart which it has published. This brief description is
useful in a comparative study of different microorganisms.
BRIEF CHARACTERIZATION
Mark + or o, and when two terms occur on a line, erase the one which does not
apply, unless both apply.
L.4B0RAT0RY AND TEACHING METHODS 639
Notes. —^The
morphologic characters shall be determined and described from
growths obtained upon at least one solid medium (nutrient agar) and in at least
one liquid medium (nutrient broth). Growths at 37°C. shall be in general not older
than twenty-four to forty-eight hours, and growths at 2o°C. not older than forty-
eight to seventy-two hours. To secure uniformity in cultures, in all cases prelimi-
nary cultivation shall be practised as described in the revised Report of the Com-
mittee on Standard Methods of the Laboratory Section of the American Public
Health Association, 1905.
The observation
of cultural and biochemic features shall cover a period of at
days and frequently longer, and shall be made according to the revised
least fifteen
standard methods above referred to. All media shall be made according to the same
standard methods.
Gelatin stab cultures shall be held for si.x; weeks to determine liquefaction.
Ammonia and indol tests shall be made at theend of tenth day, nitrite tests at
end of fifth day.
Titrate with —n NaOH, using phenolphthalein as an indicator; make titrations
at times from blank. The difference gives the amount of acid produced.
The titration should be done after boiling to drive off any CO2 present in the
culture.
Generic nomenclature shall begin with the year 1872 (Cohn's first important
paper). Species nomenclature shall begin with the year 1880 (Koch's discovery of
the poured plate method for the separation of organisms).
Chromogenesis shall be recorded in standard color terms.
LESSON 29
DIRECTIONS FOR THE STUDY OF PATHOGENIC FUNGI
The directions given below for the study of the fungi which cause diseases in
plants have been made as general as possible so that the student will find enough
flexibility in the outline that it may be applied to description of any of the patho-
genic fungous organisms which may be presented to him in his laboratory or field
work. The use of such directions is with the best teaching methods in this
in line
country at the present time. The student is given the diseased organ or plant for
study and by following the outline an acquaintance is obtained not only with the
diseased conditions of the host, but with the morphologic character of the fungus as
well. Some away from the study of
teachers emphasize the importance of getting
systematic details and concentrating the attention of the members of the class in
mycology upon the plant diseases on the basis of the pathologic phenomena exhibited.
Perhaps this is the best plan with advanced students, who have some knowledge
of the morphology and classification of the fungi, a knowledge which should precede,
it seems to the writer, a more detailed study of these interesting plants. It is recom-
mended to the teacher that this outline be used closely in connection with the study
of the diseases described in part III of this book. The teacher, of course, is at liberty
to select other forms for study as the geographic locality may afford. The following
outline is suggestive of such study, where the heading suggests the question which
the students ask themselves in their examination of the diseased plants.
Serial number of type. Place of collection.

Habitat and soil condition. Date.
Name of host. Common names of disease.
History and geographic distribution.
Additional data (Here may be given the nature and amount of loss).
SYifPTOMS
Under this head should be described the general structural changes (morphologic,
or histologic) which are manifest in the diseased host plant, and which distinguish it
from a healthy individual. They may be treated under the following captions:
1. General appearance of the diseased plant.
2. Change in form of part diseased.
3. Change in taste and odor.
4. Change in color as contrasted with healthy part.

(a) Pallor (chlorosis), yellow or white instead of normal green. (Do such
names and yellows apply?)
as mosaic, calico
(b) Colored spots or areas on leaves, stems, fruits (black, brown, orange,
red, variegated, white, yellow, etc.).
5. Perforation of leaves (shot-hole).
6. Damping-off, wilt, wilting, blight (blossom-blight, body-blight, leaf-blight,
twig-blight).
*
7. Death of leaves, twigs, stems, etc. (necrosis).
8. Dwarfing or atrophy. Several names have come into current use expressive
of such condition, as: curly dwarf, leaf-roll, little-peach, spindling-sprouts.
9. Increase in size: hypertrophy. Measurements should be made of the en-
larged parts as contrasted with the normal and the following names may be found
applicable in the study of the hypertrophy: crown-gall, root-gall, root-knot, root-
tubercle.
10. Replacement of parts by new parts.
11. Mummification, character of.
12. Change in position of organs.
13. Disappearance or non-formation of plant parts.
14. Excrescences and malformations.' The following names may be found

suggestive in the description of excrescences and malformations: Cankers, corky
outgrowths, pustules, rosettes, scabs and witches' brooms.
15. Exudations.
Slime flux.
Gummosis.
Resinosis.
16. Rotting.^ — The following terms are suggestive of some kinds of rot: bud-rot,
collar-rot, crown-rot, foot-rot, heart-rot, root-rot, stem-rot and the following par-
ticular kinds given prominence here.
LABORATORY AND TEACHING METHODS 64T
Dry-rot.
Soft-rot (Gangrene).
Black-rot.
White-rot.
The incidental or experimental evidence of disease is indicated by marks or

signs. Such signs are usually afforded by the fruiting or vegetative part of the
pathogenic organism. Such terms as mildew, mould, ooze, rust and smut are indica-
tive of diseased or parasitic conditions.
General Suggestions. — In the report which is made by each student following
the above outline, drawings should, as far as possible, accompany the descriptions.
ETIOLOGY
Common Name of Pathogen.
Scientific Name.
Family.
Pathogenicity.
Additonal Data.
Cultural Character of Organism.
—
Note. In case the pathogenic organism is bacterial the directions for its
study have already been given as recommended by the Society of American Bac-
teriologists. As the outline of the Society is the outcome of years of study, it
should be followed in all cases, but study
in addition the following directions for the
of parasitic plant organisms should be kept in view by the mycologic student.
Isolation of organism in pure culture. Directions have been given for the manu-
facture of culture media and for the isolation of fungi in pure culture. These should
be followed.
Inoculation of pure culture into healthy host plants.
Recovery of organism in pure culture.
life history
The Primary Cycle. — Nature of mycelium (septate, or unseptate; presence or

absence of haustoria (nature); intercellular, or intracellular hyphae; color; contents;
penetration and destruction of host cells = pathogenic histology of host).
Kinds of spores (sexual or non-sexual; conidia; pycnospores; oidiospores;
chlamydospores; ascospores; zygospores; oospores; urediniospores; teliospores;
aeciospores; basidiospores, etc.).
Sizes, shapes and color of spores.

Importance in life cycle.
Pathogenesis of primary stage.
Saprogenesis.
The Secondary Cycles. —The same order of procedure should be observed in the
study- of the secondary cycles as in the examination of the primary.

41
Influence of Soil Factors.

Influence of Climatic Factors.
Control.
Quaranlinc measures.
Spraying.
Remedial measures (dressing wounds and soil amelioration).
Breeding (selection of resistant strains and crossing).
Eradication (burning of diseased plants, cultivation of soil by rotation;
disinfection).
Literature Relating to Disease and Organism. — The citations which are

given in this section can be arranged with reference to their importance and with
some view of theabove outline of study. For example, papers dealing with the
disease in general, with themorphology of the fungus, with the method of control,
might be listed separately under one of the above heads. It is important for the
student to get acquainted with the literature of a subject; otherwise he cannot
appreciate what has been done in his particular field of scientific endeavor.
A bibliography should be made.
\
CHAPTER XXXVIII—LABORATORY AND TEACHING

METHODS (CONTINUED)
LESSON 30
Inoculation Experiments. —The experiments recorded below need not be rigidly
followed by the mycologic teacher. Other organisms and other hosts can be used
just as satisfactorily. The types used must be determined by locality and by other
considerations of cultural methods and laboratory facilities. The directions below
may be taken as samples.
Potato Rot{Fusariiim trichothecoides). —
Take Green Mountain potato tubers and
sterilize surfaceby soaking in 2 per cent, formalin for two hours. The tubers are
then held with towels that have been boiled in water, and are wrapped in these steri-
lized wet towels after having been inoculated with Fusarium trichothecoides by
pricking the surface of the tubers and dipping them in distilled water which holds ^
the spores of the fungus in suspension. The potato tubers wrapped with wet towels
are then surrounded with oiled paper and kept at a temperature not lower than
10° to 1 2°C. Tubers of several varieties can be used, such as Up-to-date, Early Rose,
Irish Cobbler. If the inoculation has been successful, results will be noted in ten
to fifteen days. A transfer to potato slant test-tubes will result in a fungus which
has powdery-rosy appearance. Consult Jamison, C. O., and Wollenweber,
H. W.: An External Dry-rot of Potato Tubers caused by Fusarium trichothecoides.
Journal of th& Washington Academy of Science, II, No. 6, March 19, 191 2.
After the normal lesions have been obtained and the fungus studied mor-
phologically under the microscope, take small slices of potato tuber showing healthy
and diseased tissues in proximity and fix in chromacetic acid. Wash ofi the fixa-
tive in running water, and carry through the alcohol, etc., to paraffin. After im-
bedding in paraffin, section and mount as usual (see Lesson 42).
Crown-gall {Pscudomonas tumefaciens) (Fig. 227). —
Inoculate the stem of a
geranium (Pelargonium zonale) with the organism in pure culture by first
washing the stem at the intended point of infection with i per cent, formalin and
then with distilled water. Place some of the pure culture on the stem by means of
a platinum needle and prick the organism into the stem with a sterile needle mounted
in a wooden handle. The part of the geranium stem selected should be a young
actively growing leader (consult the Bulletin of Erwin F. Smith, and the book
of DuGGAR, Diseases of Plants, pp. 114-118).
This organism can be successfull}^ grown on beef agar which is made as follows.
To 1000 c.c. of peptonized beef bouillon add i per cent, of agar flour, steam three-
quarters of an hour and cool down below 6o°C. Then add neutralized white of two
eggs to clarify. Made to -f 15 Fuller's scale by adding 4NaOH. The test-tubes
are autoclaved fifteen minutes at iro°C.
643
•
644 LABORATOEY EXERCISES
For this and other experiments consult Melhus, T. E.: Culture of Parasitic
Fungi on Living Hosts. Phytopathology, ii: 197-203, October, 1912.
—
Pear Blight (Bacillus amylovoriis, Burrill) (Fig. 228). Take some pear twigs
long enough to be accommodated easily under an ordinary bell jar. Cut off
these stems under water and transfer to a jar under water, so that the cut ends are
not exposed to the air. Then make slanting cuts at the upper end of the twigs
with a sterile knifeand inoculate the cut ends with the organism. Cover the twigs
and jar in which they are placed with a bell jar, as shown in the accompanying
Fig. 227. —
Crown gall artificially produced in greenhouse of University of Penn-
syl by inoculation of Pelargonium zonule with Pseudononas tumefaciens. {Photo
by Charles S. Palmer.)
illustration. Note the result of the inoculation on the tissue of the twigs and on the
health of the leaves. Consult Duggar, B. M.: Fungous Diseases of Plants, pp.
121-129.
Lettuce Drop (Sclerotinia Libertiana, Fuckel.).—Lettuce leaves may be in-
oculated by means of the sclerotia of fungus,' or by the mycelium laid upon the sur-
face of scarified areas of the leaf. As inoculation produces a virulent form of the
disease control, plantsof lettuce should be kept for comparison (Duggar, pp. 190-200).
Wilt of Sweet Corn {Bacterium {Pseudomonas) Stewarti E. F. Sm. (Fig. 229). —
This organism was furnished on beef agar and is best inoculated by applying small
quantities of a pure culture to a stem ofyoung sweet corn and then pricking it in by
means of a sterile needle. Some have inoculated the young sweet corn plants
by placing the organism in the drops of water which exude from the tips of the corn
leaves early in the morning, but the inoculation by means of needle pricks is more
certain. Sections should be made of the stem at various stages of growth after
inoculation. This is done by using a number of plants. Free-hand sections, or
paraffin sections, will show the presence of the organism in the vascular bundles.
Stain with carbol fuchsin (Duggar, pp. 111-113).
Fig. 228.
J^
— Arrangement of
W
experiment for inoculation of pear twigs with blight
organism, Bacillus amylovorus.
LESSON 31
Black-rot of Cruciferous Planls {Bacterium (Pseudomonas) campesiris, Pammel) (see

Smith, Erw. F.: Bacteria in Relation to Plant Diseases, pp. 300-334; Duggar,
B. M.: pp. 107-111). —This organism is best inoculated into the stem of young
cabbage plants below the upper last three leaves, because of the tendency of these
leaves to drop off before the disease has progressed to
its fullest extent. The stem
washed, the organism is smeared on at the point of inoculation and pricked
is first
by a sterile cambric needle into place. It is recommended that several sections

be made, and that to secure the several stages, a number of different inoculations
be made.
Clicstmil Bliglit {Endolhia {Diaporlhe) parasitica (Murrill) Anderson). — Inocula-

tion into the chestnut tree should be made made by
into scarifications of the bark
means of a sterile scalpel. The bark should be washed before inoculation by means
of a weak formalin solution followed by distilled water. The summer spores can
be rubbed into place by means of a sterile platinum needle.
AppeVs Potato Rot {Bacillus phyto-
phthorus, Appel.). —
This organism read-
ily grows on beef agar. It is inocu-
lated into washed parts of the potato
stem by smearing some of the culture
on the stem and pricking into place by
means of a sterile cambric needle into
the young growing tissue.
LESSON 32
Sleepy Disease of Tomatoes {Fusarium

lycopersici Sacc). —This organism can
be cultivated on steamed rice, or on
potato slants. Inoculate just above the
lower leaves of the young stem by first
washing the stem with distilled water.

Place some of the culture on the part
of the stem to be inoculated and prick
the fungus into the stem with a sterile
In ten to fifteen days, the
needle.
tomato plants begin to wilt and in
three weeks the diseased conditions are
unusually good for study. The culture
growths show pale orange spore masses
and a whitish mycelium. The tomato
variety Consate is not susceptible.
Wollenweber used the variety Stone
and found it satisfactory.
Egg Plant Wilt {Verticillium albo-
Fig. '229. Young corn plant showing ^ ^ ,
^ ^v 1 ^1 i
atrum) .—Inoculate the hypocotyl near
places for inoculation th Pseudomonas
Stewarti. or below the soil level with spores sus-
pended in water of a ten days old cul-
ture. Egg plants of any age may be inoculated. Black sclerotia are found in
from ten to fourteen days after the inoculation. This organism is readily grown
on potato slants.
Wilt Disease of the Cotton Cowpeas and Watermelon {Neocosmospora vasinfecta
(Atkinson) E. F. Sm.).— See Duggar, B. M.: Fungous Diseases of Plants, pp. 233-
239; also Smith, Ekw. F. Wilt Disease of Cotton, Watermelon and Cowpeas. Bull.
:
17, U. Division of Vegetable Physiology and Pathology, 1899.

S.
As plants of cowpea, cotton and watermelon have been grown in the greenhouse
and are ready for inoculation, experiments may be tried on all three of these plants.
Inoculation with this fungus should be made into the roots of these plants, just below
the soil of the experimental pots. The soil should be removed and the tops of the
roots laid bare. Inoculation can be made by incisions into the root into which the
mycelium or spores of the fungus are rubbed. After inoculation the soil can be
returned to its place.
LESSON 33
Knot of Citrus Trees {S pliaropsis tumefaciens) .—Successiul inoculations have been

made on lime, pomelo, lemon, tangerine and hardy orange {Citrus trijoliata).
First Method.— Make a small T-shaped cut in the back of a lemon or orange tree
with a sterile knife and insert some mycelium. Smooth the bark down and bind
the stem with raffia to cover the wound completely.
—
Second Method. Inoculate by pricking the stem three times with a sterile cam-
bric needle fixed in a wooden handle, then place a little mycelium over these punctures
and bind with raflia.
Third Method. —Inoculate by cutting off a very small amount (2 or 3 sq. mm.) of
the outer bark, then spread the mycelium over this injury and bind it with irafl&a.
A may elapse before the galls are fully formed.

year
Consult Hedges, Florence, and Tenny, S. S.: A Knot of Citrus Trees Caused
by Sphaeropsis tumefaciens. Bull. 247, Bureau of Plant Industry, 191 2.
Clover Disease.- — Select either red, white, or alsike clover plants somewhere in
a protected place in the garden, or as potted plants in the greenhouse, and inoculate
with Bacillus lathyri. The inoculation may be made by an atomizer. Make a
suspension of the organism in distilled water by means of several loopfuls stirred in
the water. Spray the clover plants with the water and cover with a bell jar for a
few days (J. J. Taubenhaus).
LESSON 34
Sweet Pea Diseases (J. J. Taubenhaus). —

Take several potted sweet pea plants
and spray the leaves by means of an atomizer, which has been sterilized previously
by boiling in water. Make
a suspension of the spores of Glomerella rufomaculans
in water and spray water upon the sweet pea plants which should then be
this
covered with a bell jar. Study the stages of spore germination and spore inoculation
by sacrificing daily one of the sprayed plants.
Inoculate the seeds of sweet pea varieties with cultures of Fusarimn sp. and
Corticium vagum by immersing the seeds in water containing a suspension of fungous
spores. To get this suspension stir up the separate cultures in a sterile watch glass
in distilled water. Then dip the seeds in this water and plant the seeds in loamy
soil in pots for greenhouse culture.Follow the germination of the peas and the
progress of the disease, thus communicated to the plants.
Inoculate the sweet pea by placing a pure culture of root-rot, Thielavia basicola
on the roots of sweet pea plants. Another method adapted to prove the patho-
genicity of the fungus is to sow pure cultures of it on sterilized seeds (seeds treated
with 5 per cent, formalin for one-half hour) in sterile pots and soils.
Inoculate seedlings of sweet pea with Chcetomium crispatum by soaking the seeds
in distilled water containing the spores of the fungus. The seeds should be pre-
viously sterilized, as described above, and the suspension of spores made as above
directed. Healthy plants should be raised from uninoculated seeds as checks on
the progress of the disease in inoculated plants.
Inoculate sweet pea plants with Sclerotinia libertiana by introducing pieces of
the fungus into pots in which sweet peas are growing. Have a potted plant as a
check and cover both plants with a bell jar in order to imitate the moisture condi-
tions of the greenhouse. After four to si.x days, wilting of the inoculated plants
will be noted, while the check remains in a perfectly healthy state.
LESSON 35
Experiments with Artificial Wounding of Plants.

1. Take any herbaceous plant such as hyacinth, snowflake, daffodil, and by
means of a pair of scissors make a short cut into the tissues of the leaves of these
plants, into enough of leaves, so that a serial study can be made of the formation
of healing tissue. Pieces of the leaf are taken from time to time and sectioned by
any of the methods described in Lesson 42.
2. Take any living shrub or tree and make the following cuts:
(a) With a knife cut out a thin longitudinal piece of bark down to the cambium.
{b) Make an irregular tear in the bark by removing a small piece down to the
wood.
(c) Cut out a ring of bark half way around the stem.
(d) Make incisions into a pine tree and by means of sections study the flow of
resin and the healing operation.
(p) Make incisions into the ordinary rubber plant Ficus elastica, and study
with sections the effect of the injury on the cells affected.
(/) Make incisions into any of the woody euphorbiaceous plants of the greenhouse
and study the injuries produced in a similar analytic manner.
3. Cut out larger pieces of bark from deciduous trees and shrubs and by sections
study the formation of cells. By several trips to the fields much of the material
illustrating the healing of wounds can be obtained making of sections and
for the in
all stages of development without waiting for the slow development of new tissue in
the experimental plants. Cut with sliding microtome.

Note the formation of tyloses in many of the woody stems studied. Linden is an
especiallygood tree to show their formation.
Study callus formation of various cuttings, for e.xample, Ficus, Geranium, Ostrya,
Populus, Quercus and Ulrnus. Place the ends of these cuttings in different media,
as follows:
1. One end in water, the other end in dry air.
2. One end in water, the other end in moist air.
3. Both ends in moist air.

4. Both ends in water.
5. One end in moist air, the other in dry air.
6. One end in water, the other in moist sand.
:
7. One end in moist air, the other in sand.

8. Two ends in wet sphagnum.
9. One end
wet sphagnum, the other in moist air.
in
10. One end
in wet sphagnum, the other in wet sand, etc.
Try wounding the cotyledons oiPhaseolus, Vicia, etc.; also young seedling plants.
Use plaster casts to envelope the cut ends. Cf. Tittman: Physiologische Unter-
suchungen uber Callusbildung an stecklinger holziger_ Gewachse. Pringsheim
Jahrb. fur wissensch. Bot., xxvii: 164, 1895.
After securing callus under experimental treatment, then cut, stain and mount for
microscopic study. See Kuster, Ernst:, Pathologische Pflanzenanatomie, 2d.
Edition.
LESSON 36
Gas Injuries. — See Exper.

Rec, xxx, 131, February, 1914.
Sta.
Take and introduce into the soil by means of the hole,
a series of potted plants
in the pot bottom different quantities of illuminating gas by means of a rubber tube
connected with the gas pipe. Note the effect of the illuminating gas on the health
of the plants. Set willow cuttings in water treated and untreated with gas; note
the effect.
Take another set of potted plants and place them beneath bell jars, as follows:
Plant A beneath a bell jar with a beaker of water containing illuminating gas
introduced into the water from the gas pipe.
Plant B beneath a
bell jar into which free gas is conducted by a rubber pipe from
the gas Stone, G. E.: Effects of Illuminating Gas on Vegetation. 25th
jet. Cf.
Annual Report Mass. Agric. Exper. Sta., 1913: 13-28; The Effect on Plant Growth
Carbon Dioxide. Science, new sec. xl: 792, Nov. 27, 1914.
of Saturating a Soil with
—
Smoke Injuries. See Clevenger, J. F. Mellon Instit. Bull. No. 7.
:
Take a series of potted plants of different species and expose them to smoke
conducted to them by means of glass tubes or rubber tubes from the receptacle
where the smoke is generated. Study sections of the smoke-injured tissues.
Tobacco smoke may be tried on tender plants likewise. Consult Bakke, A. L.
The Effect of Smoke and Gases on Vegetation. Iowa Academy Sciences, 1913
(xx): 169-188.
As to smoke injuries, consult also Bakke, A. L.: The Effect of City Smoke on
Vegetation. Bull. 145, Agric. Exper. Sta. Iowa State Coll. of Agric. and Mech. Arts,
October, 1913. See also Knight, H. I. and Crocker, Wm.: Smoke and Gas Poison-
ing. Bot. Gaz., May, 1913: 337-371-
—
Acid Injuries. Treat plants with dilute solutions of various acids and note
their effect on the leaves and flowers. The common morning glories, Ipomcea
purpurea, are useful for this purpose.
Raise some morning-glory plants to flower and treat with dilute acids by spray-
ing with an atomizer. Cf. Stone, George E.: The Influence of Various Light
Intensities and Soil Moisture on the Growth of Cucumbers and their Susceptibility
to Burning from Hydrocyanic Acid Gas. 25th Annual Report. Mass. Agric. Exper.
Sta., 1913: 29-40.
LESSON 37
Enzyme Diseases. — Study these diseases of green plants by taking a series of
leaves of various variegated Anthuriums and other greenhouse species and treat
them as follows: The leaves to be tested are to be boiled for about one minute in
water, when they should be flaccid and free from intercellular air. They are then
placed in methylated spiritwarmed to 50° to 6o°C. : cold spirit will remove the chloro-
phyll, but not so quickly. To produce the iodine reaction, place the decolorized
leaves in alcoholic tincture of iodine, dilute with water to the color of dark beer.
In a few minutes they will be stained, and after washing in fresh water, they should
be spread out on a white plate so that their tint may be well seen. When full of
starch they are almost black, and with less amount of starch, the color sinks through
purple, gray and greenish-gray to the yellow tint of starchless leaves (Sach's method).
In Schimper's method prepare strong chloral hydrate by dissolving the crystals
in as much distilled water as will just cover them. The solution is now colored by
the addition of a little tincture of iodine and is ready for use.
Discoloration of Cut Pieces of Plants. — Cut slices of fresh potatoes and expose them
to the action of the air. Also grate some of the material and test the rapidity of
discoloration.
Take similar pieces and place them in distilled water for twelve hours. Then
expose the cut pieces to the air, and note the result.
These same experiments can be performed with various toadstools and fleshy
.
fungi, when these are in season.

Bibliography. —Allard, H. A.: The Mosaic Disease of the Tobacco. Bull. U. S.
Dept. Agr., No. 40, pp. 33, Jan. 15, 1914-

LoEW, O.: Catalase. U. S. Dept. Agr., Report 68.
Stone, Geo. E. Mosaic and Allied Diseases with Especial Reference to Tobacco
:
and Tomatoes. 25th Annual Report Mass. Agric. Exper. Sta., 1913: 94-104.
Woods, A. F.: Mosaic Disease of Tobacco. U. S. Dept. Agr., Bureau of Plant
Industry, Bull. 18.
Chlorosis. —
Grow vetches and peas in nutrient solution; add 2 per cent, calcium
carbonate, when chlorosis immediately appears, even if iron sulphate is present in
the solutions. A few days in iron nitrate will cause the return of the green color.
In treating plants for chlorosis, a 0.2 per cent, solution of iron nitrate sprayed on the
leaves gives good results.
Where pineapples can be grown in the greenhouse or the open the following facts
will suggest a line of experiments with them and their chlorosis.
Chlorotic pineapples in Hawaii. occur on acid or neutral soils that average 5.0
per cent. Mn304 and 0.5 per cent. CaO. Chlorotic pineapples in Porto Rico occur
on soils containing from 2 to 80 per cent, carbonate of lime and no manganese.
That the is induced by the carbonate of lime was proved by
chlorosis in Porto Rico
direct experiment. which normally produced healthy pineapples were made
Soils
to produce chlorotic plants by the admixture of carbonate of lime from different

sources. We may thus speak of one as a manganese-induced chlorosis and the other
as a lime-induced chlorosis. The lime chlorosis has been shown to be due to a lack
of iron in the plant, caused by the carbonate of lime diminishing the avaflability
1
LABORATORY AND TEACITING METHODS 65
of iron in the soil. M. O. Johnson at the Hawaiian Experiment Station has shown
that the chlorosis of pineapples occurring on highly manganiferous soils can be cured
by spraying the leaves with ferrous sulphate, similarly in Porto Rico the disease due
to calcareous soils can be cured by the application of iron salts.'
LESSON 38
—
Study of Mislldoe. Procure living material of the American mistletoe (Phora-
dendron flavescens) or European mistletoe {Viscum album) and make sections with
the sliding microtome of the stem of host and the parasitic roots of the parasite
and study in detail the association of host and parasite (Figs. 119, 120, 121).
This method of study can be used with Loranlhus Sadebeckli on Citrus niedica.
See Klebahn, Dr. H.: Grundziige der allgemeinen Phytopathologie, 191 2: no.
Cf. TuBEUF, C. von: Infektionversuche mit der rotfriictigen Mistel. Naturw.
Jahrb. Forst. und Landw., xi: 51; Bot-Centralblatt, 123: 293.
Dodder. — Gather material of Cuscuta, Orohanche, Gerardia, Lathraa and other
parasites, and study their anatomy as connected with the anatomy of the hosts on
which they occur (Figs. 117, 122, 123).
The writer has frequently made sections of the stems of the Jo-Pye weed, Eupa-
torimn purpureum, parasitized by Cuscuta Gronovii. These sections were made with
the sliding microtome and have been kept in 50 per cent, alcohol until ready for use.
As class exercises they have been double-stained with safranin and methyl green,
which brings out the relationship of host and parasite very nicely. Finally the
sections have been mounted in balsam and drawn by each member of the class.
LESSON 39
—
Wire Worms in Plants. As the subject of the injurious effects of animals on
plants is a large one and belongs rather to entomology and other departments of
Zoology only one case will be studied here.
—
Nematode Infection of Plants.- Secure material showing the root infection of
horticultural plants by the nematode worm, Heterodera radiciccla. Make sections
showing relation of parasite to host.
Take healthy plants and infect them by transplanting into a soil containing the
eggs or the live round worm. Study entry of the parasite into the hosts and by
paraffin, celloidin or sliding microtome sections, study the relation of the parasite
and host plants.
Similarly, a study of insect galls can be made and their anatomy studied accord-
ing to the description of galls previously given in the second part of this book.
Such a study of galls should be encouraged by the teacher, wherever time and the
arrangement of the courses makes it practicable to do so.
^ GiLE, P. L.: Chlorosis of Pineapples Induced by Manganese and Carbonate
of Lime. Science, new ser., 44: 856, Dec. 15, 1916. Maze, P., Ruot, M. and
Lkmoigne, M.: Calcareous Chlorosis ofGreen Plants: The Role of Root Excretions
in the Absorption of Iron in Calcareous Soils. Compt. Rend. Acad. Sci. (Paris),
157 (1913), No. 12, pp. 495-498 (Exper. Sta. Rec. xxix: 826).
LESSON 40
Relation of Light to Pathologic Conditions. — While light plays an important part

in the development of normal tissue, a lack of it is responsible for man}' abnormal
conditions, and there are a number of diseases common to plants under glass which
are traceable to insufficient light. Plants, such as cucumber, grown under the poor
light common to November and December, have leaves of poor color, slender and
elongated petioles, and little mechanic or resistant tissue, and when subjected to the
bright sun in the early spring every plant in the house will wilt. Poor light also
renders cucumber plants more susceptible to powdery mildew and often causes the
tender edges of the leaves to wilt, turn brown and die. The larger number of leaves
produced in lettuce plants prevent light from reaching the stem, and stem-rot (Sclero-
tinia) or "drop" could undoubtedly be prevented, if the stem were continually
exposed to sunlight. The leaf blights of chrysanthemum and tomato, caused by
Cylindrosporium, are associated with insufficient light and circulation of air at the
base of the stem. Cf. Stone, George E.: The Relation of Light to Greenhouse-
Culture. Bull. 144 (July, 1913), Mass. Agric. Exper. Sta.
—
Experimental Work. Grow cucumbers and lettuce plants from seed and expose
the potted plants to various light intensities in the greenhouse by shading with
several thicknesses of glass, by placing in shaded places in the greenhouse, by growing
next to the glass in the best lighted places. Note the effect on the growth and general
health of the plants. Grow morning glories in pots during winter and study growth.
Etiolation and the Health or Vigor of Plants. —
In order to study the tonic influence
of light upon a plant, we must study its growth in darkness. We find that a plant
grown in the dark is modified both in form and structure. The woody and scleren-
chymatous elements are much reduced, and the parenchyma of the cortex is in-
creased in bulk. The stem becomes very much elongated and remains slender.
It is more succulent than a normal stem, and bears extremely small leaves which grow
out from it at a more acute angle than those which rise upon a normally illuminated
stem. The reaction of its sap is much more acid. The chloroplasts do not become
green, the pigment, which they contain, known as etiolin, being a pale yellow. In
the leaves, the differentiation of the mesophyll into palisade and spongy parenchyma
does not take place. Plants thus affected by darkness are said to be etiolated.
Experimental Work. —
Grow the following plants in light and in total darkness:
Arisama triphyllum, Asparagus officinalis, Caladium esculcntum, Castanea dentata,
Aesculus hippocastanum, Hyacinthus, Onoclea sensibilis, Osmunda cimtaniomea,
Polystichum acrostichoides, Quercus rubra, Sarracenia purpurea, etc. Contrast
influence of etiolation by a determination of water content, dried material, ash,
starch (by iodine method) duration of etiolated organs and plants, structure of leaves,
development of emergences, stomata, lenticels, collenchyma, schlerenchymatous
and other histologic structures. Sections can be made by paraffin and celloidin
methods, etc.
LESSON 41
Withering, or Wilting of Plants. — When the amount of water given off by plants
in transpiration is excessive, the leaves and branches lose their turgescence, become
flaccid and droop, in other words they wilt, or wither. This withering may be due
to the lacli of water in sufficient quantities, in the soil, or it may be due to the pres-
ence of salts of high osmotic equivalent in the soil, which render the absorption of
water difficult, or impossible. Plasmolysis may induce wilting.
Experimental Study. —Take two potted plants and wrap the pot in rubber dam,
or oiled paper, so as to cover the pot and soil to prevent evaporation from their
surfaces. Weigh both potted plants carefully. Water one each day with a meas-
ured quantity of water and let the other remain unwatered until the plant begins to
wilt, then weigh it carefully to determine the amount of available water transpired.
Then knock out the plant and weigh the soil after drying in an oven to determine
the amount of hygroscopic water present.
We now make the following very instructive experiment with Hdianthus tuhero-
SKS. We bend down a long shoot without separating it from the plant, and without
cracking it, so that a portion 20 cm. from the summit dips into water contained in a
vessel placed below it, the summit of the stem and the leaves not being wetted.
We cut through the stem with a sharp knife under water, so that the cut surface
remains under water. Our shoot keeps fresh for days, while other Helianthiis
shoots cut off in the and then at once placed in water, rapidly wither. We may
air,
make them turgescent again by placing a withered shoot in the shorter limb of a
U-shaped glass tube containing water fixed in place in the tube by a rubber
cork fitted air-tight about the stem. Mercury is now poured into the longer limb
of the tube and its pressure is sufficient to revive the withered shoot. Consult
Shive, John W. and Livingston, B. E.: The Relation of Wilting Plants. The
Plant World, No. 4, April, 1914: 81-129.
Plasmolysis and Wilting. —
Prepare 250 c.c. of 0.5 gram-molecular (M) solutions
of potassium nitrate and of sodium chlorid as stock solutions. From these solutions
make dilutions in small vials, capacity about 25 c.c. to contain the following strengths
ofeach of the above solutions, namely o.io, 0.20, 0.30, and 0.40 molecular (M); also
one vial with distilled water as a control. In each of the dilutions place a seedling
of some plant (root as nearly entire as possible) with delicate stem or leaf stalks,
such as lettuce, radish or mustard. Water plants can also be used, such as Elodea
gigantea, Vallisneria spiralis, Trianca bogotensis and the staminal hairs of Trades-
cantea and the filaments of Spirogyra nilida. Observe the dilutions in which wilting
occurs and note the time required in the solutions in which it occurs. Compare
the equivalent strengths of the two salts (The Country Gentleman, Dec. 6, 1913:
1781).
LESSON 42
—
Methods of Sectioning. By the time that this lesson is reached some of the plants
which have been wounded or have been inoculated with the various bacterial and
fungous organisms, or have been treated in various ways experimentally, will begin
to show growth reactions. Such material can be studied by the making and mount-
ing of sections. The sections can be made in one of three ways: (i) By free-hand
sectioning, the razor ground flat on one side being held in the hand; (2) by the slid-
6S4 LAJ30RATORY EXERCISES
ing microtome (Fig. 230); (3) by the rotary microtome, the material having been
imbedded in paraffin. If desirable, the material to be cut on the sliding microtome
can be prepared by the celloidin method. Where the sections to be made are of
woody material they can be cut directly on the sliding microtome, and the sections^
as fast, as they are cut, should be placed in 50 per cent, alcohol. Where free-hand
sections are used they should be placed immediately in 50 per cent, alcohol.
Crlloidin Method. — It is customary to use two solutions of celloidin, a "thick"
and a "thin." The thick solution (about 10 or 12 per cent.) should have the con-
sistency of thick syrup. The thin may be made by mixing equal parts of thick
and ether alcohol. The material inoculated as described in the preceding lessons is
fixed in chrom-acetic acid solution prepared as follows.
Chrom-acetic Acid Fixative.
Chromic acid, i gram

Glacial acetic acid, i c.c.
Water, 98 c.c.
Fle.mming's Fluid (Weaker solution).

[ r per cent, chromic acid, 25 c.c.
A. { 1 per cent, acetic acid, 10 c.c.
[ Water, 55 c.c.
B. I per cent, osmic acid, 10 c.c.
Keep the mixture A made up, and add B as the reagent is needed for use, since
it does not keep well.

Wash the fixed material carefully in running water for several hours and put into
30 per cent, alcohol, then by successive steps into 50 per cent. 75 per cent., 95 per
cent, and absolute alcohol. After dehydrating in absolute alcohol, the succeeding
steps are taken.
1. Ether alcohol, i to 2 days.
2. Thin celloidin, 2 to 6 days.
3. Thick celloidin, 3 to 10 days.
—
Use of Alcohols and Celloidin. The celloidin is dissolved in equal parts of ether
and absolute alcohol about i part by weight of celloidin to 15 parts of the solvent.
After the material is thoroughly penetrated by this solution, it is passed to a stronger
solution, containing i part of celloidin to 11 parts of the solvent and finally to a
solution containing i part of celloidin to 8 parts of the solvent.After remaining a
suitable time in the last solution, the object is ready for imbedding. For this
purpose, a paper strip may

be wound tightly about the end of a small block of suit-
able size and material, so as to form the sides of a box open above, with a bottom
the end of the block of wood. This box is now filled with the thickest celloidin
solution, and in it the object is placed and oriented carefully by needles wet with the
ether-alcohol mixture. As soon as a strong film has developed over the surface of
the celloidin, the whole block of material is plunged into 80 per cent. After the
celloidin has hardened in the alcohol, the paper ring is removed and the mass is
trimmed to the desired size.
In cutting, the block is clamped in the sliding microtome, where the knife is set
obliquely, so that the celloidin sections may be cut with a long drawing stroke.
The knife and top of the block should be kept wet with 80 per cent, alcohol, and as
rapidly as the sections are cut, they should be placed in the alcohol (Fig. 230).
The sections are attached to the slide by placing the slide in a closed chamber
over ether. The ether vapor quickly dissolves the celloidin to cause the sections
to adhere firmly to the slide on removal from the chamber. After the removal of
the celloidin, the sections can be stained with appropriate stains. For mounting
inCanada balsam, celloidin sections may be cleared with a mixture of 3 parts xylol
and 1 part phenol.
Paraffin Method. —
The fixing and dehydrating of material for imbedding in
parafl&n is performed in a manner similar to that for work with celloidin up to the
dehydration in absolute alcohol. The following schedule should be followed
subsequently.
Transfer from absolute alcohol to pure xylol, allowing at least two hours in each
of the following three mixtures, î alcohol +
3^^ xylol; 3^2 alcohol xylol; +M
î xylol + H
alcohol, xylol. Add to the mixture of paraffin dissolved cold in xylol.
Place in melted paraffin in the bath, kept at 55°C., two to twenty-four hours as
convenient. Imbed in paper capsules, or in small shallow glass dishes. Section
with rotary microtome; about 6 to lo/i is a good thickness.
See Lesson 43 for details of cutting frozen section by the microtome and the
method of freezing each section. Lesson 43 may be introduced here.
—
Fastening of Sections to Slide. After cutting, fasten section to slide by using
Meyer's albumen, or by the process of drying on the slide after treatment with tepid
water to remove the wrinkles.
Dissolve off paraffin in xylol.
Pass down through 100 per cent., 95 per cent., 85 per cent., 70 per cent., 50 per
cent., 30 per cent., alcohol, thirty seconds each.
Delafield's ha?matoxylin, fifteen minutes.
Rinse in water five minutes.
Pass up through 30 per cent., 50 per cent., 70 per cent., 95 per cent., and absolute
alcohol.
Put in xylol at least one minute.
Mount in balsam.
Note. — All of the material obtained in the inoculation experiments should be
studied microscopically. The above methods of fixing, imbedding, sectioning and
staining are applicable in all of this work.
If time permits, all of the organisms inoculated in the plants should be recovered
and in pure culture by the methods outlined in Lesson 22. Direct inoculation of
media in plugged test-tubes can be used. A reinoculation of the recovered organisms
is desirable, if time permits the class to undertake such additional work.
LESSON 43
Freezing of Material and Cutting. —Freezing Microtome. —The material may be

imbedded in a thick solution of gum arable which is frozen on a metal plate cooled
to the freezing temperature by conducting under the plate a mixture of ice water
and salt. This is accomplished by filling a glass vessel full of a mixture of ice and
saltand conducting the water from the jar by a tube (.4) through metal a box {B)
on which the sections are placed in the mucilage.
The circulation of the ice-salt water is accomplished by allowing it to drip from

a small orifice at the end of the glass tube C.
The block of frozen mucilage with the contained substance held on the freezing
plate is then cut with the hand microtome or with the design of microtome shown
on the next page.
Or the material may be frozen in the design of freezing chamber shown on page
659 and sectioned by Spencer automatic laboratory microtome No. 880, as indicated
in the accompanj-ing figures. If mucilage is used it can be removed by placing the
sections as rapidly as cut in warm water.

CO2 Freezing Allachmcnt. —The freezing device in this attachment consists of a
small metal cylinder. The object is placed on the flat disk top of the cylinder,
Fig. 231. — Freezing attachment for use of CO2 in freezing microtome.

which measures 36 mm. in diameter, and is frozen by the expansion of the CO2.
This device is connected with the gas cylinder by a flexible copper tube, provided
with a connecting nut for joining to the cylinder and the necessary adapter for fitting
to the microtome. It is furnished also with an extra valve, which can be placed at
either end of the tube.
CO2 gas furnishes the most rapid and convenient medium for freezing specimens
and can be used in this attachment with either the table or physician's microtome
(Figs. 231, 232). An ether attachment is also used (Fig. 233).
LESSON 44
—
Use of Drawing and Projection Apparatus. The author has found it an excellent
training for students to learn the use of the drawing apparatus designed by Edinger,
as well as the new Spencer photomicrographic camera. These pieces of apparatus
can be used for drawing, for projection and for photomicrography.
42-
6s8 LABORATORY EXERCISES.
The Edinger drawing and projection apparatus^ (Figs. 234, 235) projects micro-
scopic objects even under a high magnification directly upon the drawing board so
that the outline can be traced in pencil. The image thus projected can be used for
demonstrating to a small audience and also for photomicrography. For such work
a powerful illuminant is used with a hand-fed electric arc taking 4 amperes. It may
be used with a suitable plug connected with the direct-current house supply (alter-
nating current may be used by special arrangement). The crater in the positive
|_f^||t4LOCo
Fig. "232. — Clinic microtome with freezing attachment.
carbon from which light emanates is brought to coincide with the optic axis of the
apparatus by means of the two screws (o) as in Fig. 234, and the lamp with the con-
densing system K can be moved along the optic axis by the lever G. The distance
between the carbons is regulated by the milled head (6) which if out of reach of
the operator can be turned by the long handle connected to (c). The smaller car-
bon which is placed horizontally should not project into the optical axis, or crater
area of the larger vertical carbon.
The apparatus proper consists of a cast-iron pillar S, Fig. 234, mounted upon a
1 May be had of E. Leitz, 30 East icSth Street, New York City.
rectangular frame into which a drawing board is fitted. The fitting is grooved to
allow the adjustment of the illuminant L by the lever G, the stage 0, and the objec-
tive holder //, the face being graduated to ]î cm. in order that the correct position of
the stage O, which varies according to the objective in use (see Table A), can be
determined. The same table gives the correct size of diaphragm, five accompanying
each outfit, viz.: 12, 18, 24, 32 and 46 mm. diameter. The cover-glass faces the ob-
jective when the slide with object is placed in position. The objective carrier H
which has a rack and pinion for coarse adjustment and a micrometer screw for fine
adjustment occupies a constant position on the fitting B, viz., i cm. from the lower
Fig. 233. — Ether or rhigoline freezing attachment for freezing microtome.

end, but can be removed if necessary. The fine adjustment can be controlled by
a long rod similar to that used for the setting of the arc.
Above the stage two lenses of different foci are mounted in a swing-out {K2,
Fig. 234) which has a sliding focussing adjustment and iris diaphragm, and is so
contrived that either of the condensers or the diaphragm only can be interposed in
the optic axis. The microscope body T can be removed from the fitting M, into which
it pushes, and the triple nosepiece is mounted on a sliding attachment, so that it can
be interchanged from a similar slide carrying the microsummar lenses. The draw
tube should always be set at 152 mm. when working with the nosepiece; otherwise,
at 170 mm. Should the apparatus be required for projection the whole optical
66o LAEORATORY EXERCISES
system can be rotated from the vertical to the horizontal position by j)ulling out the
spring catch E, Fig. 234.
Fig. 234.— Details of Edinger's drawing apparatus. Z, Drawing board; T, micro-

scopic attachment; K\ and A'2 condensers; L, electric lamp attachment.
For photomicrographic work a camera is clamped to the pillar S, Fig. 234, the
plate holder, which will take plates of any size up to 24 by 30 cm., resting on the
LAEORATORY AND TEACHING METHODS 66l
drawing board Z (Fig. 234). Having determined the camera extension required by
means of a special set screw provided, an allowance of 2.8 cm. is made for the
height of the plate above the drawing board. The arm clamping the camera to
Fig. 235. — Edingti s diawm^ icroscopic drawing.
the pillar is then raised until the collar fits over the draw tube of the microscope
body T, or over M, when working with the niicrosummars, thus ensuring a light-
tight connection. It is advisable to support the bellows by the strap pieces shown in
Fig. 236, when extended. Correct focus is determined by the observation of the
image upon a paper surface in place of the usual ground glass.
Fig. 236. — Edmgcr b di i\\int ith altachment for photo-micrography.
The following tables have been prepared with the view of simplifying the use of
the apparatus as much as possible, and the best results can only be obtained when
:
LABORATORY AND TEALHING MKiTlOUS 663
the instructions given for the height of the stage and lamp, and the use of condenser
and diaphragm for each objective, are strictly adhered to
Table A
Objective
Table C
Of the Achromatic Objectives with the Huyghenian Eyepieces at 250 mm. distance
from the Drawing Board
SUGGESTIONS TO STUDENTS
Concerning Notes.
1. The laboratory notes or descriptions should embody only such facts as have
been gathered from your own observation and study of the object. Any collateral
notes written up from lectures or reading should not be mingled with those of your
own observation, but should be kept distinct and under separate headings.
2. The may be gathered first on "scratch
facts observed in the laboratory or field
[)aper" as temporary notesand subsequently be written on the note tablet in per-
manent form; but such temporary notes should be promptly written up and not be
allowed to accumulate.
3. The permanent notes or descriptions should be an original account of your
own observation.The statements should be scrupulously accurate and free from
figurative expression and rhetoric embellishment; the style should be simple, clear
and concise.
4. Frequent reference should be made to the drawings and diagrams which
accompany the study so that these and the notes may be mutually helpful.
5. The ability to give a clear and accurate account of one's own observations
and conclusions is an essential in scientific work, and is also of much value in prac-
tical life.
Concerning Drawings and Diagrams.

1. A drawing is intended to show the size and shape of the object, and the pro-
portions and relations of its parts. In case the drawing is to be smaller or larger
than the object, the size of the object may be indicated by symbols, as for example:
" X î" or " X 4," the former signifying that the drawing is reduced to one-fourth
and the latter that it is enlarged to four times the actual size of the object.
2. A diagram is intended to show only the relation of the parts of the object and
does not pretend to represent their size, shape or structure.
3. In making either drawing or diagram, do not aim at anything ornamental, or
artistic in effect. Let your aim be to represent clearly and distinctly certain facts
of your observation.
4. First, carefully examine the object and have definitely in mind what you wish
to show in your diagram or drawing and omit everything else.

5. Decide in advance what view of the object you wish to represent and the size
of your drawing. If the object be an animal or a plant, represent it whenever

practicable in its most natural position.
6. With a fine-pointed hard pencil, make a very faint outline of the object, step
by step comparing the drawing with the object, and omitting at first all details.
See that the proportions are correct, revising your drawing, if necessary, by sub-
stituting new lines and ignoring or erasing old ones.
7. The details may now be worked. Avoid much shading and omit it altogether
whenever possible. If the drawing is merely an outline it may be improved by trac-
ing Its lines, and the effect of shading may be produced by tracing more heavily
those lines which are opposite the direction of the light.
8. In diagrams no shading is needed, but in many cases the use of flat tints,
produced with colored pencils or preferably water colors is very helpful.

drawings and diagrams should be accurately and intelligibly labeled.

9. All
Generally it is also desirable that the parts of the drawing, especially the parts of a
diagram, be designated in a way that is convenient for reference.
Drawings should be made either entirely in ink, or entirely in pencil, and the
10.
lettering also, which should be uniform, not one style, then another.
11. Large headings should be more especially emphasized by larger letters, and
the lettering of the larger and smaller headings should be of the same style.
12. All drawings presented to the teacher for examination should be placed
between the two sides of a folder of stiff manila paper.
13. The grade of pencil should be determined by the kind of finish or surface of
the drawing paper, but in general for science work, the harder grades of lead, say
from 4H to 6H, are preferable.
14. The name of the student, the number and the subject, as well as the year,
should in all cases be placed on the outside of the manila cover.
Method and Materials of Photomicrography (Fig. 236). The photographic —
plates which best meet the requirements in photomicrographic work with the
Edinger apparatus are Lumier Sigma 9 by 12 cm. plates, or the ordinary 4 by 5
plates. Another good plate is known to the trade as Seed Special 27.
Whatever plate is used, it is placed in the plate holder of the photomicrographic
camera in a dark room, the dull side of the plate being outermost. The holder is
then placed in its proper position in the photographic camera. Before the insertion
of the holder, however, the object to be photographed must be focussed on the
ground-glass plate of the camera until a sharp image is obtained, then the focussing
screw should be moved a trifle, say one of the divisions of the screw, so that the
object is focussed up a slight amount. The light being regulated properly, the
exposure is made by withdrawing the shutter of the plate holder. The length of
time to expose the plate can be determined only by several trials until the operator
learns the length of time by the experience thus gained.
The most satisfactory developer is made as follows:
Rodinol, i part.
Water, 12 parts.
Potassium bromide, 10 drops of 10 per cent, solution.
The advantage of this developer is that the process is sufficiently slow, so that the
operator may be able to study the photograph, as it makes itself evident.
After washing in water, the negative is placed in a rather strong hyposulphite
solution as a fixing bath. The advantage of rodinol over metol is that the develop-
ment is more even and sure.Where the photomicrographs have been made ob-
scure, or where it is desirable to convert them into outline drawings for diagrammatic
purposes the following method can be used.
Draivings on Photographic Prints. —All pen-and-ink drawings of photographic
prints must be made with water-proof India ink after which the photographic part
is bleached out by exposure for a few minutes in water containing cyanide of potash
(i : 500, more or less). The drawings should be exposed in this bath as long as
necessary. If any part of the print refuses to bleach, it should be moistened with
iodine-potassium iodide and returned to the cj'anide bath. It is then passed tlirough
pure water and dried face up on blotting paper in a place free from dust.
Bibliography. —
For details the student is referred to a book by W. H. Walmsley,
entitled, The AB C
of Photomicrography. A Practical Handbook for Beginner.
New York, Tennent and Ward, 1902.
Complete details will be found in Erw. F. Smith's Bacteria in Relation to Plant
Diseases, Vol. i: 130-151; Barnard, J. Edwin: Practical Photomicrography, 1911:
xii -f 322, London, Edward Arnold; Hind, H.Lloyd and Randles, W. Brough:
Handbook of Photomicrography, 1913: xii + 292 with 44 plates. New York, E. P.

Dutton & Co.
Lesson 46
The course mycology will not be complete without the introduction of

in
field trips and excursions which supplement in an important way the laboratory
and lecture work, and which will show the student how mycology touches
practically the sciences of bacteriology, chemistry, engineering, and the other
technologic sciences. Besides the trips into the woods and fields for various
kinds of fungi and to the market houses to collect the fungous diseases of the
food plants sold there, trips can be planned to include slaughter houses, cold
storage plants, meat extract factories and dairies where the cooling, filtration,
Pasteurization, and bottling of milk can be demonstrated. Mushroom farms
should not be omitted, nor should the farms where vaccine and other biologic
products are made be overlooked. Cheese, butter, oleomargarine and soap
factories should be included in the schedule, as well as the sugar refineries.
The are employed should be investigated, such
industrial plants where yeasts
as bread bakeries, beer breweries, wine and pressed yeast factories. The estab-
lishments where pickles, sour krout and vinegar 'aTE^made should not be omitted.
The disposal of the sewage of our large cities will pay inspection. The con-
servation of manure in the city and on the farm, the general problems of soil
mycology and the preparation of silage ought to be introduced by the field
trips. The health laboratories of our large cities should be included in the
itinerary. These are only a few of the places that might be visited profitably
near such large cities as Boston, New York, Philadelphia, Baltimore, Chicago,
St. Louis, New Orleans, Denver, and San Francisco, and smaller places where
manufacturing is important.
References
Bergey, D. H.: The Principles of Hygiene, Philadelphia, 1914.

CouN, H. W.: Bacteria, Yeasts and Molds in the Home, New York, 1903.
Fuhrmann, Dr. Franz: Vorlesungen iiber technische Mykologie, Jena, 1913.
GiLTNER, Ward: Laboratory Manual in General Microbiology, New York, 1916.
Kossowicz, Dr. Alexander: Einfiihrung in die Mykologie der Gebrauchs-und
Abwasser, Berlin, 19 13.
Kossowicz, Dr. Alexander: Einfiihrung in die Agriculturmykologie, Berlin.
Kossowicz, Dr. Alexander: Lehrbuch der Chemie Bakteriologie und Tech-

nologic der Nahrungs-und Genussmittel, Berlin, 1914-
LiPMAN, Jacob G.: Bacteria in Relation to Country Life, New York, 1908.
LoHNis, Dr. F.: Handbuch der landvvirthschaftlichen Bakteriologie, Berlin.
Lafar, Dr. Franz: Technical Mycology, Landon, 1898-1910.
Marshall, Charles E.: Microbiology, Philadelphia, 1911.
Prescott, Samuel C. and Winslow, Charles-Edward A.: Elements of Water
Bacteriology, New York, 3 edit., 1913.
RosEMAN, Milton J.: Preventive Medicine and Hygiene, New York, 1914.
Whipple, George C: The Microscopy of Drinking Water, New York, 3 edit.,
1914.
APPENDIX I
Perhaps what follows may be looked upon by some teachers as hardly forming
appropriate laboratory exercises, and, therefore, should be treated as in the nature
of appendices. In agricultural and horticultural schools, the manufacture and
use of fungicides and sprays may very well form a part of the curriculum designed
for laboratory, and especially for field purposes, where in the experimental farm, or
garden, the spraying apparatus and its construction can well be experimented with
as a regular part of the instruction. Hence the making of sprays is given prominence.
Fungicides. — Definition of Terms. — Fungicides are substances which are capa-
ble of destroying, or preventing, the growth of spores, or the mycelia of fungi. Germi-
cides are those substances used for a similar purpose with germs, or bacteria. Such
materials may be used as a spray, in the form of a powder dusted on the plant, or in
the form of a steep into which the plant, or plant part, is dipped. A substance to
be useful as a fungicide must not only not injure the plant, but must at the same
time destroy' or hold in check the parasite. Usually the material is most effective
when the fungous parasites can be reached directly by the spray. If the fungus
works internally, as the chestnut blight fungus, such fungicides usually do harm to the
host without touching the parasite and are, therefore, ineffectual.
The chemic substances used are naturally of a poisonous character and should
be used with precautions taken to prevent their injurious effects upon human beings.
.\n up-to-date agriculturist, horticulturist, or orchardist considers the use of
fungicides, germicides, or insecticides, as essential, as any of the other major opera-,
tions on the farm.
For convenience of treatment and ease of reference the following fungicides and
insecticides are arranged alphabetically. The formulae have been taken from a num-
ber of reliable sources and they may be considered as dependable in ordinary work.
—
Ammoniacal Copper Carbonate. This is not as good for general purposes as
Bordeaux mixture. It is used instead of Bordeaux when it is desirable to avoid the
spotting of leaves or fruit. It is prepared as follows:
Copper carbonate, 5 ounces.

Strongammonia (26° Baume), 2 to 3 pints.
Water to make 50 gallons.
Dilute the ammonia with about 2 gallons of water, as it has been found that
ammonia diluted seven or eight times is a greater solvent for copper carbonate than
the concentrated liquid. Add water to the carbonate to make a thin paste, pour on
about half of the diluted ammonia and stir vigorously for several minutes: allow it
to settle and pour off the solution leaving the undisturbed salt behind. Repeat
this operation, using small portions of the remaining ammonia water until all the
669
:
670 ADDITIONAL EXERCISES
carbonate is dissolved, being careful to use no more ammonia than is necessary to

complete the solution. Then, after adding the remainder of the required quantity
of water, the solution is ready for application.
Caution. —
Plants likely to be injured by Bordeaux mixture are more susceptible
to the clear light-blue solution of ammoniacal copper carbonate, which upon drying
leaves little or no stain.
Arsenate of lead is one of the best arsenical insecticides. It has in many cases
entirely displaced Paris green orchard spraying, and there are at least three good
reasons for its use.
First. —The arsenate

of lead has great adhesive qualities. It will not wash off
even heavy showers of rain. Some of the experiments at the Minnesota Experi-
in
ment Station showed the presence of this arsenate on the leaf in sufficient quantity
to kill insects, ten weeks after spraying.
Second. —
It can be used in any strength without burning the foliage of the plant
sprayed, except peach leaves which are burned, if it is too strong.
Third. —
It has some fungicidal properties that are increased when added to lime
sulphur. The home-made preparation is made as follows
22 ounces acetate of lead (sugar of lead) dissolved in 2 gallons of warm water in
a wooden pail.
8 ounces arsenate of soda dissolved in i gallon water in another wooden pail.

These two solutions are poured together and make sufficient quantity of poison for
50 gallons of spray.
Arscnite of Lime. —A home-made preparation much cheaper than Paris green
and just as good. It is prepared as follows:
White arsenic, i pound ]
Crystal sal soda, 4 pounds [

Stock solution
Water, i gallon J
Boil these in an iron kettle for twenty minutes until thoroughly dissolved. The
kettle must be kept exclusively for this purpose. The soluble material obtained is
arsenite of soda and can be stored away in jugs or bottles, labeled poison, for future
use. For 40 or 50 gallons of spray, take 1 3^ to 2 pints of this solution, and 4 pounds
of freshly slaked lime. Dilute the lime and stain: then add the stock solution.
Pour into the spray barrel, and it is ready for use.
—
Bordeaux Mixture. This is the most valuable fungicide in use for combating
plant diseases and consists of a mixture of copper sulphate (blue stone) and stone
lime slaked in water. It is used in various strengths.
Standard Bordeaux Mixtures (Fig. 237) (6-4-50 formula).
Copper sulphate, 6 pounds.

Lime, 4 pounds.
Water to make 50 gallons..
This mixture can be used successfully on many plants, but on others like the peach
and Japanese plum, it injures the foliage. It also sometimes russets the fruit of
apples and pears. It can be increased in strength for certain purposes by reducing
APPENDIX I 671
the proportion of water, but the formula given above has been regarded as the
standard with which all others should be compared, at least in experimental work.
—
The 5-5-50 Formtda. Here the preparation consists of

Lime, 5 pounds.
The use of this formula is desirable where the purity of the lime is in doubt, as
it makes certain, with lime of any reasonable quality, that all of the copper is properly
neutralized. The danger of scorching, or russeting fruit is, therefore, less. With-
holding I pound of the copper sulphate also cheapens the mixture by a few cents.
For these reasons the 5-5-50 formula has come to be quite generally used in orchard
spraying. In fact, it has almost replaced the old standard Bordeaux mixture in
spraying for the apple scab, bitter-rot, pear and cherry leaf-blight and similar diseases.
The 4-4-50 and Other Formulas.— The strength of the mixture is often further
reduced by using the 4-4-50 formula, but it is questionable whether it pays to reduce
the strength. For use as a whitewash, a very concentrated mixture, 6-4-20, may
be desirable and for certain diseases Bordeaux mixture can be diluted so as to be
equivalent to 6-4-100.
The form of Bordeaux mixture most harmless to foliage is 3-9-50, having a con-
siderable excess of lime. This may be known as the "peach Bordeaux mixture."
Various modifications of the original Bordeaux mixture have been suggested and
tried. The principal ones, however, are the "soda Bordeaux mixture" and the
"potash Bordeaux mixture." The former consists of 6 pounds of copper sulphate,
2 pounds of caustic soda and 50 gallons of water. The latter is the same except an
equal quantity of caustic potash is substituted for the soda. Other materials are
sometimes added to Bordeaux mixture to increase its spreading power. The most
successful is ordinary hard soap, dissolved in hot water and added at the rate of 4
pounds to the barrel, and this modified Bordeaux mixture is known as "soap
Bordeaux."
Bordeaux Resin Mixture (N. Y. (Geneva) Bull. No. 188, 1900).
Resin, 5 pounds.
Potash lime, i pound.
Fish oil, I pint.
Water, 5 gallons.
Add to Bordeaux as directed below. To prepare a stock resin solution proceed

as follows: "Place the oil and resin in the kettle, heating them until the resin is dis-
solved, then remove the kettle from the fire and allow the mass to cool slightly, after
which the solution of lye is added slowly, the whole being stirred while adding the
lye. After adding the lye the kettle should be again placed over the fire and the
required amount of water added. The whole should be boiled until the solution
will mix with cold water forming an amber-colored solution. Care should always
be taken to have the resin and oil cool enough, so that when the solution of lye or the
water is added the whole mass will not boil over and catch fire.
"Dilute this stock resin solution with 8 parts of water before adding to the
Bordeaux mixture, that is in preparing a 50-gallon barrel of the mixture, the copper
sulphate and lime are diluted enough to make 40 gallons after which 2 gallons of
stock resin solution are diluted to 10 gallons, then added to the Bordeaux."
This solution exceeds ordinary Bordeaux in adhesive properties and has been
highly recommended for asparagus rust.
Method of Making Small QiiantUies of Bordeaux Mixture. Two half-barrel tubs —
are made by sawing a barrel through the middle. One tub is used for the blue-stone
solution and the other for the milk of lime, and each tub should contain 25 gallons.
One man dips the blue-stone solution with a bucket and pours it into a barrel and
another man simultaneously dips up and pours in bucketfuls of the milk of lime.
DIP EQUAL PARTS FROM

-
I
^3ss5^ anp2into3
'EST0KE*^SVrHEN5Tifi
IGOR-
,
0U5LY
fine mesh screen

nelto
leoux
Dipper-* mil-:
"t/se thi5 miiiturf at once m&pi^cr- Sprayer
FiG. 237. — Diagramshowing easy method of making-small quantities of Bor-
deaux mixture. {After Coons, G. H., and Levin, Ezra, Spec. Bull. 77, Mich. Agric.
Coll. Exper. Stat., March, 1916.)
The lime solution should be kept well stirred. If only a single barrel is to be made,
the materials be dissolved in the dilution tubs, but if a number of lots are re-
may
quired the materials can be kept in stock solutions and simply transferred by dipping.
No matter what quantity of mixture is to be made up, it is necessary to strain the
materials through a wire strainer. The best type is made of brass wire with 18 to
20 meshes to the inch (Fig. 237). For details see Waite, M. B.: Fungicides.
U. Farmers' Bull. 243 (1906).
S.
In large operations stock solutions should always be used, as the time required to
dissolve the material is saved. These can be prepared of both copper sulphate and
the lime. Dissolve copper sulphate in water at the rate of i pound per gallon and
lime in the same ratio. Then measure ofT the required quantity of each and dilute
with water before mixing. If possible the dilution tanks should be raised so high
on an elevated platform that the mixture can be conducted by gravity into the
spray tank on wheels or in a wagon beneath. An available water supply is necessary.
APPENDIX 1 673
Testing Bordeaux Mixliirc.—When Bordeaux mixture is properl)' prepared it is of
a brilliant sky-blue color. If the lime is air-slaked, or otherwise inferior in quality,

resulting in a bad mixture, the preparation have a greenish cast, and if this is
will
very pronounced the mixture will injure In order to make certain that
the foliage.
the copper sulphate is properly neutralized by the lime, the yellow prussiate of potash
test may be used. A small bottle containing a 10 per cent, solution of yellow
prussiate of potash can be secured from a druggist. After stirring the Bordeaux
mixture a drop of this solution is allowed to fall on the surface of the preparation.
If free copper is present, the drop will turn reddish brown in color immediately.
Lime should then be added until the brown color fails to appear. If the reaction
is complete, the yellow prussiate of potash solution will remain a clear yellow until
it disappears in the mixture.
—
Bordeaux Mixture and Inseclicides. One advantage of Bordeaux mixture is the
possibility ofadding arsenical insecticides to the preparation and thus of spraying
at the same time for :(,ungous diseases and for the codling-moth and leaf-eating in-
sects. Paris green at the rate of yi pound to 50 gallons of Bordeaux mixture, may
be considered as the standard formula for this purpose. London purple, arsenate of
lead and other arsenicals may Bordeaux mixture may be
be used in the same way.
considered as so much water in the formulas for this class of insecticides. As a
matter of fact, the slight excess of lime in the standard mixture renders it an espe-
cially suitable medium for distributing these insecticides.
Dust Bordeaux Mixture. — This mixture is prepared as follows:
4 pounds of copper sulphate in 4 gallons of water.

4 pounds of lime in 4 gallons of water.
60 pounds of slaked lime dust.
Dissolve the 4 pounds of copper sulphate in 4 gallons of water and slake 4 pounds
of lime in 4 gallons of water, when cold pour the two solutions together simultaneously
into a tub. Allow the resulting precipitant to settle, decant off the liquid, pour
the wet mass of material into a double flour bag, and squeeze out as much water as
possible. Then spread the dough-like mass in the sun to dry. After a day's dry-
ing can be crumbled easily into an impalpable powder by crushing with a block
it
of wood.This powder should be screened through a brass wire sieve having at least
83 meshes to the inch and should be mixed thoroughly with 60 pounds of slaked lime
dust. The lime dust is best prepared by slowly sprinkling a small quantity of water
over a heap of quick lime, using barely enough water to cause the lime to crumble
into a dust. The heat generated will soon drive off the excess of moisture, and the
dust should be passed through a screen of 80 meshes to the inch. This powder is
applied by means of a blower. If desired 4 pounds of sulphate and i pound of Paris
green may be added to each 60 pounds of Bordeaux mixture dust. For details,
consult Waite M. B.: Fungicides. U. S. Farmers' Bull. No. 243 (1906).

Copper Sulphate Wash.

Water, 50 gallons.
:
This is used as a wash on dormant trees, for the prevention of such diseases
as apple scab. It must never be used on trees after the buds have burst.
- Copper Acetate.
Copper acetate (dibasic acetate), 6 ounces.
Water, 50 gallons.
First make a paste of the copper acetate by adding water to it, then dilute to
the required strength. Use finely powdered acetate of copper, not the crystalline
form. It may be used as a substitute for copper carbonate mixtures.
Copper Saccharate. — Consult Freemen, E. M.: Minnesota Plant Diseases, p. 220.
Corrosive Sublimate.
Mercury bichloride (corrosive sublimate), 2 ounces..
Water, 15 gallons.
This is an extremely poisonous mixture and should be handled with great care.
It is very effective against potato scab. It should not be made in tin vessels, as it
corrodes them.
Formalin.
Formalin (40 per cent, formaldehyd), J-2 pound.
Water, 15 gallons.
This is used in treating seed for prevention of such diseases as potato scab.
Iroti Sulphide Mixture. —
This is a new, but very promising fungicide. It was
tried on apples, and gave splendid results in preventing fungous diseases, being non-
injurious to the fruit. In preparing this fungicide, it is recommended that a self-
boiled lime-sulphur mixture be prepared, as later described, except that 10 pounds
of lime and 10 pounds of sulphur are used. The mixture is diluted to 40 gallons,
and then 3 pounds of iron sulphate (copperas) dissolved in about 8 gallons of water,
is added.
Potassium Sulphide (Liver of Sulphur).
Potassium sulphide, 3 to 5 ounces.

Water, 10 gallons.
This is used in place of Bordeaux mixture to avoid spotting of foliage and fruit.
It is considered to be especially effective against powdery mildews. It is quite ex-

tensively used in greenhouses and on shrubbery.
Sulphur. — Is used as a fungicide in a pure state. The flowers of sulphur is the
highest and usually the purest chemically. It is dusted on plants as a remedy for
mildew, especially the rose mildew and the powdery grape mildew.
Sulphur and Resin Solution. —It is made up as follows
Sulphur (flowers, or flour), 16 pounds.

Resin (finely powdered), 3^2 pound.
Caustic soda (powdered), 10 pounds.
Place the sulphur and resin, thoroughly mixed, in a barrel or smaller vessel,
APPENDIX I 675
and make a thick paste by the addition of about 3 quarts of water. Then stir in
the caustic soda. After several minutes, the mass will boil violently, turning a
reddish-brown, and should be stirred thoroughly. After boiling has ceased, add
about 2 gallons of water and pour off the liquid into another vessel, and add to it
sufficient water to make 6 gallons. This form of stock solution may be used at the
rate of i gallon to 50 of water for spraying mOst plants and for soaking seeds.
Eati Celeste (Modified). — It is made as follows:

Ammonia, 3 pints.
Sal goda, 5 pounds.
Dissolve the copper sulphate in 10 or 12 gallons of water, add the ammonia, and
dilute to 45 gallons; then add the sal soda and stir until dissolved. Eau celeste is
an dormant spray for the peach leaf-curl and other similar diseases, but it
effective is
unsafe to use on the foliage of most plants.

Polassinm Permanganate. (Not used in the United States.)
Potassium permanganate, i part.

Soap, 2 parts.
Water, 100 parts.
Recommended in France for black-rot and mildew of grape, etc.

Iron Sulphate and Sulphuric Acid.
Water (hot), 100 parts.

Iron sulphate, as much as will dissolve.
Sulphuric acid, i pint.
Prepare the solution Just before using. Add the acid to the crystals and then
pour on the water. Valuable for treatment of dormant grape vines affected with
anthracnose, applications being made with sponge or brush from wooden vessels in
which it is made. The solution will destroy the foliage, so it must be used in late
fall, or early spring, or applied only to tree trunks.
—
Within the last few years this wash has come into prominence as
Lime-sulphur.
one of the best Scale insecticides discovered. Several forms of it are excellent
fungicides. Three formulae are here given.
The Boiled Mixture (home-made).
Best stone lime, 15 pounds.

Flowers of sulphur, 15 pounds.
Water, 15 gallons.
Slake the lime in a small quantity of hot water, add the sulphur gradually and
stir thoroughly. Dilute the mixture to 15 gallons with water, and boil in an iron
kettle, or cook by steam in a barrel for forty-five minutes. Fill the vessel with water
to the required 50 gallons; strain the wash through a fine-mesh strainer and apply
hot. This wash should be applied in the fall after the leaves have dropped, or in the
spring before the buds open." Spray thoroughly, covering all parts of the tree.
Concentrated Mixture.
Sulphur, 80 pounds.
Best stone lime (95 per cent, calcium oxide), 40 pounds.
Water, 50 gallons.
Live steam run in a barrel, or fire under an iron kettle may be used in boiling.
Place 5 gallons of water and 40 pounds of the sulphur in the vessel, and apply heat
until the sulphur becomes a smooth paste, stirring constantly. Now add 10 gallons
of water and 20 pounds of lime and boil for forty-five minutes. Add water to make
25 gallons. . When cooled to 3S°F. test with Baume scale; the reading should be
about 33°F. As a scalecide to use in the dormant season, this should be diluted
I to 10 (i.e. I part of the above formula diluted with 9 parts of water) and 6 to 10
pounds of stone lime added to every 50 gallons of the spray. As a fungicide for
summer use, dilute i to 30 (i part of stock solution to 29 parts of water). When
away it is best
stored an eighth of an
to cover the solution with a layer of oil about
inch thick. This will prevent evaporation and the forming of a crust on the
material. The material should not be stored where the temperature will go very
low.
Self-boiled Lime Sulphur.
Lime, 8 pounds.
Sulphur, 8 pounds.
Water, 50 gallons.
This spray is valuable in cases where Bordeaux is injurious to foliage or fruit.

The stone fruits, such as plums, are particularly susceptible to Bordeaux injury,
while some varieties of apples are badly russeted by it.There is slight danger of
injury by the self-boiled lime-sulphur preparation, and is an efficient fungicide
it
when properly made. It stains the fruit as does Bordeaux. In making it 8 pounds
of lime of good quality should be placed in a barrel, and enough water to nearly
cover it should be added. While the lime is slaking, add sulphur which has run
through a sieve to break up the lumps. The sulphur should be stirred thoroughly
into the slaking lime, enough water being added to make a pasty mass. The barrel
should now be covered, in order to retain its heat, and the contents should be occa-
sionally stirred. The time required varies with the quality of the lime; if the lime
acts quickly, five to ten minutes would be sufficient, while if it acts slowly, fifteen
minutes may be necessary. It should not be allowed to stand too long, because it
may in that case be injurious to foliage. Now add water, stirring the mixture
while it is being poured in. Then add enough water to bring the total up to 50
gallons. In applying the spray, it is necessary to have a good agitator in the sprayer.
Consult RuGGLES, A. G., and Stakman, E. C. Orchard and Garden Spraying. Bull.
:
No. 121, Agric. Exper. Sta. Univ. Minn., March, 191 1. Also Duggar, B. M., and
CooLEY, J. S.: The Effect of Surface Films and Dusts on the Rate of Transpiration.
Ann. Mo. Bot. Gard., I: pp. 1-22, March, 1914.
APPENDIX I 677
Lime-sulphur Salt Wash. — This wash, although rarely used, is made as follows:
Lime, unslaked, 20 pounds.

Sulphur (flour, or flowers), 15 pounds.
Salt, 10 pounds.
Many different formulas are used in making up

this wash but the above formula
seems to be the best, and has been extensively used. If the lime is high-grade stone
lime, 15 pounds will be sufficient to dissolve all the sulphur. With average lime
20 pounds is the better quantity, but with poor or partly air-slaked lime 25 to 30
pounds are necessary. Lime absorbs an equal weight of water in becoming air-
slaked.
To prepare small quantities of this wash proceed as follows: Place about 10 gal-
lons of water in an iron kettle over a fire, make the sulphur into a paste with a little
water, and when the boiling point is nearly reached add the fresh lime and the sul-
phur together. The mixture should be constantly stirred, and the boiling continued
for forty to sixty minutes. The object of the cooking is to dissolve the sulphur and
when this is accomplished further boiling is useless, but not harmful. The salt may
be added at any time during the process of boiling, or entirely omitted. It is gener-
ally conceded, however, that salt increases the adhesiveness of the wash, as it does
ordinary lime whitewash, and for this reason, it is perhaps advisable to use it, al-
though it is not supposed to strengthen the fungicidal property of the mixture.

Possibly also the salt hastens the solution of the sulphur by raising the boiling point,
or by its solvent action.
It has been found that the sulphur dissolves more readily in a concentrated mix-
ture with lime, and the quantity of water used during the process of boiling should,
therefore, be reduced to aminimum. The mixture should not be allowed to become
pasty, however, and water, preferably hot, should generally be added until the barrel
is nearly full when finished. When the cooking is completed, pass the mixture
through an iron wire strainer (not brass or copper), and dilute with the required
amount of water. For details, see Waite, M. B.: Fungicides and Their Use in
Preventing Diseases of Fruits. U. S. Farmers' Bull. No. 243 (1906).
The wash may be applied either hot or cold with practically the same results,
though the warm mixture is less likely to clog the nozzles. If allowed to stand over
night, sulphur crystals will form on the bottom and sides of the containing vessel.
It is difficult to dissolve the lime-sulphur crystals after they have once formed. For
this reason, it is better not to prepare more than can be used the same day.
Steeps. — Solutions in use for dipping seeds, fruits and the like in order to control,
or check fungous diseases.
Formalin. — For oat smut
(.4) and stinking smut of wheat. Add }/-}, pound of
formalin to 30 gallons of water and immerse the seed grain for two hours, then spread
out and dry: or sprinkle the grain with the formalin solution until thoroughly wet,
shoveling over rapidly to distribute the moisture evenly, then place in a pile (covered
with sacking) for two hours and finally spread out to dry as in the first method.
{B) For potato scab. The formalin treatment of seed potatoes practically frees
the seed from scab with slight expense and trouble. Add ^^'2 pound of formalin to
15 gallons of water and immerse the seed tubers for two hours. The seed tubers
are then spread in thin layers to dry promptly. After removing from the solu-
tion, cut and plant as usual-.
Hot Water Method for Smuts (Jensen) (consult Freemen, E. M.: Minnesota
.Plant Diseases, p. 225). —
Provide two large vessels, preferably holding at least 20
gallons. Two wash kettles, soap kettles, wash boilers, tubs or even barrels, will do.
One of the vessels should contain warm water, say at 110° to i2o°F. and the other
scalding water, at 132° to i33°F. The first is for the purpose of warming the seed
preparatory to dipping it into the second. Unless this precaution is taken, it will
be keep the water in the second vessel at the proper temperature. A

difficult to
pail of cold water should be at hand, and it is also necessary to have a kettle filled
with boiling water from which to add from time to time to keep the temperature
right. Where kettles are used, a small fire should be kept under the kettle of scald-
ing water. The seed which is to be treated must be placed, half a bushel or more at
a time, in a closed vessel that will allow free entrance and exit of water on all sides.
Hence a gunny bag, or sac, can be used for this purpose. Now dip the basket, or
bag, of seeds into the water at 110° to i2o°F. and lifting it out plunge it into the
second vessel containing water at 132° to i33°F. After removing the grain from the
scalding water, spread it on a clean floor, or piece of canvas to dry.
Corrosive Sublimate.
Corrosive sublimate, 2 ounces.
Water, 15 gallons.
Dissolve the corrosive sublimate in 2 gallons of hot water, then dilute to 15
gallons, allowing the same to stand five or six hours, during which time thoroughly
agitate the solution several times. Place the seed potatoes in a sack and immerse
in the solution for one and a half hours, and then spread to dry.
Insecticides Used to Kill Insects
Carbon Bisulpkid.— This inflammable and volatile liquid is used against grain
weevils and against the insects that are destructive to herbarium specimens.
Crude Petroleum.— This is an oily inflammable liquid used against scale insects.
Hellebore.-—This is a stomach or internal insecticide. It is not poisonous to man
as are the arsenical insecticides, and is used where there is danger of poison remain-
ing on parts to be eaten. It is often used on currants and gooseberries when the
berries are beginning to ripen. It is used in the dry form, and must be fresh
when used.
Hydrocyanic Gas. — This gas is made by dropping potassium cyanide into sul-
phuric acid and water. The fumes are deadly to all kinds of animal life, and the gas
is used only in special cases.
Kerosene. —This is an excellent contact insecticide. Pure kerosene, however,
will ordinarily burn the leaves of plants, consequently it is used in pure form when
trees are dormant, or against insects off' of plants as grasshoppers, household insects,
etc.
Kerosene Emulsion. — This is probably the best form in which kerosene can be
used. A stock emulsion is made as follows:
APPENDIX I 679
Hard laundry soap (shaved fine), ^^ pound.

Water, i gallon.
Kerosene, 2 gallons.
Dissolve the soap in boiling water, remove from the stove, and immediately add
the kerosene; churn with a bucket pump until a soft, butter-like, clabbered mass is
obtained. One part of this stock is added to 10 to 12 of soft water. If the stock
solution is properly made this can be used on tender foliage of plants for such insects
as plant-lice, etc.
Lime Sulphur. — See ante.
Miscible oils mix with water. There are several oils on the
are those that will
market that are miscible These make a good winter spray for scales and
in water.
are also excellent summer sprays against the same insects. Great care, however,
must be taken to get the right dilution, or burning of the leaves will result.
Paris Green is used by many where an arsenical insecticide is necessary. It is
generally used at the rate of pound to 50 gallons of spray. In using, always first
i
make a paste of the Paris green and water, and then add to the spray material.
Pyrethrum, or Insect Powder (Persian insect powder, Dalmatian powder, or
—
Buhach). This is a powder from the ground-up flowers of the pyrethrum plant.
•It is a contact insecticide and is used against fleas, cockroaches, etc. If the powder
is burned in a room the fumes will destroy mosquitoes and flies.
—
Resin Lime Mixture. Used with a fungicide, or insecticide, to insure sticking
of poisonous material to smooth, glossy leaves.
Pulverized resin, 5 pounds.
Concentrated lye, i pound.
Fish, or other animal oil, i pint.
Water, 5 gallons.
Place the oil,and i gallon of water in an iron kettle and heat until the
the resin
resin softens; thenadd the lye and stir thoroughly. Add to this 4 gallons of hot
water, and boil until a little mixed with cold water gives a clear, amber-colored
liquid. Add water to make up to 5 gallons. This is a stock solution. In spraying
with Paris Green, or Bordeaux mixture, take 2 gallons of this mixture, dilute it to
10 gallons, and add 40 gallons of spray.
Soap. — Ordinary soap is a valuable contact insecticide.
Ivory soap, i pound.
Water, 14 gallons.
Boil the soap in 5 to 6 gallons of water until dissolved, dilute with water to 14
gallons and spray while still warm. It is recommended for plant-lice, red spiders, etc.
Sulphur. — Flowers of sulphur is often dusted on ornamental plants to prevent
such diseases, as powdery mildews, and spots, 2 parts of sulphur and i part of
air-slaked lime.
Tobacco is a very important contact insecticide. As a powder it is one of the best
remedies for root-lice on trees. As a decoction it may be used as a spray against

plant-lice. Tobacco smoke kills soft-bodied insects.
—
Whale Oil Soap (Fish-oil Soap). This is a commercial product, and is a good
contact insecticide, particularly for soft-bodied insects, like plant-lice.
.2
g
•a
a
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APPENDIX II 68l
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APPENDIX II 683
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ADDITIONAL EXERCISES
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i
APPENDIX II
Fig. 238. — Spray pumps isolated and with bucket attachments.
^
Spraying Apparatus.- — Various forms of spraying apparatus are upon the market
for use in the different operations of spraying. The student is directed to trade
catalogs and to special treatises on the subject for details.
We may, as an introduction to this subject, classify the types of spraying outfits
into: Bucket pumps (Fig. 238), knapsack sprayers (Fig. 238), barrel pumps (Fig.
239), the tank outfit, geared sprayers, steam and gasoline outfits, etc.
The question of details resolves itself into a consideration of hose, extension
rods, nozzles, forcepumps, wagons, push carts and receptacles for the spray materials
(for outfit see page 672). For these details and a list of firms dealing in spraying
apparatus, consult a bulletin by C. A. McCue entitled Plant Protection, Bull. No. 97,
Del. Col. Agric, Exper. Sta., June 15, 191 2.
APPENDIX III
Antisepsis and Disinfection. — An antiseptic is a substance which acts to the ex-

clusion from wounds of living organisms that cause putrefaction, or decay.
Liquor Antisepticus. —
155 grains of boric acid should be dissolved in 113-^ ounces
of water, and 7 grains of benzoic acid in 2 3'2 ounces of alcohol, and the two liquids
then mixed. After dissolving 7 grains of thymol in a mixture of 8 drops of oil of
peppermint 4 drops each of eucalyptol and oil of gaultheria and i drop of oil of thyme,
triturate with 155 grains of purified talc and add the solution of benzoic and boric
acids. Shake occasionally during forty-eight hours, filter and add to the clear fil-
trate first iî ounces of alcohol, and then sufficient water to bring the volume up
to I pint.
Formalin. —Has powerful antiseptic properties. It is sold in 40 per cent,
solution and can be distilled with water to the required strength.
Corrosive Sublimate (Bichlorid of mercury). — It is used in solution in water in
a strength of i : 1000.
Definition of Disinfectant. —A disinfectant is a substance used to destroy the
germs of infectious diseases. The common disinfectants are formaldehyd (liquid,
gaseous), carbolic and (phenol) cresol, chlorinated lime (chlorid of lime), corrosive
sublimate. See Dorset, M.: Some Common Disinfectants. U. S. Farmers' Bull.
No. 345 (1908).
Preservation of Wood by Impregnation. —
Impregnation tends to increase the dura-
bility of wood by injecting an antiseptic liquid and may mean a desirable, or un-
desirable, change of color, and in some cases fire-proofing. Little is known about
he latter. Four principles may be applied.
A. Immersion.
I. Immersion in a salt. Corrosive sublimate (kyanizing).
II. Metalized wood by dipping in a solution of iron sulphate.
B. Boiling.
I. In salt water or solution of borax.
II. Frank's mixture, 95 per cent, liquid manure and 5 per cent, of lime.
III. Injection of copperas (siderizing).
IV. With exhaust steam.
APPENDICES III, IV, V 693
C. Use of Hydrostatic Pressure. —

Boucherie method with sulphate of copper.
D. Use of Air Pressure (Open-tank treatment).
E. Use of Steam Pressure. —
The liquids commonly used are chloride of zinc,
coal-tar creosote, mixture of chloride of zinc and of creosote, gases of tar
oils (thermo-carbolization), heavy petroleums.
Preservation of Wood by Air Drying or Kiln Drying. Bibliography. — Schenck,

C. A.: Logging, Lumbering or Forest Utilization, 1913, and the following bulletins:
Bureau of Forestry and late Forest Service, U. S. Dept. Agr.: No. 41, Seasoning of
Timber; No. 50, Cross Tie Forms, Etc. with Reference to Treated Timbers; No. 51,
Condition of Treated Timbers Laid in Texas, February, 1902; No. 78, Wood Preser-
vation in the United States; No. 84, Preservative Treatment of Poles; No. 107,
Preservation of Mine Timbers; No. 118, Prolonging Life of Crossties; No. 126,
Preservative Treatment of Red Oak and Hard Maple Cross' Ties, etc.
APPENDIX IV
CULTURE OF MUSHROOMS
Tissue Culture of Fleshy Fjingi.— Consult Duggar, B. M.: The Principles of

Mushroom Growing and Mushroom Spawn Making. Bull. No. 85, Bureau of Plant
Industry, 1905: 18.

This method is applicable to the mushroom and to 68 other species of fleshy
fungi listed by Duggar.
A young sporophore of Agaricus campestris is taken and broken open longitudi-
nally. A number of pieces are carefully removed with a sterile scalpel to a sterile
Petri dishon a number of nutrient media such as bean pods, manure and leaf mould.
From and numerous other similar tests it was ascertained that when the mush-
this
rooms, from which the pieces of tissue are taken, are young and healthy, there is
seldom an instance in which growth does not result. It was easily shown that failure
to grow was generally due to advanced age of the mushroom used, to an unfavorable
medium, or to bacterial contamination.
APPENDIX V
SYNOPSIS OF THE FAMILIES AND PRINCIPAL GENERA OF THE MYXOGASTRALES
Suborder I. Exosporeae. — Spores developed outside the sporophore.

of
Family I. Ceratiomyocace^. — Sporophores membranous, branched; spores

white, borne singly on filiform stalks arising from the areolated
sporophore.
Suborder II. Endosporea;.— Spores developed inside the sporangium, sthalium or
plasmodiocarp.
A. Spores violet-brown, or purplish gray (ferruginous in Slcmonitis fcrruginea
and S.flavogenita, colorless in Echinostelium).

(a) Sporangium provided with lime (Calcium carbonate),
Family 2. Phvsarace^. —
Lime in the form of minute round granules, innate
sporangium wall.
in the
Capillitium charged with lime throughout. Badhamia.
Capillitium of hyaline threads with lime knots.
Sporangia single, subglobose, or plasmodiocarps; capillitium without free,
hooked branches. Physarum.
Sporangia forming an sethalium. Fidigo.
Plasmodiocarps; capillitium with free, hooked branches. Cienkowskia.
Sporangia goblet-shaped or ovoid; stalks cartilaginous. Craterium.
Sporangia ovoid, shining, clustered; stalks membranous. Leocarpus.
Capillitium without lime.
Sporangial wall opaque. {Chondriodcrnia ( = Diderma).
Sporangial wall hyaline. Diachaa.
Family 3. Didymiace^. —^Lime in superficial crystals deposited outside the
sporangial wall.
Crystals stellate, sporangia single. Didymium.
Crystals stellate, sporangia forming an sthalium. Spumaria ( = Mucilago).
Crystals lenticular. Lepidoderma.
(b) Sporangia without lime.
Family 4. —
Stemonitace^. Sporangia single, provided with a stalk and
columella.
*Sporangial wall evanescent.
Capillitium spreading from the column and forming a superficial net.
Siemonitis.
Capillitium as above, but not forming a superficial net. Comatrkha.
Capillitium spreading from the apex of the sporangium. Enerthenema.
**Sporangial wall more or less persistent. .
Capillitium radiating from.the columella. Lamprodcma.

Capillitium scanty, colorless, branching from a short columella, sporangia
very minute. Echinostclium.
Family 5. Brefeldiace^. — Sporangia combined into an aethalium.
Capillitium irregularly branched. Amaurochate.
Capillitium with chambered vesicles. Brcfddia.
B. Spores variously colored, not violet {except Cribraria violacea).
(a) Capillitium wanting, or not forming a system of uniform threads.
Family 6. CribeariacEvE. — Sporangial wall membranous, beset with micro-
scopic round plasmodic granules.
Sporangia asthalioid, the wall not forming a persistent net. Lindbladia.
Sporangial wall forming persistent net. Cribraria.
Sporangial wall forming numerous parallel ribs. Diclyditim.
Family 7. — Sporangial wall cartilaginous.
Liceace^.
Sporangia solitary, sessile.
Licea.
Family Tubiferace^.— Sporangial wall membranous,
8. without round plas-
modic granules.
Sporangia tubular compacted. {Tuber ifera ( = TuhuUna).
APPENDICES V, VI 695
Family 9. Reticulajiiace^.— Sporangia closely compacted and usually forming

an sethalium, true capillitium none.
Sporangia columnar, inner walls reduceri to straight slender threads.
Dictydalhalium.
Sporangia interwoven, inner wall reduced to broad bands. Enlcridium.
Sporangia interwoven, inner walls laciniated. Relicularia.
(b) Capillitium present; a system of uniform threads.
Family 10. Trichiace^. —
Sporangia single, rarely in an athalium. Peridium
without thickenings, without lime. Capillitium of tubular simple, or
branched, free threads. Spore mass as capillitium, yellow or red,
rarely white or brown, never violet.
*Capillitium of free elaters, or an elastic network of spiral thickenings.
Elaters free, spirals distinct. Trichia.
Elaters free, scanty, spirals obscure. Oligonema.
Elaters combined into a web or network. {Hemitrichia ( = Hemiarcyria).
**Capillitium a profuse network of threads (usually scanty and free in Peri-
chana populina), thickened with cogs, half rings, spines or warts.
Sporangia stalked, sporangial wall evanescent above. Arcyria.
Sporangia sessile, clustered, the walls single, persistent. Lachnoholus.
Sporangia sessile, the walls usually double. Perichana.
***Capillitium coiled and hairlike, or straight, and attached to the sporangial
wall.
Capillitium straight. Dianema.
Capillitium penicillate, spirally banded. Prototrichia.
****Sporangia forming an aethalium; capillitium consisting of branched color-
less tubes.
Capillitial tubes, thick- walled where they traverse the cortex, thin- walled
among the spores. Lycogala.
APPENDIX VI
KEY FOR THE DETERMINATION OF SPECIES OF MUCOR
Laboratory Work. — The teacher will find it good educational practice to supply
the class with material of the commoner moulds in order that they may become
familiar with the general morphology of the ZYGOMYCETALES.
From the standpoint of taxonomy the columella is an organ of the first im-
portance. The position of the columella in relation to the wall of the sporangium
has been described as "free,"- "subjacent," "infundibuliform."
Terms which have been applied in systematic works to the different shapes of
the columella^ are illustrated in Fig. 240, a to /, inclusive.
The whether sporangiospores, conidiospores, chlamydospores, oidiospores
spores,
or stylospores (as in Mortierella), have been described b)^ special names, as spheric,
ellipsoidal, oval, dumbbell-shaped, spindle-shaped, bottle-shaped, bead-shaped, etc.
'Lender, Dr. Alf.: Les Mucorinees de la Suisse, 1908: 29.

Several solid culture media recommended by Lindner can be used

in the growth
of various moulds and in Petri dishes for class use. Such is grape juice
in test-tubes
exactly neutralized and combined with 10 per cent, gelatin. Another medium is
prepared by taking i liter of white wine, heating it over a flame for one-half hour to
drive off completely the alcohol. The liquid lost by evaporation is replaced to bring
the volume up to i liter. It is neutralized exactly and 10 per cent, gelatin is added.
On this medium moulds grow luxuriantly. The gelatin can be replaced by agar-
agar, using 1.5 per cent., and the advantage of this medium is that it does not
liquefy. The writer has found baker's bread a useful medium for the growth of
moulds under bell jars, the air of which is kept moist by filter paper. If the bread
is used in Petri dishes, it can be sliced, cut into a circular form, soaked in water, or
beerwort, placed under cover in the Petri dish, which should then be sterilized one
or two times. He has found beerwort agar extremely useful in raising moulds and
other filamentous fungi. A supply of the -\- and — races of heterothallic moulds
Fig. 240. — Forms of columella, a. Spheric; b, spheric with collarette; c, oval; d,

depressed oval; e, piriform; /, panduriform; g, conic; h, cylindro-conic; i, mammiform;
k, I, spinescent. (After Lendner.)
should be kept in culture, so that the students may experiment with the formation
of the gametes and zygospores. These can be mounted in acetic acid with a ring
of asphalt about the cover-glass, or they can be fixed and carried up through the
alcohols to such materials as Venetian red in which they are not only beautifully
stained, but also keep indefinitely. The Venetian red can be softened in a water
bath and a little placed in the center of a slide with the addition of a little balsam
to fill out the space beneath the cover.
The systematic study of the moulds should begin after their general morphology
and physiology have been considered. Cultures, the names of which are known to
the teacher, should be then given to the members of the class in mycology, as un«
known moulds, which the members of the class should mount and determine. Such
mounts may be made in 2 per cent, acetic acid after treating first with a weak alcohol
(10 per cent.) to wet the mycelium, so that the acetic acid will cover the specimen
without air bubbles and without the hyphae massing together, as happens frequently
APPENDIX VI 697
when acetic acid is applied without the preceding application of the alcohol. The
identification of the "unknown" moulds can be made by the use of the following
key, which is a translation of the one given by Lindner in his work on the Swiss
moulds, and which includes most of the important moulds of the world. Pure
cultures of various moulds can be obtained from Johanna Westerdijk, Director of
the Phytopathological Laboratory, Amsterdam, Holland; from Krai's Bacteriologis-
chen Laboratorium, Prague, Bohemia, i., Kleiner Ring, 11; and from Mrs. Flora
W. Patterson, Bureau of Plant Industry, Washington, D. C. Some of them can be
obtained by exposing various articles to the air under a bell jar with filter paper.
Transfers of these moulds to fresh culture media should be made every two or three
months. During the summer and even during the winter the cultures can be kept
on ice in a refrigerator, so that the transfers need not be made so frequently during
the hot weather of the summer, or while the teacher is off on his vacation. The
janitor should be instructed to look after the ice supply during the year. Cf Povah,
.
A. H. W.: A Critical Study of certain Species ofMucor. Bull. Torr. Bot. Club,
44: 241-259, May, 1917, continued.
Key for the Determination of Species of Mucor

Sporangiophores not branched, i Group Mono-mucor.
Sporangiophores branched.
(a) Branches rare, or more numerous and indefinite, in racemes, or corymbs.

2 group Racemo-mucor.
(h) Branches definite in sympodia. 3 Group Cymo-mucor.
I Group Mono -Mucor
Sporangiophores unbranched. (E.xceptionally unless the conditions of nutrition

are unfavorable, they form branches. These are anomalous cases.)
1. Sporangiophores at first erect, afterwards weak, finally drooping and trans-

formed into a woolly felt of a rusty color, i M. rufesccns Fischer.
Sporangiophores always erect and forming a matted growth. (2)
2. Sporangiophores never exceeding 2 cm. (3)
Sporangiophores longer than 2 cm. (7)
3. Sporangiophores never e.xceeding 300 (4) /j..
Sporangiophores exceeding o cm. (maximum 2 cm.). (5)

5
4. On solid media matted growth very short, velvety, color at first brownish
red-carmine then grayish, sporangia small (20;u maximum). 2 M. Raman-
niamus MoUer.
Matted growth scarcely visible, sporangiophores 210^1, colorless, septate;
sporangia 40 to 45^l diameter. 3 M. siibtilissimus Oudemans.
5. Wall of sporangium not diffluent; on breaking it leaves an irregular, ragged
collarette, sporangia 36 to 42yu diameter, spores elliptic 6/x by 8^. Matted
growth 1.5 tall. 4 M. hygrophiliis Oudemans.
Wall of sporangium not diffluent, sporangia large, 80 to q8;u in diameter,
spores elliptic sû by Sju.
.
Matted growth 2 cm. high. 5 M. advcnlitius Oudemans.

Columella with orange-red contents; variety auranliaca Lendner.
6. Spores mixed with oil drops and intersporal granular protoplasm. 6 M.
plasmaticus van Tieghem.
Without drops of oil in the sporangium. (7) J
7. Sporangiophores cm. long.
2 to 3 (8)
Sporangiophores more than 3 cm. (9)
8. Sporangia 8om diameter, columella oval, spores 8^ by loî (except 8 by 14).
7 M. hiemaUs Wehmer.
Sporangia larger than 250 to 3 50M, columella pyriform, large, spores 4 to 8m
by 5 to 13m- 8 M. piriformis Plscher.
9. Wall of sporangium ruptured rapidly, columella frequently with yellow con-
tents, spores 3 to 6m by 6 to i2m. 9 M. mucedo Linn. (Fig. 13).
Wall of sporangium ruptured slowly, columella colorless, spores very large,
ISM by 30 to ^^n. 10 M. mucilagineus Brefeld.
2 Group Racemo-Mucor .
Branching indefinite, in racemes or in corymbs.
1. Branching secondary verticillate, these last have at their nodes the verticil-
late branches. 11 M. glomerula Lendner (Bainier).
Branching open in racemes, or in corymbs.
2. Columella hemispheric, covered with colorless threads resembling the capil-
litium of certain Myxomycetes. 12 M. comatus Bainier.

Columella round or oval, never presenting capillitial character, (3)
3. Sporangiophore at first erect, then curved toward the substratum, and then
fading. 13 M. de Baryanus Schostakowitsch.
Sporangiophores always erect and forming a matted growth. (4)
4. Species parasitic on other Mucorace^. 14 M. parasiticus Bainier.

Species not parasitic. (5)
5. Sporangiophores of two kinds, one with a terminal large sporangium with

diffluent wall, the others lateral, bearing sporangioles with persistent walls.
15 M. agglomeratus Schostakowitsch.
Species not possessing the above characters. (6)
6. Sporangiophores bearing laterally the branches with normal sporangia

(or abortive), or with zygospores.
Suspensors unequal. (7)
Sporangiophores normally laterally (i.e. all terminated by sporangia).
Zygospores with suspensors approximately equal. (8)
7. Sporangiophores straight, simple or branched bearing one or two opposite
branches terminated by sporangia. Columella depressed, spores elliptic 2 to
;in by 4 to 5m. 16 M. Moelkri Vuillemin (Fig. 241).
Sporangiophores straight, branched, bearing verticillately two to four
sporangia, columella roundish, spores spheric 2 to 2>t^ diameter, it M
heterogamus Vuillemin.
—
APPENDIX VI 699
Spores unequal (mixture of numerous small spores with others twice as

large). (9)
Spores approximately equal in size. (10)
Sporangiophores 0.5 to 1.5 cm., straight. Sporangia 80 to 125/1 diameter,
spores spheric or angular of diverse forms, 4 to 15/^ diameter. 18 M. heiero-
sporus Fischer.
Sporangiophores ordinarily 3 to 4 mm. (i cm. maximum), sporangia 70/x
diameter as maximum. Spores oval or subcylindric 2 to 6ju by 6 to 8/x.
Chlamydospores along the course of the sporangiferous hyphas. ig M.

sylvaticus Hagem.
Fig. 241. Mucor Moelleri. Stages in zygospore formation. {After Lendner.)
Sporangiophores i cm. Sporangia 40 to 54M, wall dehiscent. 20 M. lau-

sannensis Lendner.
10. Wall of sporangium not diffluent, but breaking into pieces. (11)
Wall diffluent. (13)
11. Spores spheric 7m diameter. 21 M. corymbosus Harz.
Spores oval. (12)
12 Sporangiophores frecjuently unbranched, chlamydospores provided with
very fine points; azygospore formation the normal process. 22 M. tenuis
Bainier.
Sporangiophores branched, chlamydospores with smooth walls, zygospores
and azygospores. 23 M. racemosus Fresenius (Fig. 30).
13. Spores spheric, 3 to 3. 5m- 24 M. piisillus Lindt,
Spores oval or elongated. (14)
—
14. Large species 6 to 8 cm. tall (exceeding in all cases 2 cm.). (15)
Small species never exceeding 2 cm. in height. (16)
15. Sporangiophores 6 to 7 cm. in height, sporangia 300 to 400/i (exceptionally
5oom), spores 7.5 by i7.5m- 25 M. proliferus Schostakowitsch.
Sporangiophores 6 to 8 cm. in height, sporangia 140 to 150^ diameter,
spores 4.2fi by 9 to 12^. 26 M. flavus Bainier.
16. Columella largely subjacent and concrescent with the wall of the sporangium,
diameter looyu, spores 2 to 4^. 27 M. mollis Bainier.
Columella free and slightly flattened at base. (17)
17. Spores oval, small 2.1^1 by 4.2/^, a grayish-blue. 28 M . Jragilis Bainier.
Spores elongated plano-convex, unequal, 2 to 5yu by 5 to ion. (18)
18. Sporangia never exceeding So/x, zygospores frequent, forming (on bread)
special branches. 29 M. genevensis Lendner.
Sporangia a mean of 80// frequently 120^1 diameter, suspensors bearing the
sporangiophores as with M. racemosus (Fig. 30). 30 M. erectus Bainier.
3 Group Cymo-Mucor
Sporangiophores branched in sympodial cymes.

1. Sporangiophores of two kinds, the one straight and bearing the normal
• spheric Sporangia, the other creeping, circinate branches sympodial, bearing
piriform sporangia. 31 M. pirelloidcs Lendner.
Sporangiophores of a single kind. (2)
2. Sporangiophores circinate. (3)
Sporangiophores straight not circinate. (6)
3. Sporangiophores never exceeding i cm., spores oval, maximum length 6/1- (4)
Sporangiophores exceeding i cm. sometimes 3 cm., spores spheric, loî or
more. (5)
4. Wall of sporangium brown, sporangium frequently subsessile, spores 3 to
4n by s to 6/x long. 32 M. circinelloides van Tieghem.
Sporangia wall bluish-black, sporangia carried on long pedicels, frequently
circinate, spores 4^1 by 5 to 6^. 33 M. griseo-cyanus Hagem.
5. Sporangiophores creeping, Yz to 2 cm., sporangia black 120 to 2ooju, spores
10.5JU to I4M in diameter. 34 M. angariensis Schostakowitsch.
Sporangiophores straight not circinate, the others short, freely branched and
circinate, sporangia small 6om (mean), 12/i (maximum). 41 M. laniprosporus
Lendner (Fig. 242).
6. Spores spheric or very unequal of diverse forms. 35 M. heterosponis sibiricits
Schostakowitsch.
Spores spheric appreciably equal. (7)
Spores oval. (12)

7. Species poorly cultivated on grape- juice gelatin, forming on bread a short
mat of 2 to 3 mm., sporangia 50 to 70/11, spores spheric, 5 to 6ju. 36 M.
Jansseni Lendner.
Species readily cultivated on grape-juice gelatin, forming a taller matted
surface (i to 3 cm.). (8)
—
APPENDIX VI 701
8. Columella spinescent. (9)

Columella smooth. (10)
9. Sporangiophores never exceeding 2 mm., sporangia 60 to 8o/u, spores smooth
7 37 M. spinescens Lendner.
to 8^.
Sporangiophores over i cm. and more tall, spores frequently punctate,
5 to 8m. 38 M. plumheus Bonorden.
Fig. 242. Mucor lam prosper us. a, b, c. Columella; d, sporangiole; e, sporangium;

/, branched sporangiophore. (After Lendner.)
Sporangia 75 to 120/x, columella piriform or campanulate, spores 4 to 8m

diameter. 39 M. globosus Fischer.
Sporangia ordinarily smaller {iion maximum), columella spheric, oval or
campanulate. Spores larger lo^ (mean). Species with sporangioles near
the substratum. (11)
Sporangia 70 to nop diameter, sporangioles not caducous, spores spheric,
shining, lo^. 40 M. spharosporus Hagem.
Sporangia never exceeding 80 to 90/1, spores lo^.
Sporangioles circinate, caducous, sporangiophores more elevated than in

preceding species. 41 M. lamprosporus Lendner (Fig. 242).
Sporangia 60 to 8o/i, spores normally 8 to 10, spheric or accompanied by
abnormal spores, oval 8 to iom by 30/i long, without sporangioles. 42 M.
dimorphosporus Lendner.
12. Large species 9 to 12 cm. high. (13)
Small species. (14)
13. Sporangiophores 9 to 10 cm., sporangia up to i mm. diameter, spores 10.5
by 28;u. 43 M. irkutensis Schostakowitsch.
Sporangiophores 10 to 12 cm., sporangia 500M, spores 5;u by 8.6. 44 M.
Wasnessenskii Schostakowitsch.
14. Wall of sporangia not diffluent, breaking into pieces. 45 M. brevipes Riess.
Wall of first sporangia diffluent. (15)
15. Spores elongate with punctate spore walls, sporangia blackish, 100^1 diameter.
46 M. amhiguus Vuillemin.
Spores subspheric with smooth walls. (16)
16. Species forming on bread or grape-j-uice gelatin a mycelium somewhat
raised and of a yellow color. 47 M. Rouxianus Wehmer.
Species forming a matted growth of i to 3 cm. tall. (17)
17. Species branched but little. (18)
Species copiously branched. (19)
18. Sporangia 50 to 350M, columella spheric, spores spheric or elliptic or angular,
4.2 by 6.SM with chlamydospores. 48 M. geophilus Oudemans.
Sporangia 90^1 to 170M diameter, columella ovoid, spores subspheric 5 to 6m
by 6 to 8m rarely iom. 49 M. strictus Hagem.
19. Sporangia 35 to 70M (90M diameter), spores 6m by 8m or 8 to iom diameter,
yellow pigment in hyphae weakly developed. 50 M. Prainii Chodat &
Nechitch.
Sporangia 50M, wall more diffluent, spores more frequently oval and very
small, 4 to 5m by 5 to 7m, also 4 to 7m diameter. 51 javanicus^ Wehmer. M .
APPENDIX VII
Keys for the Determination of Species of Aspergillus and Penicillium

For student use in systematic study, or identification of the green moulds be-
longing to the genus Aspergillus, the teacher will find the following key, adopted
from "Household Bacteriology" by the Buchanans, pages 76 and 77, of great value.
Lafar in his "Technical Mycology," Vol.II, Part 2, also gives on page 308 a useful
specific summary. The different species may be kept in culture for distribution
as unknown to the members of the class.
key to common species of ASPERGILLUS

I. White spores, or nearly white.
A. Sterigmata unbranched. Aspergillus catididus.
1 M. dubius is a variety of M. javanicus.
—
APPENDIX VII 703
B. Sterigmata branched. Aspergillus albus.

II. Colored spores.
A. Spores yellowisli-green, bluish-green, grayish-green, green.
1. Sterigmata unbranched.
(a)Perithecia produced readily.
1. Perithecia not imbedded, naked. A. herbariorum.
2. Imbedded perithecia.
With slightly swollen conidiophore tips, sterigmata club-shaped, later-
ally placed. A. clavaius.
With hemispheric conidiophore tips, sterigmata terminal. A.fumigatus.
(b) Perithecia unknown.
1. With large conidiophore tip, elongate 80 to ioom by 500 to Soo/x. A.
giganteus.
2. With smaller conidiophore, end spheric, or hemispheric.
With rough worty conidiophore. A flavus. .
With smoother conidiophore. A. oryzea.

2. Sterigmata branched.
(a) With rusty-brown myceUum. A. versicolor.
(b) Mycelium not rusty-brown.
End of conidiophore, club-shaped with lateral and terminal sterigmata.
A. pseudoclavatus.
End of conidiophore hemispheric with terminal sterigmata. A . nidulans.
B. With black, or dark-brown conidiospores.
1. Sterigmata unbranched. A. calyptratiis.
2. Sterigmata branched. A. niger.

C. With reddish-brown, yellowish-brown, or yellow conidiospores.
Sterigmata unbranched, spores coffee-brown. A. Wenlii.
Sterigmata branched, spores yellow-brown. A. ochraceus.
The genus Penicillium is closely related to the genus Citroniyces, which includes
fungi causing citric acid fermentation in sugar media and which has a single whorl
of conidia-bearing cells (sterigmata) at the tip of the conidiophore. All of the
fungi with the penicillate type of fructification are grouped together in the form
genus Penicilliiiin. The small and delicate conidiophore differs from that of Asper-
gillus in being divided into a row of short cells by transverse septae. The conidio-
phores are branched and the upright branches bear the sterigmata as tufts of termin-
ally disposedsecondary branches. The conidiospores are pinched off from the ste-
rigma and are arranged in chains. The whole inflorescence suggests a whisk, or a
broom. The spores are of various shapes and sizes from spheric to ellipsoidal.
Some have smooth walls, others are roughened. Several species show the tendency
to form coremia (coremium), which are tufted forms of inflorescence. Four, or
five, species are known to produce perithecia and ascospores, so that no satisfactory
key can be based on perithecial and ascosporic characters. The number of species
which are associated with the ripening of cheeses, or which produce decay in fruits
of various kinds is about six or seven. The species usually designated as Penicillium
glancnm and P. crustacenm are included in the most recent paper by Thom under
—
PeniciUinm cxpansum (Fig. 243) which can always be obtained from apples decaying
in storage. Colonies of this mould upon gelatin and potato, or bean agar, are green,
becoming gray-green and later brown. The conidiophores are tufted into corem-
ium-like clusters.
The conidia fructifications consist of one to three main branches bearing verticils
of branchlets supporting crowded whorls of sterigmata. Conidiospores are elliptic
2 by 3.3A1, green, persisting in chains, when mounted.
'*oi;f/'/II
MM ;,/
Fig. 243. Penicillium expansum. a, b, f. Arrangement of branches of conidial
fructification; c, d, e, conidiiferous cells and chains of conidiospores; g, h, j, k, I.

sketches of fructification; m, n, o, germination of conidiospores; r, s, sketches show-
ing in ^ loose aggregations of conidiophores, r coremmm. (After Thorn.)
Penicillium Roqiieforli (Fig. 244) is the agent in the ripening of Roquefort,

Gorgonzola and Stilton cheeses. Colonies on potato agar quickly become green,
becoming a dirty brown when old. The velvety mycehum consists of radiating
branching hyphae giving an indefinite margin. The conidiophores arise separately
and in acropetal succession from the growing parts of submerged hyphas, 200 to 300M
—
APPENDIX VII 705
long and septate. The conidiospores are bluish-green, globose-cylindric, 4 to 5m in

diameter. Roquefort cheese is a hard rennet cheese made from the milk of sheep.
Some imitations are made from cow's milk. The most striking characteristic of
this cheese is the mottled, or marbled appearance due to the develop-
of the interior
ment which is the principal ripening agent. The manufacture of
of this fungus,
Roquefort cheese has been carried on for at least two centuries in the southeastern
part of France, in the Department of Aveyron and the village of Roquefort. The
curd is put into hoops, which are filled in three layers, a layer of bread crumbs
penetrated with the hyphje of Penicillium Roquejorli being placed between the first
\m %
Fig. 244. Penicillium Roqueforti. a, part of a conidiop.hore; h, c, other types

of branching; d, young conidiophore, just branching; e, /, conidiiferous cells; g, /j,i,
diagrams of types of fructifications; k, I, m, n, germinating spores. {After Thorn.)
and second and the second and third layers.The bread is prepared from wheat and
barley flour, with the addition of whey and a trace of vinegar. It is baked and
kept moist from a month to six weeks during which time it is penetrated by the
green mould above mentioned. For use the bread is crumbled and sifted. The
cheese is subjected to pressure, which is gradually increased for ten to twelve hours.
It is turned usually one hour after putting into hoops. It is wrapped in cloth at
the end of twelve hours and taken to the first curing room. The cloths are fre-
quently changed during ten to twelve days. Formerly, the manufacture was
carried on by shepherds but now as the industry is commercialized, the ripening is
carried on in caves in the Roquefort region in which the air circulates freely and the
45
—
7o6 ADDITIONAL EXERCISES
temperature is 40° to 45°C. When ripe, the cheeses are prepared for shipment by a
covering of tin-foil properly inscribed with the manufacturer's name.
Penicillium Camemherli (Fig. 245). —The colonies of this important fungus on
potato agar are at first effused and white changing in five to eight days togray-
FiG. 245. Penicillium Camemherli. a, Conidiophore with common type of

branching with conidiospores; b, a common less-branched form; c, d, /, diagrams of
large fructifications; g, i, j, germinating conidiospores. {From Bull. 82, Bureau of
Animal Industry, also After Thom.)
green. The hyphae are loosely felted, about 5m in diameter. The septate conid-
iophores are 300 to 8oom in length and 3 to 4yu in diameter, thin-walled often
collapsing with age. Fructification about 175^ tall, consisting of one main branch
and one lateral branch, sparingly branched to produce the sterigmata which abstrict
off ellipsoidal conidiospores, smooth and bluish-green by transmitted light, thin-
—
APPENDIX VII 707
walled and commonly guttulate, 4.5 to S-Sm in diameter. The growing and fruiting
period about two weeks. This green mould grows in Camembert and other soft
is
cheeses, where it causes a breaking down of the casein. Camembert cheese is a soft
rennet cheese made from cow's milk. A typic cheese is about four and a half inches
in diameter and one and a quarter inches thick, and is sold in this country wrapped
in paper and inclosed in a wooden box of the same shape. The cheese has a rind of
Fig. 246. Penicillium stoloniferum. a, b, c, e, f, the types of branching at the

tips of the "stolons" by which the species spread in substrata; d, conidial fructifica-
tion; h, j, k, I, sketches of conidial fructifications of various ages; g, formation of
conidial spores; i, ripe conidiospores; m, n, germination of conidiospores; o, rough
diagram of habit. (After Thorn.)
considerable thickness, which consists of moulds and dried cheese surrounding a

yellowish, waxy, creamy, or almost fluid interior depending upon the ripeness of
the cheese. Probably originated about 1791 in the Department of Orne in north-
western France, the industry has extended into other departments of the French
Republic. It is made from whole fresh milk, or from milk which has been skimmed
in part. The curd which forms at about 8o°to 85° is transferred to perforated tin
forms, or hoops. These rest upon rush mats, which permit free drainage. After
—
7o8 ADDITIONAL EXERCISES
draining, the cheese is frequently turned and in two or three days, it is carried to a
well-ventilatedroom where the ripening process begins. Here it remains fifteen
to twenty days when the surface becomes covered with Penicillitim Camemberli,
which gradually breaks down the casein.
Pig. 247. Penicillium iialicum. a, b, c, d, e, f, g, types of branching in verticils

and chains of conidiospores; j, k, sketches of conidial I, m, n, swelling
fructifications;
and germination of conidiospores. (After Thorn.)
Penicillium stoloniferum (Fig. 246) grows on decaying fungi, Boleti, Polypori and
in cultures from milk and ensilage. It has been collected repeatedly at Storrs,
Conn., and once upon decaying Boletus scaber at the Jardin des Plantes in Paris, and
—
APPENDIX VII 709
hence, it is probably widely distributed. Its stolon-producing character is very

characteristic and diagnostic.
PenicilUum italicum (Fig. 247) and P. olhaceum occur on tropic fruits, including
pineapples, lemons, oranges, etc. The fungus causes extensive putrefaction in such
fleshy fruits as the pineapple.
PenicilUum brevicaidc (Fig. 248) grows on decayed paper and it has been recom-
mended by Gosio when grown in media with traces
for the detection of arsenic, since
of arsenic, it forms the pungent compound diethylarsine. None of the species of
PenicilUum are pathogenic. About six to seven species of this genus are connected
with the ripening of cheeses. For example, a little-known Norwegian cheese
"Gammelost" has its ripening, according to Johann Olsen, a green
associated with
mould, PenicilUum aromaticum, and so showing the unsatisfactory state of our
Fig. 248. PenicilUum brevicaule. a, Conidiophores and simple chains of conidi-

ospores; b, f, more complex conidial fructifications; c, two young chains of conidio-
spores; d, e, echinulate conidiospores; g, h, j, sketches of forms and habits of conidial
fructifications; k, germinated conidiospores. (After Thoyn.)
knowledge about these fungi, this fungus may prove on close investigation to be
identical with the one which works in Roquefort cheese.
As all of the species of PenicilUum are readily cultivated and kept for some time in
a satisfactory condition for study, they are especially useful in the systematic exercises
which are essential in the training of competent mycologists. As the time which can
be devoted to such a study is limited, the work can be varied by assigning, as un-
knowns, cultures of the different species of the genus Aspergillus to certain members
of the class and cultures of PenicilUum as "unknowns" to other members, and it may
be advisable to interchange the material, so that all of the students in the class in
mycology become acquainted with the similarities, as well as the differences dis-
played by fungi of the genera Aspergillus and PenicilUum. It is better to distribute
these moulds to the class in culture media in Petri dishes than in test-tubes, because
7IO ADDITIONAL EXERCISES
is more easily accomplished, and because the

the removal of the material for study
whole growth can be examined readily by placing the Petri dish on the stage of the
microscope and examining with the low power. In mounting such fungi for study
beneath a cover-glass lo per cent, alcohol should be used to wet the spores and
hyphae, otherwise difficulty will be encountered with spores flowing together in mass
and the hyphse becoming knotted together. Thom, in his paper on the " Cultural
Studies of Species of Penicillium," published as Bull. ii8 of the U. S. Bureau of
Animal Industry in 19 lo, recommends that the following media be prepared for the
study of the species as his key for the identification of the species given below is
based on their behavior upon the recommended culture media. For this purpose
prepare the following media: (i) 15 per cent, gelatin ("gold label") in distilled water;
(2) 15 per cent, gelatin in distilled water plus 3 per cent, cane sugar; (3) either bean
or potato decoction plus 1.5 per cent, cane sugar; (4) bean or potato agar plus 3
per cent, cane sugar. Litmus solution may be added, if desired, when cultures are
Fig. -Penicillium claviforme. a, Coremium grown upon sugar media;

coremium on gelatin free from sugar. {After Thom.)
made. Prepare Petri dishes with 10 c.c. of each of the media used and allow them to
cool. Inoculate two or more Petri dishes of each medium with spores of the species
to be distributed to the class. Incubate at 2o°C. (the laboratory temperature is
usually satisfactory). Have the members examine at intervals of three

of the class
days, or less, making naked-eye observations from above and below also with a
hand lens and with the low power of the compound microscope. A drop of litmus
solution at the margin of a colony can be used to test acidity, or alkalinity.
Have the class examine i and 2 for liquefaction; 2 and 4 for coremium amd sclero-
tium formation which will call for continued examination for at least two weeks.
Below will be found two separate keys. One, after Thom, is a general key
of species of Penicillium grown upon the above-recommended agar and gelatin
media. The second key, after Buchanan, which includes the species of most eco-
nomic importance, is based on the character of the substratum on which the
fungi are found growing in a state of nature.
—
APPENDIX VII 711
Fig. 250. Penicillium Duclauxii. a, b, Conidial fructifications with young

smooth conidiospores; c, d, e, conidial fructifications from potato-agar plate culture,
more complex types-; /, g, h, j, sketches of habit upon potato agar; k, ripe spores
highly magnified to show delicate markings; I, m, n, germination of spores; si,
coremium. (After Thorn.)
Fig. 2 si. ^-Penicillium chrysogenum: a. b, c, d, e, branching of conidial fructifica-

tion from gelatin plates; /, g. h, j, I, m, sketches of conidial fructifications from
potato-agar plates; n, o, germination of conidiospores. {After Thorn.)
—
I. Key of Species Grown on Agar and Gelatin Media
A. Species fruiting typically by coremia (vertical and definite).

a. Coremia long (3 to 15 mm.).
1. Conidial masses strictly terminal, olive-green, fragrant. P. daviforme
(Fig. 249).
2. Upper third of coremia fertile, conidia green. P. Duclauxii (Fig. 250).
aa. Coremia small.
Fig. 252. Penicillium roseum. a, b, c. Branching of conidial fructification,

showing few cells of each verticil; d, e, conidiiferous cell and conidiospores; g, h, j, k,
sketches of ripe fructification showing agglutination of conidiospores into slimy
masses. {After Thorn.)
1. Coremia definite, densely crowded, colony orange below. P. granu-

latmn.
2. Coremiform character indicated in cultures by clustering of conidio-
phores, definite coremia only in old cultures, becoming large and definite
upon apples. P. expansum (Fig. 243).
AA. Species not (or rarely) producing coremia in culture.
B. Species constantly producing sclerotia, or ascigerous masses.
b. Producing ascigerous masses, yellow, or reddish. P. hUeum.
bb. Sclerotia appearing as white masses in old cultures. P. ilaliciim (Fig.
l-
247).
bbb. Sclerotia reddish or pink, globose or elliptic, 500^ or less in diameter.
—
APPENDIX VII 713
Fig. 253. Penicillium atranienlosmn. a, b, c, d, branching of conidial fructifica-

tions showing unequal length of branching; e, /, conidiiferous cell and chain of co-
nidiospores; g, h, j, sketches of conidial fructifications; i, conidiospores; m, n, o, r,
germination of spores. {After Thorn.)
Fig. 254.—Penicillium lilacinum. a, h, c, Short conidiophores and verticils of

conidiiferous cells; d, conidiiferous cell, solitary and sessile; e, conidia;/, g, h, sketches
of conidial fructifications. {After Thorn.)
—
BB. Sclerotia not (or rarely) produced (under special conditions), Use gelatin
cultures (i) andcompare agar cultures.
(2),
C. Rapid liquefiers (abundant liquid in five to twelve days).

D. With definite, strong ammoniacal odor.
1. Yellowish brown, spores rough. P. brevicaide (Fig. 248).
2. White or cream, spores rough. P. brevicaide var. album.
Fig. 255. Penicillium funiculosutn. a, b, c, d, e, f, conidial fructifications with

conidiiferous cells and conidiospores; g, h, k, I, m, n, fructifications separate and
borne upon hyphae and ropes of hypha3; o, r, germination of conidiospores. (After
Thorn.)
3. White or cream, spores smooth. P. brevicaide var. glabrum.

DD. Without ammoniacal odor.
E. With yellow coloration of liquefied gelatin (not of mycelium in reverse).
1. Colonies small, conidiophores 100 to 150^ in length. P. citrinum.

2. Colonies broadly spreading, conidiophores 250 to 300^. P- chrysogcnum
(Fig. 251).
—
APPENDIX VII 715
EE. Without yellow color in liquefied gelatin (or slight traces only).
e. Colonies white to pink or salmon. P. roseiim (Fig. 252).
ee. Colonies some shade of green.
/. Colonies floccose, margin spreading by stolons. P. stoloniferum (Fig. 246).
//. Colonies velvety; surface growth of fruiting hyphse only; conidiophores
200 to 400^1 long, with a verticil of branches; reverse and medium
darkened in sugar media. P. alramcntosum (Fig. 253).
CC. Liquefaction of gelatin none or slower than ten to twelve days, or only partial.
G. Colonies never green.
"=^"=^--.ii
Fig. 256. Penicillilun decumbens. a, h, c, d, Conidial fructification with a
single verticil of conidiiferous cells; h, j, k, sketches of conidial fructifications. (Aftet
Thojn.)
g. Colonies yellowish-brown, spores elliptic. P. dlvaricatum.

gg. Colonies white to lilac, slow liquefier, fourteen to sixteen days. P. lilacinum
(Fig. 254).
ggg. Colonies floccose white or creamy; conidiophores long, typically penicillate.

P. Camemberti var. Rogcri.
GG. Colonies some shade of green.
H. Surface with hyphae definitely in ropes or trailing, bearing numerous conidio-
phores, as short branches, distinctly traceable to their origin in such
hyphcc.
h. Colonies usually red below and reddening the substratum.
—
7i6 ADDITIONAL EXERCISES
1. Fruiting areas dark green. P. funiculostim (Fig. 255).

2. Fruiting areas mixed yellow and green. P. pinophilum.
hh. Colonies not producing red color.
Fig. 257. Penicillium biforme. a, b, g. Branching of conidial fructification;
c, conidiiferous cells and conidiospores; h, j, k, sketches of conidial fructifica-

d, e, f,
tionson potato agar; I, m, sketches of conidial fructifications on sugar gelatin; o, r>
germination of conidiospores. {After Thorn.)
1. Colonies gray, rarely greenish, very loose floccose. P. intricalum.

2. Colonies gray to green, hyphie scattered, creeping. P. decumbens (Fig.
256).
HH. Surface hyphae not in well-defined ropes, nor trailing.
—
APPENDIX VII 717
i. Surface hyphae woven floccose, course of hyphae not traceable.

1. Gray-green, long conidiophores, no odor. P. Camemberti (Fig. 245).
2. Gray-green, shorter conidiophores, strong odor. P. biforme (Fig. 257).
Fig. 258. PenicilUum commune, a, b, c, d, e, Conidial fructification with conidio-

spores;/, g, h,j, k, I, sketches of fructifications in various stages. (After Thorn.)
ii. Surface growth at margin simple conidiophores, in older parts both floccose
hyphae and conidiophores.
I. Gray-greenish, branching of conidiophore rather loose, odor none or
slight. P. No. 22.
—
7i8 ADDITIONAL EXERCISES
Fig. 259. Penicillium spinulosum. a, b, Conidial fructifications, consisting of

single Verticils of conidiiferous cells; c, conidiiferous cell with chain of conidiospores
(smooth); d, f, ripe echinulate conidiospores; c, swollen end of conidiophore; g, /;,
sketches of conidial fructifications. (After Thorn.)
Fig. 260. Penicillium rubrum. a, b, c, d, e. Whole conidiophores and the

branching of conidial fructifications;/, g, conidiiferous cells and conidiospore forma-
tion; h, j, sketch of habit of growth; m, diagrammatic figure of a series of conidial
fructifications. (After Thorn.)
—
APPENDIX VII 719
2. Green, conidial fructiikations rather compact, odor definite, "mouldy."

P. commune (Fig. 258).
Hi. Fruiting surface velvety of simple conidiophores, or conidiophores borne
so close to surface of subtratum as to appear simple.
j. Conidial mass a dense column of conidial chains.
1. Column from a single verticil of sterigmata. P. spimilos'itm (Fig. 259).
2. Column from a verticil of branchlets with verticillate cells and chains.
P. nibrum (Fig. 260).
jj. Elements of conidial fructifications not in a column.
k. Conidiospores smooth.
I. Green, broadly spreading, ripe conidia globose, 4 to 5^1. P. Roqiicfoiti

(Fig. 244).
I
CI
Fig. 261. Penicillium purpurogenum. a, h, c, Conidial fructifications; d, e,f, g,
conidiiferous cells and conidiospores; h, j, k, I, m, sketches of whole fructifications.
{After Thorn.)
2. Green, less spreading, conidiospores elliptic, uredium commonly purpled.

P. purpHrogcnum (Fig. 261).
3. Gray or olive-green, conidiospores 5 to 7 by 3 to 5/u. P. digitalum
(Fig. 262).
kk. Conidiospores delicately rugulose. P. rugulosum (Fig. 263).
2. Key of Species Determinable from Substrata. (After Buchanan.)
Cheese (Camembert and Brie).
1. Floccose, white unchangeable, no odor. P. Camemberti var. Rogeri.
2. Floccose, white to gray-green, no odor. P. Camemberti (Fig. 245).
3. Powdery, yellowish-white, spores smooth, ammoniacal odor. P.
brcvicaule var. glabnim.
— .
4. Powdery, yellowish- white, spores tuberculate, ammoniacal odor. P.

brevicaule var. album.
5. Forming yellowish-brown areas, spores rough, ammoniacal odor. P.
brevicaule (Fig. 248).
Cheese (Roquefort).
I. Green streaks inside the cheese. P. Roquejorli (Fig. 244).
Fig. 262. Penicillium digilalum. a, Whole conidiophore and_ fructification; b,

c, d, e, types of branching and formation of conidiospores; m, n, o, germination of
conidiospores. {After Thorn.)
Citrus fruits.
1. Colonies of mould, blue-green. P. italicum (Fig. 247).
2. Colonies of mould, olive-green. P. digitatum-olivaceum
Pomaceous fruits (apples, pears, etc.).
I. Blue-green colonies finally producing P. expansum (Fig. 243).
Polyporaceae (Boleti, Polypori, etc.).
I. Colonies green (yellowish-green), spreading by stolons. P. stolonijerum

(Fig. 246).
—
APPENDIX VIII 721
Wood (pine).
I. Producing orange to red stains in pine wood. P. pinophilum.
Fig. 263. Penicillium rugulosutn. a, b. Branching of conidiophore; c, d, e,

conidiiferous cells and conidiospores; /fully ripe conidiospore; g, h, j, swelling and
germination of conidiospore;./, m, diagram of conidial fructifications. (After Thorn.)
APPENDIX VIII
Keys to the Genera of the ERYSiPHACEiE

(See Salmon, Ernest S.: A Monograph of the Erysiphaceae Mem. Ton. Bot.
Club IX, 1900.)
A. Perithecium inclosing only a single ascus.
(a) Appendage simple, filamentous, unbranched. r Spharolheca.
(b) Appendage dichotomously branched at end. 2 Podosphccra.
B. Perithecium containing many asci.
{a) Spores unicellular.
I. Perithecia with appendages.
* Appendages often basally swollen, never enlarged into a plate,
t Appendage unrolled at the end, or only slightly and irregularly curled.
*J Appendages simple, or only irregularly branched.
§ Appendages mycelium-like, unbranched, or slightly irregularly

branched. 3 Erysiphe.
§§ Appendages stiff, bristly, radially arranged, numerous. 4 Pleoch-
ceta.
tX Appendages frequently dichotomously branched at apex. 5 Micro-

sphara.
46.
tt Appendages more or less spirally coiled at the apex. 6 Uncimila.

** Appendages united at the base into a plate. 7 PhyllacHnia.
2. Perithecia without appendages, sessile or mycelium. 8 Erysibella.
(b) Spores divided. 9 Saccardia.
Key to the Species of Sph^rotheca (After Salmon)
Brief Characterization. —Perithecia subglobose, ascus solitary, eight-spored.

Appendages floccose, brown or colorless, spreading horizontally and often interwoven
with the mycelium, simple or vaguely branched, frequently obsolete.
1. Mycelium persistent, thick, pannose, forming dense patches of special hyphae
in which the perithecia are more or less immersed. (2)
Mycelium without these characters. (4)
2. Persistent mycelium usually satiny and shining, white, sometimes becoming
gray, or pale brown. 2 pannosa.
Persistent mycelium dark brown. (3)
3. Inner wall of perithecium separating from the outer, hyphse of persistent

mycelium very tortuous. 4 lanestris.
Inner wall not separating, hyphas straighter. 3 mors-iivm.
4. Perithecia 60 to ySju in diameter, ascus 60 to 75 by 42 to 50^1, inner wall of
perithecium separating from the outer. 5 phytoptophila.
Perithecia 50 to 120M in diameter, ascus 45 to 90 by 50 to 72/*; inner wall
scarcely separating. (5)
5. Cells of outer wall of perithecium 10 to 20M wide, averaging 15/x. i hutmili.
Cells 20 to 30 (rarely 40) /x wide, averaging 25/^. i humuli var. fuUgnea.
Key to Species of Podosph^ra (After Salmon)
Brief Characterization. — Perithecia globose, or globose-depressed; ascus solitary,

subglobose; spores eight. Appendages equatorial or apical, branches simple and
straight, or swollen and knob-shaped; very rarely of two kinds: one set apical,
brown, rigid, unbranched or rarely one to two times dichotomous at the apex; the
other set basal, short, flexuous, simple, or vaguely branched, frequently obsolete.
1. Basal appendages present, apical appendages usually unbranched. 4

leiicotricha.
Basal appendages absent. (2)

2. Appendages erecto-fasciculate, springing from near the apex of the peri-
thecium. (3)
Appendages more or less spreading and equatorially inserted. (4)
3. Appendages six to twelve and one-half times the diameter of the perithecium,
colorless, or occasionally pale brown toward the base. 2. Schlectendalii.
Appendages one to eight times the diameter of the perithecium, dark brown
for more than half their length, i oxyacantha var. tridactyla.
4. Appendages colorless, or faintly tinged with brown at the base, branches of

apex not swollen. 3 biuncinata.
APPENDIX VIII •
723
Appendages dark brown for more than half their length, ultimate branches
of the apex knob-shaped, i oxyacantluc.
Key to Species of Erysiphe (After Salmon)
Brief Characterization. — Perithecia globose, or globose-depressed, sometimes be-

coming concave; asci several, two- to eight-spored. Appendages floccose, simple
or irregularly branched (never with a definite apical branching) sometimes obsolete,
usually more or less similar to the mycelium and interwoven with it, very rarely
{E. tortilis) brown, assurgent and fasciculate.
1. Asci (of mature perithecia) not containing spores on living host plant. (2)
Asci (of mature perithecia) containing spores.
2. Perithecia large, 135 to 280^ in diameter, averaging 200/x, more or less im-
mersed in the lanuginose persistent mycelium.
4 graminis.
Perithecia smaller, 80 to 140M, not immersed in the lanuginose mycelium. (3)
3. Haustoria lobed. 3 galea psidis.

Haustoria not lobed. 2 cichoracearum.
4. Asci two-spored, rarely (and never uniformly) three-spored. (5)
Asci three- to eight-spored, rarely (and never uniformly) two-spored. . (8)
5. Perithecia 52 to 6oju in diameter; asci three, 48 to 50 by 28 to 36/i. 8 trina.

Perithecia 80 to 240^ in diameter; asci more than three, larger. (6)
6. Perithecia large, becoming pezizoid, 135 to 240^1 in diameter, usually about
2oo/u; asci seven to thirty-eight, usually about twenty, 75 to iiom long,
averaging gofi, spores 28 to 40/i»long, averaging 32 by i8î long. 6 taurica.
Perithecia 80 to 140/i (very rarely 100 to 175); asci four to twenty-five (very
rarely as many as thirty-six), usually ten to fifteen, 58 to goti long; spores
20 to 28/i long, averaging 34 by i4ju. (7)
7. Haustoria lobed. 3 galeopsidis.

Haustoria not lobed. 2 cichoracearum.
8. Perithecia 65 to i8om in diameter, usually about 90^1; asci usually few, two
to eight, rarely as many as twenty-two, 46 to 72 (rarely 80) ju long. (9)
Perithecia larger, 130 to 280M in diameter, averaging 180 to 200/*; asci, nine
to forty-two, 70 to 11 5m long.
9. Appendages very long, ten to twenty times the diameter of the perithecium,
assurgent and fasciculate. 5 tortilis.
Appendages long or short, spreading horizontally, often interwoven with the
mycelium, i polygoni.
10. Perithecia more or less immersed in the lanuginose persistent mycelium.
4 graminis.
Perithecia not immersed in a lanuginose persistent mycelium. (11)
11. Spores four to six, 20 to 22 by 10 to i2iu. i polygoni var. sepuUa.
Spores eight, rarely six or seven, somewhat roundish, 16 to 20 by 10 to is/x.
7 aggregata.
Key to Species of Microsph^ra (After Salmon)
Brief Characterization. — Perithecia globose to globose-depressed; asci several,

two- to eight-spored. Appendages not interwoven with the mycelium, branched
in a definite manner at the apex, which is usually several times dichotomously
divided, and often very ornate, rarely {M. astragali) undivided, or once dichotomous
1. Asci two-spored, appendages densely crowded, flaccid, about equalling the

diameter of the perithecium. 6 Mougeottii.
Asci more than two-spored. (2)
2. Appendages two and one-half to seven times the diameter of the perithecium,
usually much contorted and angularly bent, apical branching very irregular
and lax, with the branches very flexuous and more or less curled. 9 euphorbia.
Appendages long or short without the above characters. (3)
3. Tips of some or all of the ultimate branches of the appendages recurved. (4)
Tips not recurved. (11)
4. Appendages eight to twelve times the diameter of the perithecium. 10
Guarinonii.
Appendages less than eight times the diameter of the perithecium. (5)
5. Appendages long and flaccid. (6)
Appendages short, not exceeding two and one-half times the diameter of the
perithecium, not flaccid. (8)
6. Apex of appendages much branched, branching ornate, more or less close
spores 22 to 26 by 4 alni var. extensa.
12 to iS/x.
Apex less branched, more or less widely, forked, or branching close and simple,
by 9 to 13M.
spores 18 to 23 (7)
7. Appendages usually three and one-half, not exceeding five and one-half times
the diameter of the perithecium, asci three to seven, ovate-globose, 38 to
48ju long. 4 alni var. divaricata.
Appendages two and one-half to eight times the diameter of the perithecium,
asci two to sixteen, ovate-oblong, 45 to 72/i long. 4 alni var. vaccinii.
8. Appendages more or less contorted, apical branching very lax and irregular.
4 alni var. liidens.
Appendages not contorted, apical branching closer and regular. (9)

9. Tips of the ultimate branches of the appendages not all regularly and dis-
tinctly recurved. 4 alni var. lonicera.

Tips all regularly and distinctly recurved. (10)
10. Axis pf some of the appendages not dividing dichotomously at the apex, but
bearing sets of opposite branches. 4 alni var. calocladophora.
Appendages regularly dichotomous at apex. 4 alni.
11. Appendages three to seven times the diameter of the perithecium, colored
nearly to apex. 8 Russdlii.
Appendages colorless. (12)
12. Appendages long and penicillate. (13)
Appendages not penicillate. (15)
APPENDIX VIII 725
13. Apex of appendages often undivided, or irregularly one to two times dichotom-
ous. 3 aslragali.
Apex more divided. (14)
14. Appendages four to six times the diameter of the perithecium, branching
diffuse and irregular. 13 Bdumleri.
Appendages two and one-half to five and one-half times the diameter of the
perithecium, apex more divided, branching closer. 2 euonymi.
1$. Branching of the appendages lax, irregular. (16)

Branching closer and regular. (17)
16. Appendages two to four times the diameter of the perithecium, not contorted,
ultimate branches long, forming a narrow fork. 7 dijfusa.
Appendages one to two times the diameter of the perithecium, more or less
contorted, branching more irregular, with short ultimate branches. 4 ahii
var. ludens.
17. Apex of appendages with very short primary and secondary branches more or
less digitate. 5 grossiilaria.
18. Apex with short, widely spreading, usually curved ultimate branches. 4 alni
var. lonicerce.
Apex with long, straight ultimate branches, not widely spreading, i berberi-
dis.
Key to the Species of Uncinula
Brief Characterization. — Perithecia globose to globose-depressed; asci several,

two- to eight-spored; appendages simple, or rarely (U. aceris) once or twice
dichotomously forked, uncinate at the apex, usually colorless, rarely dark
brown at base or throughout.
1. Appendages colored. (2)
Appendages colorless. (3)
2. Appendages colored for half their length or more. 5 necator.
Appendages colored only at base (up to first septum). 16 australiana.
3. Asci two- to three-spored. (4)
Asci four- to eight-spored. (6)
4. Asci more than thirty, perithecia very large, 215 to 320;u in diameter. 12.
polychce-ta.
Asci four to twenty, perithecia 85 to 165^ in diameter. (5)

5. Appendages, nine to twenty-five, perithecia average 95^ in diameter, asci
three to six. 4 clandestina.
Appendages fifty to one hundred and thirty, perithecia average 130^, asci
eight to twenty. 8 macrospora.
6. Appendages all simple. (7)
Appendages some or all branched. (20)
7. Appendages delicate, narrow, 3 to 4^1 wide, asci four- to seven-spored. (8)
Appendages stouter, wider, or if narrow with asci eight-spored. (10)
8. Asci about twenty-five, perithecia 150 to 200/1 diameter. 13 conjtisa.
Asci five to eight, perithecia 86 to 12 2m in diameter. (9)
9. Appendages fifty to one hundred and sixty, one-half to three-fourths diameter

of perithecium. 7 parviila.
Appendages twenty-four to forty-six, one and one-fourth to two times,
diameter of perithecium, often geniculate. 11 geniculala.
10. Appendages stout, 7 to 8^ wide near the base. (11)
Appendages narrower near the base. (12)
11. Appendages very few, six to twelve, enlarged upward. 15 Delavay's.
Appendages crowded, twenty to thirty-six, scarcely or not at all enlarged
upward. 18 Sengokui.
12. Appendages abruptly flexuose, or angularly bent. (13)
Appendages straight. (14)
* 13. Appendages about equalling diameter of perithecium, flexuose above, not
angularly bent, spores usually eight. 9 flexuosa.
Appendages one to two, usually one and one-half to two times diameter of
perithecium, more or less angularly bent, spores four to six, rarely seven.
1 solids var. Miyabci.
14. Appendages thick- walled, refractive, or rough at base. (15)
Appendages thin-walled throughout. (17)
15. Mycelium persistent, densely compacted, perithecia 158 to 268ju in diameter.
2 aceris var. Tulasnei.
Mycelium evanescent, or subpersistent, perithecia 64 to 146;^ in diameter. ( 1 6)
16. Asci ovate or elliptic-oblong, 24 to 30^ wide, spores 16 to 20 by 8 to lo/x.
3 prunastri.
Asci broadly ovate to subglobose, 34 to 40/i wide, spores 20 to 25/1 by 10 to
I3^^. 10 CUntonii.
17. Asci four- to six-spored. i salicis.
Asci seven- to eight-spored. (18)

18. Perithecia 168 to 224/x in diameter, appendages not exceeding diameter of
perithecium. 6 circinata.
Perithecia 76 to 138/i in diameter, appendages one and one-fourth to two and
one-half times diameter of perithecium. (19)
19. Perithecium 120 to 138^ in diameter, appendages thirty-five to sixty, myce-
lium persistent, more or less densely compacted. 14 aiislralis.
Perithecia 76 to 105^ in diameter, appendages ten to twenty-eight, mycelium
evanescent. 17 fraxitiis.
20. Mycelium densely compacted, appendages mostly simple. 2 accrls var.
Tulasnei.
Mycelium not densely compacted, appendages all or nearly all branched.
2 aceris.
APPENDIX IX
Collection —
and Preservation of the Fleshy Fungi. In the collection of the higher
fungi, it is of the utmost importance that certain precautions be employed in ob-
taining all parts of the plant, and furthermore that care be exercised in handling in
order not to remove or efface delicate characters. Not only is it important for the
APPENDIX DC 727
beginner, but in many instances an expert may not be able to determine a specimen
which may have lost what undoubtedly seems to some, trivial marks. The sug-
gestions given here should enable one to collect specimens in such a way as to pro-
tect these characters while fresh, to make notes of the important evanescent char-
acters dry and preserve them properly for future study. For collecting a
and to
number of specimens under a variety of conditions the following list of things is
recommended.
Implements. — One or two oblong or rectangular hand baskets, capacity 8 to
12 quarts.
One rectangular zinc case with a closely fitting top (not the ordinary botanic
case).
Half a dozen or so tall pasteboard bo.xes, or tins, 3 by 3, or 4 by 4, by 5 inches
deep, to hold certain species in an upright position.
A quantity of tissue paper cut 8 by 10, or 6 by 8 inches. Small quantity of waxed
tissue paper for wrapping viscid or sticky plants.
Trowel, a stout knife, a memorandum pad and pencil.
In gathering specimens, care should be taken to avoid leaving finger marks where
is covered with a soft and delicate outer coat.
the surface of the stem, or cap, Also
a little remove such important characters as a frail volva, or
careless handling will
annubus, which are absolutely necessary to recognize in a species. Having collected
the plants they should be placed properly in the basket, or collection case. Those
which are quite firm, and not long and slender can be wrapped with tissue paper
(waxed if the specimen is sticky), and placed directly in the basket with some
note or number to indicate habitat, or other peculiarity, which it is desirable to
make at the time of collection. The smaller, more slender and fragile specimens
can be wrapped in tissue paper made in the form of a narrow funnel and the ends
then twisted. The specimens should be placed in the basket, or case, in such a way
as to prevent jostling with the gill surfaces downward so that any loose sand, or
other material shall not fall between the gills where it is dilScult to remove such
gritty substances.
Field Notes.- —The field notes should include data on the place where the fleshy
fungi grew, the kind and character of the soil, in open field, roadside, grove, woods,
on ground, leaves, sticks, stumps, trunks, rotting wood, or on living trees, etc.
Sorting.- —
This should be done in a room with plenty of table room. This sort-
ing should be done at once as some forms deliquesce rapidly, others are attacked by
insects, while others dry rapidly, so as to lose their shape and evanescent characters.
Specimens to be photographed should be attended to at once. Some of the speci-
mens can be kept for spore prints, others must be preserved for the herbarium.
—
Drying Method. Frequently the smaller specimens will dry well when left in the
room, especially in dry weather, or better, if they are placed where there is a draft
of air. Some dry them in the sun. The most approved method is by artificial
heat. Two methods are applicable.

I. A tin oven 2 by 2 feet and 2 to several feet high with one side hinged as a door,
^ Consult Atkinson, George F.: Mushrooms, Edible and Poisonous, Etc.,
Chapter XVH.
and with several movable shelves of perforated tin, or of wire netting; a vent at the
top and "perforations around the sides at the bottom to admit air. The object of
such an oven is to provide for a constant current of air from below upward between
the specimens. This may be heated, if not too large, with a lamp, though an oil
stove, gas jet, or heater, is better. The specimens are placed on the shelves
with the accompanying notes or numbers.
2. An old cook stove can be used with wire screens 3 by 4 feet, one above the other,
placed over it.Large numbers of fleshy toadstools can be dried on such frames.
A more approved drying oven would be the revolving gas oven manufactured by
G. S. Blodgett, Burlington, Vermont.
"When the plants are dried, they become brittle but if exposed to the air a good
many kinds absorb moisture from the air so that they become pliant and can be
pressed flat, so as not to crush the gills and placed in paper envelopes for mounting
on the herbarium sheets.
When placed in herbarium they should be poisoned with a saturated solution of
alcohol and corrosive sublimate to which a spoonful of liquid carbolic acid is added.
They should then be air-dried.
Some of the specimens when there are a number of duplicates can be placed in
museum jars in 75 per cent, alcohol.
A solution of strychnine can be used for poisoning fleshy fungi.
Sulfate of strychnine, 3^8 ounce.

Warm water, 4 or 5 ounces.
Alcohol, 2 ounces.
Paper for Spore Prints. — For the identification of many species of fleshy fungi
it is necessary to make spore prints. by breaking off the stipe, if
This is best done
present, close to the under surface of the cap, or pileus, and then placing the cap
gills down on black and white paper placed side by side. Half of the gill surface
should rest on the black paper and half on the white paper, so that if the spores are
white, they will make an impression on the black paper, and if dark-colored, they
will leave an imprint on the white paper.
In all cases where a spore print is made the plant should be covered with a bell
glass to exclude currents of air. Such unprepared paper will save time in the
identification.Where, however, it is desired to obtain fancy spore prints, perfect
caps must be cut from the stipe and placed gill downward on paper prepared with
some gum arable, or similar adhesive substance, while the paper is still moist with
the fixative, so as to glue the spores as they fall to the surface of the paper. The
specimens should then be covered by a bell jar as previously directed.
Good spore prints, thus obtained, can be used for class demonstrations by mount-
ing between a piece of heavy photographic cardboard and a piece of glass. It is
easy to passepartout the glass and the paper as a museum specimen.
Blank for Nole-taking.

No. Locality
Date '
Name of collector —
Weather
APPENDICES IX, X 729
Habitat. —If on ground, low or high, wet or dry; kind of soil; on fallen leaves,
twigs, branches, logs, stumps, roots, whether dead or living. Kind of tree; in open
fields, pastures, etc., woods, groves, etc. Mixed woods or evergreen, oak, chestnut,
etc.
Plants. —Whether solitary, clustered, tufted, whether rooting or not, taste,
odor, color when bruised or cut, and if change in color takes place after exposure
to air.
Cap. — Whether dry, moist, watery in appearance (hygrophanous) slimy, viscid,
glutinous; color when young, when old; whether free from the cuticle and easily
rubbed Shape of cap.
off.
—
Margin of Cap. Whether straight or incurved when young; whether striate, or
not, when moist.
Stem. —Whether slimy, viscid, glutinous, kind of scales, if not smooth, whether
striate, dotted, granular color; when there are several specimens test one to see if it
is broken out from the cap, also to see if it is fibrous, or fleshy, or cartilaginous
easily
(firm on the outside, partly snapping and partly tough). Shape of the stem.
Gills or Tubes. —
Color when young, old, color when bruised, and if color changes
whether soft, waxy, brittle, or tough; sharp or blunt, plane or serrate edge.
Alilk. — Color if present, changing after exposure, taste.
Veil (Inner veil). —Whether present or not, character, whether arachnoid, and
if so whether free from cuticle of pileus or attached only to the edge; whether fragile,
persistent, disappearing, slimy, etc., movable, etc.
Volva. — Present or absent,
persistent or disappearing, whether it splits at apex
or is circumscribed, orcrumbly and granular or floccose, whether the part on
all
the pileus forms warts, and then the kind, distribution, shape, persistence, etc.
—
Ring. Present or absent, fragile, or persistent, whether movable, viscid, etc.
Spores. —
Color when caught on paper.
Estimation of Spore Numbers. —
Paper containing spores is placed in distilled
water. The whole is stirred vigorously until the spores have been washed off the
paper. A Leitz counting apparatus is then employed and the number of spores
per square is counted. Another method is to count spores of Coprinus comatus, for
example in situ. For details see Buller, Researches on Fungi, p. 82.
APPENDIX X
List of Keys to Fleshy Fungi and Selected Keys of Fleshy Fungi
This list includes the common accessible keys which beginners, amateurs and
students will find useful in the determination of the conspicuous fungi. The
all
list is taken from the Mj'cological Bulletin, Vol. Ill: 174; 178-179; 182-183; 185-
186, editedby W. A. Kellerman.
Amanita. Lloyd: Volvae of U. S., 3, 4, 5, 6, 1898.
McIlvaine: One Thousand American Fungi, 6, 1900.
Morgan: Journ. Mycol., 3: 25, March, 1887.
Peck: Rep. N. Y. State Mus., 23: 68, 1873; 33: 40, 1880; 48: 310, 1895.
Amanitopsis. Beardslee: Notes on the Amanitas of So. Appalachians, Part I,
Lloyd Library, September, 1902.

Lloyd: Volvac of the U. S., 8, 9, 1895.

Agaricus. McIlvaine: One Thousand American Fungi, 332, 1900.
Peck: Rep. N. Y. State Mus., 48: 231, 1895.
Armillaria. Peck: Rep. N. Y. State Mus., 43: 41, 44, 1890.
Boletinus. NinA L. Marshall: Mushroom Book, 44, 102, 1901.
Boletus. McIlvaine: One Thousand American Fungi, 406, 421, 423, 430, 436,
438, 444, 453, 459, 471, 1900.
Peck: Rep. N. Y. State Mus., 23: 127, 1873; 2>T- 58, 1884; 48: 292, 1895.
Bull. N. Y. State Mus., i: 58, May, 1887; 2: 82, 83, 106, 114, 123, 131, 138,
145, 151, "September, 1889.
Bovista. Lloyd: Myc. Notes, 12: 114, December, 1902.
Bovistella. Lloyd: Myc. Notes, 23, 1906.
Catastoma. Kellerman: Journ. Mycol., 9: 239.
Lloyd: Myc. Notes, (214), 13: 121, February, 1903.
Cantharellus. Peck: Rep. N. Y. State Mus., 23: 121, 1873; 37: 35, 1884. Bull.
N. Y. State Mus., i: 35, May, 1887.
Claudopus. McIlvaine: One Thousand American Fungi, 266, 1900.
Clavaria. McIlvaine: One Thousand American Fungi, 513, 1900.
Clitocybe. Morgan: Journ. Cin. Soc. Nat. Hist., 6: 67, 1883.
Peckk Rep. N. Y. State Mus., 23: 76, 1873; 48: 270, 1895.
Clitopilus. Beardslee: Journ. Mycol., 11: 109, May, 1905. Mycol. Bull., 3:
146, 1905.
CoUybia. Lloyd: Mycol. Notes, 34, 37, 41, December, 1900.
Morgan: Journ. Cin. Soc. Nat. Hist., 6: 70, 1883.
Coprinus. Peck: Rep. N. Y. State Mus., 23:' 103, 1873; 48: 241, 1895.
Massee, G.: Annals of Botany, X: 123-184, 1896.
Cortinarius. Earle: Torreya, 2: 169-172; 180-3, November, December, 1902.
Kauffman: BuU. Torr. Bot. Club, 32: 333, 318, June, 1905.
Peck: Rep. N. Y. State Mus., 23: 105, 107, 108, no, 112, 1873; 48: 245, 1895.
Craterellus. Peck: Rep. N. Y. State Mus., 37: 45, 1884, Bull. N. Y. State Mus.,
i: 45, May, 1887.
Crepidotus. Peck: Rep. N. Y. State Mus., 39.
Entoloma. Morgan: Journ. Cin. Soc. Nat. Hist., 6: 99, 1883.
Peck: Rep. N. Y. State Mus., 62.
Fomes. Murrill: Bull. Torr. Bot. Club, 30: 225-6, April, 1903.
Galera. Peck: Rep. N. Y. State Mus., 23: 92, 1873; 46: 62, 1893.
Ganoderma. Murrill: Bull. Torr. Bot. Club, 29: 599-608, i9o'2.
Geaster. Lloyd: 1902: 1^44.
Hebeloma. Peck: Rep. N. Y. State Mus., 23: 95, 1873; 63.
Hydnum. McIlvaine: One Thousand American Fungi, 494, 1900.
Hygrophorus. Peck: Rep. N. Y. State Mus., 23: 112, 1873; 60.
APPENDLX X 731
iQoo-
Hypholoma. McIlvaine: One Thousand American Fungi, 353, 355,
Peck: Rep. N. Y. State Mus., 23: 98, 1873; 64.
December,
Inocybe. Earle: Torreya, 3: 168-170, 183-4, November,
1903.
Lactarius. Earle: Torreya, 2: 139-41, 152-4, October, 1902.

Peck: Rep. N. Y. State Mus., 23: 114, 1873; 38-113, 1S85.
Lepiota. Morgan: Journ. Cin. Soc. Nat. Hist., 6: 60, 1883.
Peck: Rep. N. Y. State Mus., 20: 70, 1873, 35.
Lentinus. Earle: Torreya, 3: 35-8, March, 1903.
Lycoperdacese. McIlvaine: One Thousand American Fungi, 577, iQoo-
Morgan: Cin. Soc. Nat. Hist., 12:9, April, 1889.

Underwood: Moulds, Mildews and Mushrooms, 138, 1899.
Islands, 1-42;
Lloyd: Of Australia, New Zealand and Neighboring 1905:
Of the U. S. Mycol. Notes, 20, June, 1905.

Lycoperdon. McIlvaine: One Thousand American Fungi, 590, 1900-
Morgan: Journ. Cin. Soc. Nat. Hist., 13: 6, April, 1891.

Lloyd: In Europe, Mycol. Notes, 19, May, 1905.
(Fig. 264).
Marasmius. Peck: Rep. N. Y. State Mus., 23: 124, 1873
Mitremyces. Lloyd: Mycol. Notes, (218), 13: 125, February, 1903.
Mycena. Morgan: Journ. Cin. Soc. Nat. Hist., 6: 73, 1883.
Naucoria. Peck: Rep. N. Y. State Mus., 23: 91, 1873.
Nidulariaceffi. Underwood: Moulds, Mildews and Mushrooms, 142, 1899-
White: Bull. Torr. Bot. Club, 29: 254, May, 191 2.
Lloyd: 1906: 1-32.

Omphalia. Morgan: Journ. Cin. Soc. Nat. Hist., 6: 75, 1883.
Peck: Rep. N. Y. State Mus., 23: 84, 1873; 45: 33, 1893.
Panaeolus. Peck: Rep. N. Y. State Mus., 23: 100, 1873.
Panus. Earle: Torreya, 3: 86-7, June, 1903.
Peck: Rep. N. Y. State Mus., 37: 30, 1884. Bull. N. Y. State Mus.
Pa.xillus.
i: 30, May, 1887.

Phallus. McIlvaine: One Thousand American Fungi, 571, 1900.
Pholiota. Morgan: Journ. Cin. Soc. Nat. Hist., 6: loi, 1883.

Peck: Rep. N. Y. State Mus., 61.
Pleurotus. Morgan: Jour. Cin. Soc. Nat. Hist., 6: 77, 1883.

Peck: Rep. N. Y. State Mus., 39: 59, 1886; 48: 275, 1895-
Pluteolus. Earle: Torreya, 3: 124-5, August, 1903.
Pluteus. McIlv.aine: One Thousand American Fungi, 243, 1900.
Morgan: Journ. Cin. Soc. Nat. Hist., 6: 97, 1883.

Peck: Rep. N. Y. State Mus., 23: 61, 86, 1873; 38: 134, 1885.
Polyporace®. See Murrill's bibliography.
Issue, i, February, 1908.
Polystictus. Lloyd: Mycol. Notes, Polyporoid
Psalliota (Agaricus). Peck: Rep. N. Y. State Mus., 23: 97, 1893; 36: 42, 1883.
Lloyd: Mycol. Notes, 4, November, 1899.
— ———
Psathyra. Peck: Rep. N. Y. State Mus., 64.

Psathyrella. Peck: Rep. N. Y. State Mus., 23: 102, 1873.
Psilocybe. Peck: Rep. N. Y. State Mus., 23: 99, 1873; 64.
Russula. Earle: Torreya, 2: 101-3, 11 7-19, July, August, 1902.
Stropharia. Earle: Torreya, 3: 24, February, 1903.
Tricholoma. Morgan: Journ. Cin. Soc. Nat. Hist., 6: 65, 1883.
Peck: Rep. N. Y. State Mus., 23: 73, 1873; 44: 39, 40, 44, 52, 56, 61, 1891;
48: 266, 1895.
Volvaria. Lloyd: Volvaeof U. S., 10, 1898. McIlvaine: One Thousand American
Fungi, 239, 1900.
APPENDIX XI
Key to Agaricace^
The following key to the Agaricace^ is taken from Bulletin No. 175, U. S
Department of Agriculture, 1915, entitled "Mushrooms and other Common Fungi"
by Flora W. Patterson and Vera K. Charles, as well as the descriptions of a few
of the more common forms selected by way of illustration.
The classification of the genera of Agaricaceae is based upon the color of the
spores. It is generally a comparatively easy matter to form an opinion regarding
the color of the spores, but if any difficulty is experienced a spore print may be
made. The process is very simple, and the results are quite satisfactory. The
stem is removed from the specimen from which a print is desired and the cap
placed face down on pieces of black and white paper placed side by side and
covered with a tumbler. When the spores are mature they will fall in radiating
lines on the pieces of paper. If a permanent spore print is desired, an alcoholic
spray of white shellac may be employed. This is prepared by making a saturated

solution of white shellac and then diluting it 50 per cent, with alcohol.
Whites pored Agarics

Plants soft or more or less fleshy, soon decaying, not reviving
well when moistened:
Ring or volva or both present
Volva and ring both present Amanita.
Volva present, ring absent Amanitopsis.
Volva absent, ring present
Gills free from stem Lepiota.
Gills attached to the stem Armillaria.
Ring and volva both absent
Stem excentric or lateral !*
Pleurotus.
Stem central-
Gills decurrent
Edge blunt, fold-like, forked Cantharellus.
Edge thin, stem fibrous outside Clitocybe.
—— — —
APPENDIX XI 733
Edge thin, stem cartilaginous outside Omphalia.

Gills sinuate, general structure fleshy Tricholoma.
adnate or adnexed
Gills
rather fleshy, margin incurved when young
Cap Collybia.
Plants soft or more or less fleshy, etc. Continued. —
Ring and volva both absent Continued. —
Stem central — Continued.
Gills adnate or adnexed — Continued.
Cap thin, margin of the cap at first straight, mostly
bell-shaped Mycena.
Cap fleshy, gills very rigid and brittle, stem stout
Milk present Lactarius.
Milk absent Russula.
Gills various, often decurrent, adnate or only adnexed,
edge thin, thick at junction of cap, usually distant,
waxy Hygrophorus.
Plants coriaceous, tough, fleshy or membranaceous, reviving
when moistened:
Stem generally central, substance of the cap noncontinuous
with that of the stem, gills thin, often connected by veins
or ridges (Fig. 264) Marasmius.
Stem central, excentric, lateral, or absent, substance of the cap
continuous with that of the stem
Edge of gills toothed or serrate Lentinus.
Edge of gills not toothed or serrate Panus.
Edge of gills split into two laminae and revolute . Schizophyllum.
Plants corky or woody, gills inatradig Lenzites.
Rosy-s pored Agarics
Stem excentric or absent and pileus lateral Claudopus.

Stem central:
Volva present, annulus wanting Volvaria.
Volva and annulus absent
Cap easily separating from the stem, gills free Pluteus.
Cap conflaent with the stem, gills sinuate Entoloma.
Ochres pored Agarics {Spores Yellow or Brown)
Gills easily separable from the flesh of the cap:

Margin of the cap incurved, gills more or less decurrent forked
or connected with veinlike reticulations Paxillus.
Gills not easily separable from the flesh of the cap:
Universal veil present, arachnoid Cortinarius.

—— —
Universal veil absent

Ring present Pholiota.
Ring absent
Stem central
Cap turned in Naucoria.
Cap not turned in Galera.
Stem excentric or none Crepidotus.
Browns pored A garics
Cap easily separating from the stem, gills usually free Agaricus.
Cap not easily separating from the stem, gills attached:
Ring present. Stropharia.
Ring absent, veil remaining attached to the margin of the cap. . Hypholoma.
Blacks pored Agarics
Gills deliquescing, cap thin, ring present in some species Coprinus.

Gills not deliquescing:
Margin of cap striate, gills not variegated Psathyrella.
Margin of cap not striate, gills variegated Panôlus.
The genus Amanitais, easily recognized among the white-spored agarics in typical
species, or early stages, by the presence of a volva and a veil. Young plants are com-
pletely enveloped by the volva, and the manner in which it ruptures varies according
to the species. The volva may persist in the form of a basal cup, as rings, or scales,
on a bulb-like base, or it may be friable and evanescent. The cap is fleshy, convex,
then expanded. The gills are free from the stem, which is different in substance
from the cap and readily separable from it.
This is a most interesting genus, on account of the great beauty of color and tex-
ture of many of its species and the fact that it contains the most poisonous of all
mushrooms. While there are some edible species in the genus, the safest policy
for the amateur is to avoid all mushrooms of the genus Amanita.
Amanita caesarea. CcEsar's Mushroom
Cap ovate to hemispherical, smooth, with prominently striate margin, reddish or

orange becoming yellow; gills free, yellow; stem cylindric, only slightly enlarged
at the base, attenuated upward, flocculose, scaly below the annulus, smooth above;
ring membranaceous, large, attached from its upper margin; stem and ring nor-
mally orange or yellowish, in small or depauperate specimens sometimes white;
flesh white, yellow under the skin, and usually yellow next to the gills; volva large,
distinct, white, sac-like.
Cap 2 3^^ to 4 or more inches broad stem 3 to s inches long.
;
This species is variously known as Csesar's agaric, royal agaric, orange Amanita,
APPENDIX XI 735
etc. It has been highly esteemed as an article of diet since the time of the early
Greeks. It is abundant during rainy weather and may occur solitary,
particularly
several together, or in definite rings. Although this species is edible, great caution
should always be used in order not to confound it with Amanilar Frostlana, which is
poisonous. The points of difference of these two species are conveniently compared
as follows:
Fig. 264. —
Fruit bodies of fairy-ring toadstool (Marasmiits oyeades). {After
Patterson, Flora W., and Charles, Vera K., Bull. 175, U. S. Dept. Agric, pi. xix,
Apr. 29, 1915-)
Species
volva persisting as scattered, floccose, or rather compact scales, color subject to

great variation, ranging from yellow to orange, or blood red, gills white or yellow-
ish, free but reaching the stem; stem cylindrical, at first stuffed, later hoUow, upper
part torn into loose scales, bulb prominent, generally marked by concentric scales
forming irregular ridges; ring typically apical, lacerated, lax, large.
Cap $}^ inches broad, stem 4 to 6 inches long.

33-2 to
Amanita muscaria may be found during the summer and fall, occurring singly, or
in small associations, or in patches of considerable size. It grows in cultivated soil,
partially cleared land, and in woods or roadsides. It does not demand a rich soil,
but rather exhibits a preference for poor ground. The color is an exceedingly vari-
able character, the plants being brighter colored when young, and fading as they
mature. The European plant possesses more gorgeous colors than the American
form.
This is a very poisonous species, and it has been the subject of many pharmaco-
logical and chemical investigations. Its chief poisonous principle is muscarine,

although a second poisonous element is believed to be present, as atropine d(Jes not
entirely neutralize the effect of injections of Amanita muscaria in animals.
This species has been responsible for many deaths, and numerous cases of severe
illness have been caused by persons mistaking Amanita muscaria, the poisonous
species, for Amanita caesarea, the edible species. The most satisfactory treatment
is to administer hypodermic injections of atropine beginning with a dosage of }io
grain after the giving of a strong emetic. While typical specimens of these two
species possess distinguishing characters, as already shown, it is again recommended
to shun all Amanitae.
In Siberian Russia the natives make several uses of Amanita muscaria. Pre-
served in salt it is eaten, though probably more as a condiment than as a main
article of diet; a decoction is popular as an intoxicant, and deaths are reported upon
good authority as resulting from a "muscaria orgy."
Amanita phalloides. Death Cup {Deadly Poisonous)
Cap white, lemon, or olive to umber, fleshy, viscid when moist, smooth or with
patches or scales, broadly oval, bell-shaped, convex, and finally expanded, old speci-
mens sometimes depressed by the elevation of the margin; gills free, white; stem
generally smoothand white, in dark varieties colored like the cap but lighter, solid
downward, bulbous, hollow, and attenuated upward; ring superior, reflexed, gener-
all}^ entire, white.
The large, free volva, its lower portion closely adherent to the bulb, and the large
ring are of assistance in distinguishing this species.
Cap 3 to 4 inches broad; stem 3 to 5 inches long.
This species and its forms are subject to great variation in color, ranging from
white, pale yellow, and olive to brown. Amanita phalloides is a very cosmopolitan
plant and one of very common occurrence. It is the most dangerous of all mush-
rooms, for no antidote to overcome its deadly effect is known. It exhibits no special
preference as regards habitat and is found growing in woods or cultivated land from
APPENDIX XI -
737
summer to late autumn. When fresh it is without scent, but a peculiarly sickening
odor is present in drying plants.
Armillaria
The genus Armillaria is another white-spored agaric having a ring and no volva.
The gills are attached to the stem and are sinuate or more or less decurrent. The
substance of the stem and cap is continuous and firm. This genus may be distin-
guished from Amanita and Lepiota by the continuity of the substance of the stem
and cap, and it is further differentiated from Amanita by the absence of a volva.
It contains several edible species.
Armillaria mellea. Honey-colored Mushroom {Edible)
Cap oval convex and expanded, sometimes with a slight elevation, smooth, or
to
adorned with pointed dark-brown or blackish scales, especially in the center, honey
color to dull reddish-brown, margin even or somewhat striate when old; gills adnate
or decurrent, white or whitish, sometimes with reddish-brown spots; stem elastic,
spongy, sometimes hollow, smooth or scaly, generally whitish, sometimes gray or

yellow above the ring, below reddish-brown.
Cap i^^ to 6 inches broad; stem 2 to 6 inches long, ^-'2 to % inch thick.
This species is extremely common and variable. It generally occurs in clusters
about the base of rotten stumps and is often a serious parasite of fruit trees and
destructive to props in coal mines. The fruit bodies are attached to the strands of
hyphae known as Rhizomorpha subterranea, which form a network under the bark
of the tree and out into the soil. Both ring and stem are subject to marked varia-
tions. The former may be thick, or thin, or entirely absent, and the latter uniform
in diameter or bulbous. The species is edible, though not especially tender or
highly flavored (Fig. 15).
On account of the great variation in color, surface of the cap, and shape of the
stem, several forms of Armillaria mellea have been given varietal distinction. The
following varieties as distinguished by Prof. Peck may be of assistance to the amateur:
Armillaria mellea var. flava, with yellow or reddish-yellow cap.

Armillaria mellea var. radicata, with a tapering root.
Armillaria mellea var. albida, with white or whitish cap.
Pleurotus
The genus Pleurotus is chiefly distinguished among the white-spored agarics by

the excentric stem or resupinate cap. The stem is fleshy and continuous with the
substance of the cap, but it is subject to great variation in the different species and
may be excentric, lateral, or entirely absent. The gills are decurrent or sometimes
adnate, edge acute. Most of the species grow on wood, buried roots, or decayed
stumps. This genus corresponds to Claudopus of the pink-spored and Crepidotus
of the brown-spored forms.
47
Pleurotiis oslreatus. Oyster Mushroom (Edible)
Cap either sessile or stipitate, shell-shaped or dimidiate, ascending, fleshy, soft,

smooth, moist, in color white, cream, grayish to brownish ash; stem present or absent
(if present, short, firm, elastic, ascending, base hairy); gills white, decurrent, some-
what anastomosing behind to form an irregular network.
distant,
Cap 3 to 5 inches broad;
mostly cespitose imbricated (Fig. 265).
A very fine edible species, growing on limbs or trunks of living or dead trees, of
cosmopolitan distribution, appearing from early summer until late fall.
Fig. 265. — Sporophores of oyster toadstool {Pleurotus oslreatus). {After Patter-

son, Flora W., and Charles, Vera K., Bull. 175, U. S. Dept. Agric. pi. vii, Apr. 29,
191S.)
Pleurotus sapidus (Edible)
This species very closely resembles Pleurotus ostrealus and is distinguished from
it by the lilac-tinged spores, a character difficult or impossible for the amateur to
detect. From the mycophagist's point of view, these two species are equally
attractive.
Pleurotus serotinus (Edible)
Cap compact, convex or nearly plane, dimidiate reniform, suborbicular,

fleshy,
edge involute, finally wavy, smooth, yellowish-green, sooty olive, or reddish-brown,
in wet weather with a viscid pellicle; gills close, distinct, whitish or yellowish,
minutely tomentose or squamulose with blackish points.
Cap I to 3 inches broad.
APPENDIX XI 739
In general appearance this fungus resembles Claudopiis nidulans, but is sepa-

rated from it by the color of the spores, Pleurohis belonging to the section of white-
spored agarics and Claudopus to the rosy-spored species. The plants grow on dead
branches or trunks and are gregarious or imbricate.
Pleurotus serotinus is edible but not particularly good, its chief recommendation
being the lateness of its occurrence in the fall, when other more tempting species
have disappeared.
Pleurotus ulmarius {Edible)
Cap although inclined to excentricity, convex, margin incurved,

fairly regular,
later plane, horizontal, even, smooth, white or whitish, at disk shades of tan or
brown; flesh white, tough; gills broad, rather distant or rounded behind; stem more
or less excentric, curved, ascending, firm, solid, elastic, thickened, and tomentose at
the base.
Cap 3 to 5 inches broad, stem 2 to 3 inches long.
This species occurs abundantly on dead elm branches or trunks or growing from
wounds of living trees. Though exhibiting a special fondness for this host, it is not
confined to elm trees. It is readily distinguished from Pleurotus ostreatus by the
long stem and by the emarginate or rounded gills. It is considered an excellent
edible species and occurs abundantly in the fall.
Cantharellus
In the genus Cantharellus the cap is fleshy or submembranaceous, continuous

with the stem, and has the margin entire, wavy, or lobed. The gills are decurrent,
thick, narrow, blunt, fold-like, irregularly forked, and connected by net-like veins.
Cantharellus aurantiactis. False Chanterelle
Cap fleshy, soft, somewhat shape variable, convex, plane or infundibuli-

silky,
form, margin wavy or lobed, inroUed when young, later simply incurved, dull orange
or brownish, especially in the center; flesh yellowish; gills rather thin, decurrent,
forked, dark orange; stem spongy, fibrous, colored like the cap, larger at the base
than at the apex.
Plant I to 3 inches in height; cap i to 3 inches broad.
This plant is more slender and the gills are thinner than those of Cantharellus
cibarius,from which it can be readily distinguished. The taste is generally mild,
but sometimes slightly bitter. Foreign and American mycophagists do not agree in
regard to the edibility of the species. It is common on the ground or on very rotten
logs.
Cantharellus cibarius. The Chanterelle (Edible)
Cap smooth, irregularly expanded, sometimes deeply depressed,

fleshy, thick,
opaque egg yellow, margin sometimes wavy; flesh white; gills decurrent, thick
narrow, branching or irregularly connected, same color as cap; stem short, solid
expanding into a cap of the same color.
Plant 2 to 4 inches in height; cap 2 to 3 inches broad.

An agreeable odor of apricots may
be observed, especially in the dried plants of
this species, but its absence need not be construed as affecting the validity of an
identification established by other characters. The chanterelle has long been con-
sidered one of the most highly prized edible mushrooms. The remark of a foreign
mycologist is recalled that "The chanterelle is when the most costly
included
dainties are sought for state dinners." It is a common summer species found in
open woods and grassy places.
Lactarius
The distinguishing feature of the genus Lactarius is the presence of a white or

colored milk, especially in the gills. The entire plant is brittle and inclined to
rigidity. The fleshy cap is more or less depressed and frequently marked with
concentric zones. The gills are often somewhat decurrent, but in certain species
are adnate or adnexed, unequal in length, and often forked. The stem is stout,
rigid, central, or slightly excentric.
Lactarius chelidonium (Edible)
Cap convex and depressed in the center, glabrous, slightly viscid when moist,
firm,
grayish-yellow or tawny, at length stained bluish or greenish, generally zonate, mar-
gin involute at first and naked; gills narrow, crowded, sometimes forked, and some-
times joining to form reticulations, adnate or slightly decurrent, saffron yellow to
salmon; stem short, nearly equal, hollow, colored like the cap.
Cap 2 to 2^^ inches broad; stem i to i}^ inches long, about 3'2 inch thick.
This species is closely related to Lactarius deliciosus, to which in flavor and sub-
stance it is scarcely inferior. It is paler than that species and the milk is saffron
yellow rather than orange. The plants are fragile and when wounded turn blue,
and later green. They are to be found especially in dry localities in the vicinity of
pine woods in September and October.
Lactarius deceptivus (Edible)
Cap convex umbilicate, then expanded and centrally depressed, somewhat

fleshy,
infundibuliform, white or whitish, margin at first involute, covered with a dense soft
cottony tomentum, filling the space between the margin and the stem, finally spread-
ing or elevated and more or less fibrillose; gills whitish or cream-colored, rather
broad, distant or subdistant, adnate or decurrent, forking; stem solid, nearly equal,
pruinose-pubescent.
Cap 2^^^ to 53'^ inches broad; stem % inch to 3 inches long.
Lactarius deceptivus isfound in woods and open places from July to September.
It is coarse, but fairly good after its peppery taste is lost by cooking.
Lactarius deliciosus (Edible)
Cap convex, but depressed in the center when quite young, finally funnel-shaped,
smooth, slightly viscid, deep orange, yellowish or grayish-orange, generally zoned.
1
APPENDIX XI 741
margin naked, at first involute, unfolding as the plant becomes infundibuliform;

flesh soft, pallid; gills crowded, narrow, often branched, yellowish-orange; stem
equal or attenuated at the base, stuffed, then hollow, of the same color as the cap
except that it is paler and sometimes has dark spots.
Cap 2 stem i to 2 inches long, i inch thick.
to 5 inches broad;
This fungus is on account of its orange color and the concentric zones
distinctive,
of light and dark orange on the cap and because of the saffron red or orange milk.
A peculiarity of the plant is that it turns green upon bruising and in age changes from
the original color to greenish. Lactarius deliciosus is widely distributed and of com-
mon occurrence, appearing on the ground in woods, solitary or in patches, from June
or July to October. As the name indicates, it is considered a delicious species, and
that it has a preeminent claim to the name is unchallenged. Even by the ancients
it was considered "food for the gods."
Lactarius fumosus (Suspicious)
Cap convex, plane or slightly depressed, snuff brown or coffee-colored, dry gla-
brous or pruinose, very smooth, margin entire or sometimes wavy; flesh white,
changing to reddish when wounded; gills subdistant, adnate, or slightly decurrent,
white then yellow, becoming pinkish or salmon where bruised; stem nearly equal or
slightly tapering downward, stuffed, then hollow, colored like the cap.
Cap 2 to 3 inches broad; stem i^ to 2,1/^ inches long, about 6 lines thick.
This species varies considerably in size, color, and closeness of the gills. The
distinguishing features for field identification are the coffee-colored cap and the
changeable color of the flesh and gills. Its use should be strictly avoided, as it
closely resembles Lactarius fidiginosus, a poisonous species. These two species,
L. fumosus and L. fuUginosus, are sometimes considered identical.
Lactarius indigo (Edible)
Cap at first umbilicate and the margin involute, later cap depressed or infundibuli-
form and margin elevated, indigo blue with a silvery-gray luster, zonate, fading in
age, becoming greenish and less distinctly zoned, milk abundant and dark blue;
gills crowded, indigo blue, changing to greenish in age; stem short, nearly equal,
hollow.
Cap 2 to 5 inches broad; stem i to 2 inches long.
Lactarius indigo is easily recognized by its striking blue color. It occurs in mixed
Qr coniferous woods in summer and autumn. Though not particularly abundant,
several plants are generally found in fairly close range of one another.
Lactarius piperatus. Pepper Cap (Edible)
Cap fleshy, thick, convex, umbilicate, when mature funnel-shaped, even, smooth,
zoneless, margin involute when young; flesh white; gills narrow, crowded, edge
1 BURLINGHAM, GERTRUDE S. : Study of the Lactari« of the United States.
Memoirs, Torr. Bot. Club, Vol. 14, No. i, p. 84, 1908.
obtuse, in some forms arcuate, and then extended upward, white, reported wish
occasional yellow spots; stem equal or tapering below, thick, white, sometimet
pruinose.
Cap 3)-^ to 5 inches broad, sometimes reported considerably larger; stem i to
inches long.
The mUk "pepper cap" is abundant, white, unchangeable, and extremely
in the
acrid, to which character is due the specific name. This species is very common and
abundant from June to October.
Lactarius torminosus (Poisonous)
Cap convex then depressed, surface viscid when young or moist, yellowish-red or
ochraceous with pink shades, margin involute when young, persistently tomentoes
hairy; gills crowded, narrow, often tinged with yellow or flesh color; stem cylin-
drical or slightly tapering at the base, hollow, whitish.
Cap 2 to 3H inches broad; stem iM to 3 inches long, 4 to 8 ilnes thick.
According to some authors this species is injurious only when raw. It is cooked
and eaten in Sweden. In Russia it is enjoyed dressed with oil and vinegar or it
is preserved by drying.
Lactarius volemus (Edible)
Cap convex, nearly plane or slightly depressed, glabrous, dry, azonate, brownish
terra cotta, somewhat wrinkled when old; gills adnate or slightly decurrent, close,
whitish, becoming sordid or brownish when bruised; stem more or less equal, firm,
solid, glabrous, colored like the cap or paler; milk white, abundant, and mild, be-
coming thick when exposed to the air.

Cap 2 to 5 inches broad; stem i to 4 inches long, 4 to 10 lines thick.
This species is considered delicious, and is quite common from midsummer to
frost on semicleared or sprout land.
•
RUSSULA
The genus Russulais similar in form, brittleness, and general appearance to
Lactarius,from which it differs only in the absence of milk. The species are very
abundant in the summer, extending into the fall months.
Most species of Russula are regarded as edible, but several are known to be
poisonous. It is advisable to abstain from eating any red forms until perfectly
familiar with the different species.
Russula emelica (Poisonous)
Cap oval becoming flattened or depressed, smooth, shining, rosy

to bell-shaped,
to dark red when tawny, sometimes becoming yellow, margin finally-
old, fading to
furrowed and tuberculate; flesh white, but reddish under the separable pellicle;
gills nearly free, somewhat distant, shining white; taste very acrid; stem stout,
spongy-stuffed, fragile when old, white or reddish.

APPENDIX XI 743
Cap stem 2^2 to 4 inches long.

3 to 4 inches broad;
Russula emetica is a handsome plant of wide distribution found during summer
and autumn on the ground in woods or open places. Although some enthusiastic
mycophagists testify to its edibility, it is best to consider the species poisonous.
Russula ochrophylla
Cap convex, becoming nearly plane or very slightly depressed in the center, when
old purple or purplish red, margin even, sometimes faintly striate when old; flesh
white, purplish under the cuticle; gills adnate, entire, a few forked at the base, inter-
spaces somewhat venose, at first yellowish, ochraceous buff when mature, powdery
from the spores; stem mostly equal, solid or spongy within, rosy or red, paler than
the cap.
Cap 2 to 4 inches broad; stem 2^^ to 3 Inches long.
Russula ochrophylla may be found growing singly, or in small patches on the
ground in woods, mostly under trees, according to Prof. Peck, especially under oak
trees. In Virginia, Maryland, and the District of Columbia it is abundant in July
and August and is to be found less frequently in September and the first part of
October.
Russula roscipcs {Edible)
Cap conve.x, sometimes plane or slightly depressed, at first viscid, then dry and
faintly striate on the margin, rosy red, frequently modified by pink or ochraceous
shades; gills moderately close, ventricose, more or less adnate, whitish becoming
yellow; stem stout, stuffed or somewhat hollow, white tinged with red.
Cap I to 2 inches broad; stem i3'^ to 3 inches long.
This species grows on the ground in mixed, but generally coniferous, woods. It
appears in the late summer and autumn and is reported excellent, though, as already
stated, the amateur should be cautious and avoid all red species of this genus.
Russula rubra
Cap convex, flattened, finally depressed, dry, pellicle absent, polished, cinnabar
red, becoming tan when old; flesh white, reddish under the cuticle; gills adnate,
somewhat crowded, whitish then yellowish, often red on the edge; stem stout, solid,
varying white or red.
Cap 2% to 4 inches broad; stem 2 to 3 inches long, about i inch thick.
This species is extremely acrid, and, as there are conflicting opinions concerning
its edibility, it is best for the amateur to refrain from collecting it. It is found in
woods on the ground in summer and autumn.
Russula viresccns {Edible) .
Cap at first rounded, then expanded, when

somewhat depressed in the center,
old
dry, green, the surface broken up into quite regular, more or less angular areas of
deeper color, margin straight, obtuse, even; gills adnate, somewhat crowded, equal
or forked; stem equal, thick, solid or spongy rivulose, white.
Cap 3^-^ to 5 inches broad; stem about 2 inches long.

This fungus is noticeable on account of the color and areolate character of the
cap. In Virginia, Maryland, and the District of Columbia it occurs commonly either
solitary or in small patches, but not in very great abundance,
from July to September,
but it has been found from June through the entire summer and into October. The
species is edible and of good flavor.
CORTINARIUS
The genus Corlinarius is easily recognized when young among the ocher-spored
agarics by the powdery gills and by the cobwebby veil, which is separable from the
cuticle of the cap. In mature plants the remains of the veil may often be observed
adhering to the margin of the cap and forming a silky zone on the stem. Corlinarius
contains many forms which are difficult of specific determination. Many species
are edible, some indifferent or unpleasant, and others positively injurious. The
colors are generally conspicuous and often very beautiful. Most of the species
occur in the autumn.
Corlinarius cinnamomeus (Edible)
Cap rather campanulate, when expanded almost plane, but sometimes

thin, conic
umbonate, yellow to bright cinnamon-colored, with perhaps red stains, smooth, silky
from innate, yellowish fibrils, sometimes concentric rows of scales near the margin;
flesh yellowish; gills yellow, tawny, or red, adnate, slightly sinuate and decur-
rent by a tooth, crowded, thin, broad; stem equal, stuffed then hollow, yellowish,
fibrillose.
Cap I to 2}^ inches broad; stem2 to 4 inches long, 3 to 4 lines thick.
This is a very common and widely distributed species, particularly abundant in

mossy coniferous woods from summer until fall. The color of the gills is an extremely
variable character, ranging from brown or cinnamon to blood red. A form possess-
ing gills of the latter color is known as Corlinarius cinnamomeus var. semisanguineus.
This species and variety are edible and considered extremely good.
Corlinarius liloiinus {Edible)
Cap firm, hemispherical, then convex, minutely silky, lilac-colored; gills close,
violaceous changing to cinnamon; stem solid, stout, distinctly bulbous, silky fibril-
lose, whitish with a lilac tinge.
Cap 2 to 3 inches broad;stem 2 to 4 inches long.

This is a comparatively rare but very beautiful mushroom and an excellent edible
species.
Corlinarius sanguineus {Edible)
Cap convex, then plane, or perhaps slightly umbonate or depressed, blood red,
silky or squamulose; flesh paler reddish; gills crowded, entire, adnate,dark blood
red; stem stuffed or hollow, sometimes attenuated at the base, dark as the cap and
fibrillose, containing a red juice.
APPENDIX XI 745
Cap I to 1% inches broad; stem 2 to 3 inches long.

This species is much less common in its occurrence than Corlinarius cinnamomeus,
but is distinctive because of its entire blood-red color.
Corlinarius violaceus {Edible)
Cap convex, when expanded almost plane, dry with hairy tufts or scales, dark
violet; flesh somewhat violaceous; gills distant, rather thick and broad, rounded or
deeply notched at apex of stem, narrowed at margin of cap, at first violaceous, later
brownish-cinnamon; stem fibrillose, solid, bulbous, colored like cap.
Cap 2 to 4 inches broad; stem 3 to 5 inches long.
This very attractive species is at first a uniform
violet, but with age the gills
assume a cinnamon hue. The plants appear in woods and open places during the
summer and fall, generally solitary, but often in considerable numbers. It is
esteemed as one of the best edible species.
Agaricus
The genus Agaricus is characterized by brown or blackish spores with a purplish

tinge and by the presence of a ring. The cap is mostly fleshy and the gills are free
from the stem. The genus is closely related by Stropharia, but separated from it
by the free gills and the noncontinuity of the stem and the cap. The species of
Agaricus occur in pastures, meadows, woods, and manured ground. All are edible,
but certain forms are of especially good flavor. Bright colors are mostly absent
and white or dingy brown shades predominate.
Agaricus arvensis. Horse or Field Mushroom (Edible)
Cap convex, bell-shaped, then expanded, when young floccose or mealy, later
smooth, white or yellowish; flesh white; white to pink, at length blackish-brown,
gills
free, close, may be broader toward the stem; stem stout, hollow or stuffed, may be
slightly bulbous, smooth; ring rather large, thick, the upper part white, membrana-
ceous, the lower yellowish and radially split.
Cap 3 to 5 inches broad; stem 2 to 5 inches high, 4 to 10 lines thick.
Agaricus arvensis is to be found in fields, pastures, and waste places. It is closely
related to the ordinary cultivated mushroom, but differs in its larger size and double
ring. It is an excellent edible species, the delicacy of flavor and texture largely
depending, other mushrooms, upon its age.
like
Agaricus campestris. Common or Cultivated Mushroom (Edible)
Cap rounded, convex, when expanded nearly plane, smooth, silky floccose or
squamulose, white or light brown, squamules brown, margin incurved; flesh white,
firm; gills white in the button stage, then pink, soon becoming purplish-brown, dark
brown, or nearly black, free from the stem, rounded behind, subdeliquescent; stem
white, subequal, smooth or nearly so; veil sometimes remaining as fragments on the
margin of cap; ring frail, sometimes soon disappearing.
Cap 1 3^ stem 2 to 3 inches long, 4 to 8 lines thick. (Fig. 266.)

to 4 inches broad;
This is the most common and best known of all the edible mushrooms. It is a
species of high commercial value, lending itself to very successful and profitable
artificial cultivation. It is cosmopolitan in its geographic distribution, being as
universally known abroad as in America. It is cultivated in caves, cellars, and in
especially constructed houses; but it also occurs abundantly in the wild state, appear-
ing in pastures, grassy places, and richly manured ground. The only danger in
collecting it in the wild form is in mistaking an Amanita for an Agaricus; however,
this danger may be obviated by waiting until the gills are decidedly pink before col-
lecting the mushrooms.
Fig. 266. —
Meadow mushroom, Agaricus campestris var. Columbia, showing all
stages in development of young mushrooms (fruit bodies). {From Gager, after G. F.
Atkinson.)
Agaricus placomyces. Flat-cap Mushroom {Edible)
Cap thin, at first broadly ovate, convex or expanded and flat in age, whitish,
adorned with numerous minute, brown scales, which become crowded in the center,
forming a large brown patch; gills close, white, then pinkish, finally blackish-brown;
veil broad; ring large. In the early stages, according to Prof. Atkinson, a portion of
the veil frequently encircles the stipe like a tube, while a part remains still stretched
over the gills.
APPENDIX XI 747
Stem smooth, stuffed or hollow, bulbous, white or whitish, the bulb often
stained with yellow.
Cap 2 to 4 inches broad; stem 3 to 5 inches long, 3^ to K
inch thick.
This species frequents hemlock woods, occurring from July to September.
Agaricus Rodmani {Edible)
Cap firm, rounded, convex, then nearly plane, white, becoming subochraceous,
smooth or cracked into scales on the disk, margin decurved; flesh white; gills nar-
row, close, white, changing to pink and blackish-brown; stem solid, short, whitish,
smooth, or perhaps mealy, squamulose above the ring; ring double, sometimes ap-
pearing as two collars with space between.
Cap 2 to 4 inches broad; stem 2 to 3 inches long, 6 to 10 lines thick.
Agaricus Rodmani may easily be mistaken for Agaricus campestris, but can be dis-
tinguished by the thicker, firmer flesh, narrower gills, which are nearly white when
young, and peculiar collar, which appears double. This species grows on grassy
ground, often springing from crevices of unused pavements or between the curbing
and the walk. It is to be found principally from May to July.
Agaricus silvicola (Edible)
Cap convex, expanded to almost plane, sometimes umbonate, smooth, shining,

white, often tinged with yellow, sometimes with pink, especially in the center; flesh
white or pinkish; gills thin, crowded, white, then pink, later dark brown, distant
from stem, generally narrowed toward each end; stem long, bulbous, stuffed or hol-
low, whitish, sometimes yellowish below; ring membranaceous, sometimes with
broad floccose patches on the under side.
Cap 3 to 6 inches broad; stem 4 to 6 inches long, 4 to 8 lines thick.
Agaricus silvicola has been known under various names, at one time being consid-
ered merely a variety of Agaricus arvensis. By Peck^ it has been recognized as a
distinct species, A abrupiibulbus.
. A discussion of the nomenclature of this species
may be found in Mcllvaine and Macadam.*
Agaricus subrufescens (Edible)
Cap at deeply hemispherical, becoming convex or broadly expanded, silky,

first
fibrillose, and minutely or obscurely squamulose, whitish, grayish, or dull red-

dish-brown, usually smooth and darker on the disk; flesh white, unchangeable;
gills at first "white or whitish, then pinkish, finally blackish-brown; stem rather long,
often somewhat thickened or bulbous at the base, at first stuffed, then hollow, white;
the annulus flocculose or floccose squamose on the lower surface. Two additional
1 Peck, C. H.: Report of the State Botanist, 1904. N. Y. State Mus. Bull. 94,
p. 36, 1905.
2McIlvaine, Charles, and Macadam, R. K.: Toadstools, Mushrooms, Fungi,
Edible and Poisonous; One Thousand American Fungi, rev. ed., Indianapolis
(1912), p. 728.
characters of assistance in identification are the mycelium, which forms slender

branching root-like strings, and the almond-like flav'or of the flesh.
Cap 3 to 4 inches broad; stem aj-^ to 4 inches long.
The plants often grow in large clusters of twenty to thirty or even forty indi-
viduals. They occur in the wild state and have also been reported as a volunteer
crop in especially prepared soil. Specimens collected Washington,
in the vicinity of
Fig. 267. —
Fruit bodies of Coprinus alramenlariiis (edible). {After Patterson, Flora
W., and Charles, VercfK., Bull. 175, U. S. Dept. Agric, pi. xxviii, Apr. 25, ipiS-)
D. C, were found growing near the river on a rocky slope rich in leaf mould. Agari-
cus subrufescens is considered a very excellent edible species.
COPRINUS
The genus Coprinus is easily recognized by the black spores and the close gills,
which at maturity dissolve into an inky fluid. The stem is hollow, smooth, or
fibrillose. The volva and ring are not generic characters, but are sometimes pres-
ent. The plants are more or less fragile and occur on richly manured ground, dung,
or rotten tree trunks. The genus contains species of excellent flavor and delicate
consistency. Autodigestion (page 65) is shown by them.
APPENDIX XI 749
Coprinus atramenlarius. Inky Cap {Edible) (Fig. 267).
Cap ovate, slightly expanding, silvery to dark gray or brownish, smooth, silky or
with small scales, especially at the center, often plicate and lobed with notched mar-
gin; gills broad, ventricose, crowded, free, white, soon changing to pinkish-gray,
then becoming black and deliquescent; stem smooth, shining, whitish, hollow,
Fig. 268. —Edible shaggymane, Coprinus comatus. {After Patterson, Flora W., and
Charles, Vera K., Bull. 175, U. S. Dept. Agric, pi. xxii, Apr. 29, 1915.)
attenuated upward, readily separating from the cap; ring near the base of stem,
evanescent.
Cap 13-2 to 4 inches broad; stem 2 to 4 inches long, 4 to 6 lines thick.
This species appears from spring to autumn, particularly after rains. It grows
singly or in dense clusters on rich ground, lawns, gardens, or waste places. It has
long been esteemed as an edible species. Coprinus atramenlarius differs from C.
comalus in the more or less smooth, oval cap and the imperfect, basal, evanescent
ring.
Coprinus contains. Shaggy Mane (Edible) (Figs. 268 and 270).
Cap oblong, bell-shaped, not fully expanding, fleshy at center, moist, cuticle
separating into scales that are sometimes white, sometimes yellowish or darker, and
show the white flesh beneath, splitting from the margin along the lines of the gills;
gills broad, crowded, free, white, soon becoming pink or salmon-colored and chang-
ing to purplish-black just previous to deliquescence; stem brittle, smooth or fibril-
FiG. 269. — Glistening inky cap, Coprinus micaceus. (Pholo by W. H. Walmsley.)
lose, hollow, thick, attenuated upward, sometimes slightly bulbous at base, easily
separating from the cap; ring thin, movable.
Cap usually ij'^ to 3 inches long; stem 2 to 4 inches long, 4 to 6 lines thick.
This species has a wide geographic distribution and is universally enjoyed by
mycophagists. The fungus is very attractive when young, often white, again show-
ing gray, tawny, or pinkish tints. It appears in the spring and fall, sometimes soli-
tary, sometimes in groups, on lawns, in rich soil, or in gardens.
APPENDIX XI 751
Coprinus fimetarius
Cap at first cylindrical, later conical to expanded, margin splitting, revolute or

upturned, grayish to bluish-black, surface at first covered with white scales, finally
smooth; gills black, narrow; stem fragile, white, squamulose, hollow, but solid and
bulbous at the base.
Cap I inch or more across, stem 3 or more inches high.
This is a very common and abundant species on manure or rich soil and occurs
from spring to winter. It is edible and considered excellent.
it Hi
Fig. 270. — Shaggymane

toadstool {Coprinus comatus) growing in open fields
and on lawns. Edible before it begins to deliquesce. {After Gager, C. S.: Funda-
mentals of Botany, 1916: 289.)
Coprinus micaceus. Mica Inky Cap (Fig. 269).
Cap ovate, bell-shaped, light tan to brown, darker when moist or old, often
glistening from minute, mica-like margin closely striate, splitting, and revo-
scales,
lute; gills narrow, crowded, white, then pink before becoming black; stem slender,
white, hollow, fragile, often twisted.
Cap I to 2 inches broad; stem 2 to 4 inches long and 2 to 3 lines thick.
This glistening little species occurs very commonly at the base of trees or spring-
ing from dead roots along pavements, or more uncommonly on prostrate logs in
shady woods. The plants appear in great profusion in the spring and early summer,
and more sparingly during the fall. Coprinus micaceus is a very delicious mush-
room and lends itself to various methods of preparation.
1
INDEX
A list of the common and important diseases of economic plants in the United
States and Canada be found on pages 414 to 474. The scientific names of
will
the various disease-producing organisms and their common names will be found
there, arranged alphabetically according to the host plants on which they grow.
These names have been omitted from this index.
Abnormalities, classification of, 331 Agaricus arvensis, description of, 745

Abortion, 331 Agaricus campestris, analysis of, 55;
Abrasion, 294 fat content, 56; fed to Plasmodium,

Acaulosy, 331 12; figure of, 234, 746; description
Account of specific plant diseases, ofj 745) 746; number of spores, 234,
475 et. seq. Agaricus, description of genus, 745

Acetic acid fermentation, 32 Agaricus placomyces, description of, 746
Acheilary, 332 Agaricus Rodmani, description of, 747
A-chlya, figures of species, 112 Agaricus silvicola, description of, 747
Achlya polyandra on water plants, 1 1 Agaricus spectabilis, resin in, 56
Achlya prolifera, zoospores of, 67 Agaricus subrufescens, description of,
Acid injuries, 649 747

Acid spotting of morning glories, 293 Agar-agar, 605
Acrasiales, 8 Agars, various, 606, 611
Acrasis granulata, 8 Air content of tissues and disease, 280
Actinomyces bovis, 39 Albinism, 343
Actinomyces chromogenes, 39, 266, 544 Albumen of egg, 603
Actinomyces myricarum, 39 Alcoholic fermentation 59; in yeasts, 138
Actinomycetaceae, 39 Alfalfa, leaf spot of, 476, 477; leaf rust,
Activators, 57 477
Adenopetaly, 332 Algae in lichens 78; parastic, 391
Adesmy, 332 Alteration of position, 347
Adherence, 332 Alternation of generations in rusts,
Adhesion, 332 diagram of, 194
^cidium, 188 Alternaria citri, 533; dianthi on carna-
^ciospores, 188 tions, 488, 489, "490; violae, 558;
JEduia, 188 figure of, 559
Aerobic cultivation, 625 Alternariose of carnation, 488, 489, 490;
Aerobic organisms, 27 figure of, 489
/^thalium, 13 Amanita caesarea, description of, 734,
Agalinis as root parasite, 299 735, muscaria, description
735, of,
Agaricaceae, characters of family, 231, 736; at edge of woods, 83; figure of,
232 233; phalloidea, description of, 736;
Agaricaceae, Key to, 732, 733, 734 figure of, 238; in woods, 83.
753
754
Amanitopsis vaginata, speed of spore Appel's potato scab, 646

fall, 64 Appressoria, 308
Aniaurochaete, spores of, 16 Arcyria, 15
American Phytopathological Society, Armillaria mellea, 62, 83, 530; color of,
status of, 411 S3; described, 46, 737; figure of, 47
Amidase, 58 Arrestment of cell wall development, 359
Amcebobacter, 39 Arthrospores in bacteria, 25
Amphibolips ilicifolia, gall producing on Artificialwounds, 295
Quercus nana, 399 Asci of chestnut blight, figure of, 500
Amphispores, 188 Ascobolacese, characters of, 166
Amphitrichous, 23 Ascobolus immersus, special methods of
Amygdalin, 59 spore discharge, 66
Amylase, 58 Ascobolus, spore colors of, 54
Anaeretic, 332 Ascochyta pisi, 534
Anaerobic cultivation, 625; organisms, 27 Ascogenous hyphal system, figures of,
Analysis of water, 626 125, 127

Anatomy, pathologic plant, 354 Ascoideaceae, 120
Anbury, 487 Ascomycetales, bibliography of, 174,
Ancyclistaceae, characters of, 118 17s. 176; general characters of, 121,
Animals as cause of disease, 275 122; phylogeny of, 173, 174; sexuality
Animal galls, 296; injuries, 295, 309 of, 122
Animate agents of disease, 295 Ascospores 50; germination in chestnut
Annulus superus, 233 blight, figure of, 5^1; representation
Anther smuts, 72 by figures of development, 128
Antherophylly, 332 Ascus 50; diagrams of, by Claussen, 124
Anthesmolysis, 332 Ash, heart rot of, 481, 483
Anthocyanin, 360 Ashlock, J. L., quoted, 182
Antholysis, 329, 332 Ash of fungi, analysis of, 54
Anthracnose of cotton 508; of melons, Asiatic cholera, 37
52s; of raspberry, 544 Asparagus rust, 191, 483, 484

Anthrax, 35 Aspergillacese, characters of, 143
Anthurus borealis, 252 Aspergillus, characters of the genus, 144
Anti-enzymes, 58 Aspergillus fumigatus as pathogenic,
Antisepsis, 692 147; flavus, 147; giganteus, 147;
Aphylly, 332 Key to species of, 702-703; nidulans,
Apilary, 332 figure of, 148; niger with lipase, 59;
Aplanobacter, 35 with raffinase-58; luchuensis, 147;
Apogamy, 332 oryzete, 146; figure of, 145; with
Apophysis, 332 diastase 58; tokelau, 147; Wentii, 146
Apostasis, 333 Asphyxiation of roots, 565
Apothecium, structure of, 121 Assimilation tissues of galls, 400
Apple, black-rot of, 478, 479; bitter-rot Astrffius, 244
of, 477, 478; fruit spots 570; scab, Atkinson, Geo. F., book quoted, 91;
478, 480, 481; figures of, 480; tumor quoted, 235, 236; work of, 248, 249
on stem, figure of, 390 Atrichous, 23
Appel, O., work of, 272, 273 Atrophy, 333, 342
755
Auerbach's stain, 591 ingianus, fermentation by, 320; leprae

Auriculariaceae, characters of family, 216 35; Pasteurianus, fermentation by,
Auricularia Auricula- Judae, 216 32; mallei, 35; michiganense,
35;
Autodigestion of Coprinus comatus, 65; pestis, 35; phosphoreum, 36; pneu-
of fungi, 54 moniae, 35; Rathayi, 35; tuberculosis,
Autophyllogeny, 33;^ 35; vermiforme in ginger beer, 140
Awamori, a beverage, 147 Balance, organic, ^5^
Balanophoraceae, parasites of, 299
B Banana bud-rot, 484
Bark-boring beetles, 294
Bacillus amylobocter, 36; spores in, 25; Basidiobolus ranarum on frog drug, 85
amylovorus 36, 536, 644; aroideae, Basidiolichenes, 81
36; Biitschli, spores in, 25; butyricus, Basidiomycetales, characters of, 177;
36; calf actor, 36; carotovorus, 36; Key to suborders, 177
caucasicus in Kefir, 141; coli, 36; Basidiospores, 49, 187
inflatus, spores in, 25; influenzae, Basidium, 187
length and breadth of, 22; lathyri, Bastard toad-flax, 298
547; loxosporus, spores in, 25; Beam of light method of studying spore
loxosus, spores in, 25; megatherium, discharge, 64
nuclear material in, 24; mesentericus Bean mosaic, 577
vulgatus as a milk curdler, 59; Beefsteak fungus, 230
musae, 36, 484; nitri, length and Beet leaf-spot, 484, 485
breadth of, 22; nuclear material in, Beet rust, 485
24; phytophthorus 313, 646; prodi- Beetles, bark boring, 294
giosus, 36; and high temperatures, 360; Beggiatoa, 38; alba, 38; length and
putrificus, 36; radicicola, 29, 36, 612; breadth of, 22; mirabilis, 38; length
involution forms, 30; sub tills, 36; and breadth of, 22
rapidity of cell division, 24; spores in, Beggiatoaceae, 38
25; tetani, 36; tracheiphilus, 36, 313, Benecke, W., mentioned, 54
525 Benzaldehyde, 59
Bacteria, fermentation, 32; as disease Biastrepsis, 333
producers, 275; bibliography, 40; Bibliography of Ascomycetales, 174,
characterization, 638; classification of, i75> 176; of bacteria, 40; of disease
28; in general, 21; kinds of spores in, prevention, 318; of galls, 401, 402;
25; of root tubercles, figures of, 31; of non-parasitic diseases, 580; of
systematic account, 34 Oomycetales, 118, 119; of plant
Bacteriaceae, 35 diseases in general, 353; of rusts, 214,
Bacteriology emphasized, 271; systema- 215, 216; of slime moulds, 18, 19, 20;
tic, 630, 631 of smuts, 185, 186; of works on plant
Bacteriopurpurin, 38 diseases, 412; of Zygomycetales, 105
Bacterium, 35; aceticum, 36; fermenta- Biciliate zoospores, escape of, 67
tion by, 32; acidi-lactici, 36; fermen- Binucleate hyphal cells of Gasteromy-
tation by, 32; in Matzoon, 141; cetes, 218; of Hymenomycetes, 218
anthracis, 35; campestris, 485, 486, Biochemic features of bacteria, 636
487; diptheridis, 35; gammari, nuclear Biting insects, 296
material in, 24; influenzae, 35; Kiitz- Bitter-pit of apples, 570
47
756 INDEX
Bitter-rot of apple, 477, 478 Burt, E. A., work of, 248

Birds as spore carriers, 67 Butyric fermentation, 33, 59
Black ball, 178
Black Death, 35
Blackman, O. H., work on rusts, 191
Black-knot of plum 74, 540 Cabbage black-rot, 485, 486, 487
Black-rot of apple, 478, 479; of cabbage Cabbage leaf, figure of hypertrophied
485, 486, 487; of cruciferous plants, mesophyll, 368
experiments with, 645; of grape, 512; Cacao brown-rot, 490
figures of, S13, 514; of orange, 533, Cacao pink disease, 490
of sweet potato, 548 Casoma, 188; nitens, binucleate secio-
Black-rust of wheat, 560 spores of, 196
Blakeslee, A. F.,work on moulds, 93 Calciphile plants, 277
Blanched plants, 277 Calciphobe plants, 277
Blastomany, ^^^ Calcium, influence of, 277
Blight of chestnut 491; of sycamore, 549 Calcium oxalate in sporangial walls of
Blister-rust of white pine, 537; figure of, Mucor mucedo, 53
538 Calendar for spraying, 680-690
Blood serum, 604 Calico, description of, 327
Boletoideae, 234 Callous formation, conditions of, 380,
Boletus, change of color in, 53; felleus, 381; experiments with, 648; hyper-
230; figure of, 228; manual of, 227 trophies, 368, 369
Books on chlorosis, 328; on economic Callus, 377 et seq.; definition of, 377;
entomology, 296 histology of, 379
Bordeaux mixture, figure of apparatus Calvatia cyathiformis, figure of, 242
for making, 672; formulae for, 670-674 Calvatia, species of, 242
Botrytis cinerea, chitin in sclerotia of, 52 Calycanthemy, 333
Bouillon, 601 Calyphemy, 333
Bourquelat mentioned, 53 Calyptospora species of, 199
Breeding for disease resistance, 325 Cancer in plants, 34
Brefeld, Dr. O., cited, 89 Cancer-root, figure of, 301
Bronzing, 282 Canker lesion of chestnut, figure of, 492
Broom-rape as a parasite, 299; figure of, Canker of larch, 519
300 Cankers, 342, 348
Brown-rot of cacao, 490; of lemon, 520; Cantharellus aurantiacus, description
of turnip, figure of, 486 of, 739; cibarius, description of, 739,
Brown rust of rye, 202 740
Buchner discovery of zymase, 56 Capillitium, formation of, 13; in slime
Bud-rot of banana, 484 moulds, 15
Bulboceras gallicus and underground Carbohydrates, 58
truffles, 71 Carbol fuchsin, 589
Buller, A. H. F. book of, 233 Carbon circulation, 33
Bunt ear, 178 Carnation alternariose, 488, 489, 490;
Burgeff, H., work cited, 100 figure of, 489
Burl on oak trees, figure of, 350 Carrion fungi, development of, 248
Burrs, 348 Cassytha filiformis, 306
INDEX 757
Catalase, 58, 59 Chestnut blight, 491; distribution of,

Catalyst, 56 84; spread of, 316; gelatinous threads,
Cataplasms, 376, 385; histology of, 391 figure of, 494; perithecial pustules,
Cataplastic hypertrophy, 364 493
Catastome, 243, 244 Chestnut killed by blight, figure of, 313
Cavities covered with metal, 323 Chestnut leaf mildew, 502
Cavity treatment, 321 Chestnut, V. K., bulletin of, 238
Cecidial tissue forms, 397 et. seq. Chimaeras, 329, 330; periclinal, 330;
Cecidium, 384 sectorial, 330; spontaneous, 330
Cecidologists, 385 Chimney sweeper, 178
Cedar apple, figure of, 206, 394; on Chinese yeast, 99
small twig, figure of section of, 395 Chitin in bacterial cell wall, 22
Cedar rust on apple, 209 Chlamydobacteriaceae, 37
Celidiaceae, 169 Chlamydomucor racemosus, figure of,
Cell division in bacteria, 24 98, 99
Celloidin method, 655 Chlamydospores, 50; of corn smut, ger-
Celtis occidentalis, witches' broom on, mination of, 507; of smuts, 179; of
351 Tilletia foetans, figure of, 561
Cement cavity fillings, figure of, 322; Chlamydothrix, 37
mixing and placing, 321 Chloranthy, 37, 329, 333
Cenangiaceae, 169 Chlorophyljess plants, i
Cenanthy, S33 Chlorosis, 327, 343, 650; books on, 328

Cerastium viscosum, anther smut of, 72 Choanephoraceas, brief characterization
Ceratiomyxa, spores of, 16 of, 103
Ceratomany, 333 Chondromyces, 39
Cercospora beticola, 267; on beet, 484, Cholesterin, 56
485; coffeicola, 503 Cholin, 56
Cetraria islandica on ground, 83 Chorisis, 333
Chaetocladiaceê, characters of, 103 Christman, A. H., work on rusts, 191
Chaetocladium Jonesii, loi Chromatin in bacteria, 23; in fungi, 53
Chaetocladium parasitic on Mucor, 83 Chromatium, 39; Okeni, length and
Chaetomiacese, characters of, 163 breadth of, 22
Characterization of bacteria, 638 Chromogenic bacteria, 25, 26
Charles, Vera K., bulletin of, 244 Chromoparous, 26
Cheilomany, 333 Chromophorous, 26
Chemic character of soil cause of disease, Chromosomes in fungi, 53; reduction, 53
276 Chymosin, 59
Chemic elements in fungi, 54 Chytridiaceae, characters of, 116, 117,
Chemic work on fungi, 55 118
Chemistry emphasized, 271; of fungi, Circasa lutetiana, giant cells, figure of
52; of mushrooms, 237 372
Cliemomorphosis, 404 Cladochytricce, 116, 117
Chemotaxis, 60 Cladomany, 333
Chemotropism, 60 Cladonia cristatella on dead wood, 83;
Cherry leaf-curl, 491 pyxidata on stumps, 83; rangiferina
Cherry, powdery mildew of, 491 on ground, 83
758 INDEX
Cladothrix, 38; dichotoma, 38; fungi- CoUybia dryophila, fall of spores of, 64;
formis, 38; intestinalis, 38; intrica, 38; platyphylla on decaying logs, 74
profundus, 38; rufula, 38 Colonies, types of, 626, 627
Classification, i; of bacteria, 28; of Colors of bacteria, 26; in fungi, 53; of
enzymes, 58; of fungi, 2-6 Plasmodia, 12
Clathraceae, characters of family, 251; Columella in slime moulds, 15
distribution of genera and species of, Comandra umbellata, 298
87, 88 Comatricha nigra, figure of, 14; ob-
Clathrus cancellatus, figure of, 247; tusata, 13
columnatus, development of, 248 Conchs, 342
Claussen, P., reinvestigation of Pyro- Conidiophore, 46
nema confluens, 123; work cited, 108 Conidiospore, 46, 49
Clavaria, species of, 223 Coniferin, 56, 59
Clavariaceae, characters of family, 222 Coniothyrium Fuckelii, 262
Claviceps purpurea, 546; chitin in Conopholis americana, 299; figure of,
sclerotia of, 52; described, 162; fat 301; mexicana, 299

content, 56; figures of, 160, 161; Connold, Edward T., work of on galls,
sclerotia of, 69 384

Cleanliness to prevent disease, 367 Cook, Mel. T., work of, 274
Cleavage blocks in formation of spores Coprinus, deliquescence of, 53; descrip-
in slime moulds, 14 tion of genus, 748; atramentarius,
Climatic factors of disease, 281 749; figure of, 748; corpatus, 850;
Clostridium butyricum, 36 figure of, 749, 751; fed to Plasmo-
Clotting enzymes, 59 dium, 12; liberation of spores, 65;
Clouds, influence of, 284 number of spores in, 234; fimetarius,
Clover rust, 502 751; micaceus, 751; figure of, 750;
Club-root, 487, 488; figure of on cabbage stercorarius, 61; occurrence of, 83
roots, 488; of cabbage, figure of, 10 Coprophilous fungi and their spores, 68
Coagulation, 59 Cora, a lichen, 81
Cobb's disease of sugar cane, 37 Cordyceps Hiigelii, figure of, 70; mili-
Coccaceae, 34 taris, figure of, 70; on larvas of insects,
Coconut water, 599 69; ophioglossoides, figure of, 70;
Cocoon disease of silkworms, 147 parasitic on Elaphomyces, 69; sev-
Coelonemata, 15 eral sp'ecies described, 162; sphaero-
Coenobia, 21 cephala, figure of, 70
Coffee leaf-spot, 503 Coremium, 50
Coffee rust, 503 Coriolus versicolor, occurrence of, 229
Cohesion, 333 Cork as a protective layer, 308
Collection of fungi, 726, 727 Corn dry-rot, 504
Coleosporiaceae, characters of family, Corn smut on tassels, figure of, 506;
199 smut, 504, 505, 506; wilt, 507
Coleosporium solidaginis and sickness Correlation, 404
of horses, 200 Corticium lilaco-fuscum, 490; vagum-
CoUetotrichum gossypii, 508; lagena- solani, 221, 269
rium, 525; Lindemuthianum, 264; Cortinarius cinnamomeus, description
figures of, 265; species of, 266 of, 744; description of genus, 744;
INDEX 759
lilacinus, description of, 744; san- Cutting, calloused end of, figure of,
377
guineus, 744; violaceus, color of, 53; Cutting frozen material, 656
description of, 745 Cuttings of Populus pyramidalis, 379
Coryphylly, 333 Cyathus, 245
Cotton, 508; boll anthracnose, 508; Cyclochorisis, 334
rust, 508; wilt, 646 Cylindrosporium padi, 266
Cottony cushion scale, ravage of, 316 Cj'stobacter, 40
Counter, plate, 628 Cystopus condidus, 74
Counting methods, 620, 621 Cytase, 58 .
Counting plate, Jeffer's, 628 Cytinus hypocistus as a parasite, 301
Cover-glasses, squared, 616 Cytisus Adami, a graft hybrid, 329, 330
Cow wheat as a root parasite, 299 Cytology, emphasized, 271; of fleshy
Cowpea wilt, 646 fungi, 218; of rusts, 191
Cracks, frost, 294 Cytoplasm in bacteria, 23
Cranberry, 509; gall, 509; scald, 509; Cyttaria Berterii in Patagonia, 85;
detailed figures of, 510, 511 Darwinii in Patagonia, 85; Gunnii in
Crateria, 334 Tasmania, 85; Harioti in Terra del
Craterium leucocephalum, figure of, 17 Fuego, 85; in southern Patagonia,
Crenothrix, 38; polyspora, 38 74; on Nothofagus, 171
Cribraria argillacea, lead-colored Plas- Cyttariaceê, characters of, 171
modium of, 12; purpurea, scarlet
Plasmodium of, 12; violacea, violet D
Plasmodium of, 12
Cronartium ribicola, 313, 537; figure of, Dacryomycetaceae, characters of, 219
538 Dffidalea quercina, 558; absorption of
Crown-gall experiments with, 643; phosphorus by, 54; figure of, 558;
figure of an apple with, 352; nuclear occurrence of, 230
division, figure of, 373; on geranium, Damping-oflf, 342; distribution of fun-
figure of, 644; on raspberry, figure of, gus, 84
391 Danilov, work on lichens mentioned, 78
Crucibulum, 245, 246 Dasyscypha Willkommii, 519
Crustaceous lichens, 79 Death of hosts, 314
Cryptogamic parasites, 298 Decapitation experiments, 376
Cultivation of and fungi,
bacteria De Bary, Anton, work of, 189; men-
rough method, 587; of mushroom, tioned, 7
236, 237, 693 Decay, 33; of maple, 523; of oak, 526;
Cultural features of bacteria, descriptive of timber, 553
terms of, 633 Decoctions, plant, 600
Culture media, standardization of, 613 Dedoublement, 334
Cultures of de Vries, 328 Deformation, 334
Curdling, 59 Degeneration, 334
Curly-dwarf of potato, 576 Delafield's haematoxylin, 590
Curly- top of beets, 573 Deliquescence of Coprinus comatus, 65
Curricula and plant pathology, 410 Destruction of organs, 348
Cuscuta, description of, 305; figure of, Description of methods of bacterial
305 study, 631, 639
760 INDEX
Desiccation, 566 Disinfection, 692

Determining cause of disease, 274 Displacement, 335
Detailed account of specific plant Dissemination of fungi, 314, 315
diseases, 475 et seq. Distribution of slime moulds, 18
Deuteromycetes, 258-269 Distrophy, 335
Developmental mechanics of pathologic Dittschlag, work of, on rusts, 191
tissues, bibliography of, 405, 406, 407 Divulsion, 335
Development of carrion fungi, 248; of Dodder, figure of, 305; figure of section
fruit bodies in mushrooms, 235, 236 of, 306; study of, 651
De Vries, Hugo, work of, 331 Dodge, B. O., cited, 13, 15
Dextrose, 58 Dormant fungus in seeds, 308
Diachaena strumosa, 74 Dorrance, Frances, translations by, 413,
Diagram of rust spore relations, 190 564
Dialysis, 334; of enzymes, 57 Dothideaceae, characters of, 162
Diaphysis, 334 Downy mildew of grape, 513
Diastase, 58 Downy woodpecker and spores of
Dictydin granules, 15 Endothia parasitica, 67
Dictydium, 15 Drawing apparatus, 657
Dictyophora duplicata, figure of. 249; Drawing suggestions, 664-668
origin of veil, 249, 250; phalloidea, Drop of lettuce, 522
figure of, 250; figure of structure, 251 Dropsy, 352
Dictyostelium mucoroides, 8 Dry rot, 343; of corn, 504; of larch,
Dictyonema, a lichen, 81 519; of potato, 543
Didymium melanospermum, spore for- Dry-rot fungus, 225; in timber, 553
mation in, 13, 14 Duggar, B. M., book on mushroom
Die-back of citrus fruits, 572 growing, 237
Dilution methods, 616 Duration of disease, 313
Diplasy, 335 Dust brand, 178
Diplodia zeae, 504 Dwarfing, 342, 346
Diploid chromosomes in slime moulds, 16
Diremption, 335
Diruption, 335
Discentration, 335 Earth-star, 239, 244
Discharge of spores, 233, 234; figure of, Ecblastesis, 335, 338
63; in mushroom, figure of, 64 Ecology of fungi, 69
Discoloration, 342, 343 Economic entomology, field of, 296
Discomycetiineae, characters of, 164, 165 Ectotrophic mycorhiza, figure of, 49
Diseases, list of common plant, 414-473 Edinger's drawing apparatus, figure of
Diseases, non-parasitic, 564 details, 660, 661; description of, 657-
Diseases of plants, bibliography of speci- 664
fic, 473-474 Egg albumen, 603; yolk, 603
Diseases of plants in general, 271; two Egg plant wilt, 646
groups of, 413 Elaphomycetaceae, character of, 150
Diseases of sweet pea, 647 Elaphomyces, character of various spe-
Disease prevention, bibliography of, 318; cies, 150
resistance, 325 Elaters in slime moulds, 15
INDEX 761
Eleagnus, 9 Epipedochorisis, 335

and fungi, 62
Electric arc Epidemics, 315
Elenkin work on lichens mentioned, 78 Epiphytic mycelium, 48
Embryology emphasized, 271 Epiphytotisms, 298, 315
Empusa muscae, description of, 104; Epistrophy, 335
figure of, 104; as fly cholera, 85 Ergotin, 56
Emulsin, 58, 59 Ergot of rye, 546
Enation, 335 Eriksson's mycoplasm, 190
Enerthenema papillatum, figure of, 14 Erysiphaceae, characters of family, 154;
Endocellular enzymes, 56 Key to genera of, 721, 722
Endomycetaceae, characters of, 131 Erysiple, Key to species of, 723
Endomyces decipiens, parasitic on Armil- Escape of swarm spores, 67
laria mellea, 131 Esterases, 59
Endophyllaceae, characters of, 198 Ether freezing attachment, figure of, 659
Endophyiium sempervivi, described, 196, Etiolated, 335; plants, 277; plants
198; on house leek, 348 hypertrophied, 366
Endophytic mycelium, 48 Etiolation, 360; experiments with, 652
Endospores in bacteria, 25 Etiology, 272; description of, 641, of
Endothia parasitica, 491; and downy galls, 385
woodpecker, 67; description of, 164; Eubacteriales, 34
distribution of, 83, 84; figure of peri- Eubasidii, 218; bibliography of, 252-257
thecial pustules, 493, 495 ; mycelium of, Eumycetes, i, 42, 45, 46
496; spread of, 316 Euphrasia as a root parasite, 299
Engelmann experiment with bacteria Exanthema of citrus fruits, 572
and oxygen, 27 Excrescences, 342, 348; of bark, 366
Engler cited, 2 Excursions suggested, 667
Enteridium splendens, pink Plasmo- Exoascus and witches' brooms, 72; de-
dium of, 12 scription of species, 133, 134; figures
Entomology emphasized, 271 of, 132
Entomophthoraceae, characters of, 103 Exoascus cerasi, 491; deformans, 534;
Entomosporium maculatum, 264 pruni, 74, 541
Entyloma, description of several species, Exoascaceae, characters of, 131
185 Exobasidiaceas, characters of, 220
Enumeration of means of fungous Exobasidium, distribution of species and
entry into plants, 312 their hosts, 86, 87; vaccinii, figure of,
Enzymes, 56; and heat 57; and liquid 220; various species described, 220
air, 57; and plant diseases, 326; carbo- Expansivity, 335
hydrate splitting, 58; classification of, Explosions of smut, 182
58; clotting, 59; definition of word, Extracellular enzymes, 56
56; detection of, 59; diseases, 650; Exudations, 343, 350
distribution in fungi, 58; fat splitting, Eyebright as root parasite, 299
59; fermenting, 59; glucoside split- Eyepiece micrometer, 582
ting, 59; oxidizing, 59; protein-split-
ting, 59; solubility of, 57; urea-split-
ting, 59 Fabre, J. H., cited, 71
Epanody, 335 Facultative parasite, 42; saprophyte, 42
762 INDEX
Fairy ring, figure of, 75, 735; fungus, 74; Free, E. E., work of, 407
toadstool, 735 Freezing attachment for microtome,
Fasciation, 329, 335 figure of, 657
Fats in fungi, 53, 56 Freezing material, 656, 657; micro-
Fat-splitting enzymes, 59 tome, figure of, 658
Faull, J., work of, 172 Fries mentioned, 7
Fermenting power of yeasts, 595; en- Frondescence, 336
zymes, 59 Frost cracks, 294; influence of, 283;
Ferments, 56 necrosis of, 569
Fermentation, acetic acid, 32; alcoholic Fruit-pit of apples, 570
in yeasts, 138; butyric, 32, 59; Fruit-rot of orange, 533
by bacteria, 32; by mould, 96; in Fruticose lichens, 79
yeasts, 137; in fungi, 307; lactic acid, Fuhrmann, F., cited, 22, 24
32, 59 Fuligo septica, as flowers of tan, 17;

Ferrobacteria, 28 yellow Plasmodium, 12
Field of economic entomology, 296 Fuligo, spore formation in, 14
Field trip suggestions, 667 Fungi as cause of disease, 306, 307; as
Figure of rod-shaped bacteria, 22 disease producers, 275
Filar micrometer; figure of, 583 Fungicides, definition of terms, 669
Film formation in yeasts, 137 Fungi imperfecti, characters of, 258-
Final outcome of disease, 314 269
Fingers and toes, 487 Fusarium batatatis, figure of, 267;
Fink, Bruce, quoted, 78 heterosporium, 61; hyperoxysporum,
Fire-blight of pear, 536 figure of, 267; lycopersici, 646;
Fission, 336 putrefaciens, infection by, 273; species
Fistulina hepatica, 230; on tree trunks, of, 267, 269; trichothecoides, 543,
83 643; violae, figure of, 268
_
Fistulinoideas, 230
Fixatives, 655
Flagella of slime moulds, 16
Flecks of pith, 294 Galactose, 58
Fleshy fungi, 218 et seq. Gallionella, 37
Flies and spore distribution, 67 Galls, 342, 348, 384 et seq.; aeration
Flowers of tan, 17 tissues, 400; animal, 296; and insect
Flowering plants as cause of disease, 275 producers, 396; assimilation tissues of,
P'luckiger mentioned, 56 400; bibliography of, 401, 402;
Fogs, influence of, 284 cataplasmic, 385; formation, 72; his-
Foliose lichens, 79 tology of, 396, 397; hyperplasia, 384;
Fomes applanatus, 313; fomentarius, hypertrophy, 370, 371, 384; mechanic
523; figure of, 229; fraxinophilus, tissue of, 398; nutritive tissue of,
figures of, 481, 482; on beech, figure 398; of cranberry, 509; protective
of, 524; igniarius, figure of, 554; tissues, 398; secretory reservoirs of,
of rot by, 555, 556 400; vascular tissues of, 400

Forms of rust life cycles, 189 Gamomery, 336
Fossil fungi, 82 Gangrene, 343, 352
Fraser, Miss H. C, work on rusts, 191 Gas injuries, 649
763
Gases,efifect of, 289 Gray mould, figure of, 42

Gasteromycetes, character of, 239, 240 Green mould described, 45
Geaster, 239, 244 Griffiths, David, work cited, 163
Geaster fornicatus, figure of, 243; Grove, W. B., book of, 189
hygrometricus in sandy soil, 83 Guenther mentioned, 54
Gelatin, nutrient, 604 Guignardia Bidwellii, 512; of grape, 163;
Gelatin, sugar, 604 vaccinii, 509; figure of, 510
Gelatinous threads of chestnut blight, Guillermond, M. A., work cited, 142
figure of,494 Gummosis, 343, 350
Gemmiparity, 336 Guttulina rosea, 8
Genera of smuts, 182 Guying, 323 •
Genetics emphasized, 271 nature of, 271

; Gymnaxony, 336
Gentian violet, Ehrlich's anilin-water, Gymnoascaceae, characters of, 143
589 Gymnoconia interstitialis, figure of, 202
Geoglossaceae, description of, 169 Gymnosporangium biseptatum, figure
Geoglossum glutinosum, 170; hirsutum, of swelling caused by, 347; clavariae-
169; figure of, 170; range of, 85 forrrte, mycocecidia of, 393; Ellisii
Geographic distribution of fungi, 82 73, 210; figure of, 205; globosum, 21c;
Gerardia as a root parasite, 299 Gymnosporangium juniperi-virginianae,
Germination of smut chlamydospores, 74, 2 1 1-2 14; mycocecidia, 393; species
507; of smut spores, 181; of spores, of, 208-210
61; of spores and bacteria, 25; of Gynophylly, 336
spores of slime moulds, 16 Gypsum blocks and yeast spores, 622
Germination studies, 615 Gyromitra, 171
Gerry, Eloise, work on tyloses, 370
Giant cells, 371
Gilg cited, 2 Hackberry, witches' broom, 73, 351
Gilson, research of, 52 Haas, Paul, book of, 57
Ginger beer, 140 Haeckel, Ernst, cited, 7

Girdling of trees, 295 Haematimeter, Thoma's, 617; details
Glanders, 35 of, 619
Gloeosporium venetum, 544 Hail and plants, 286
Glomerella cingulata, 477, 478; gossypii, Hailstones, bruises by, 294
508; rufomaculans, 264, 477, 478 Hanbury, 487
Glucose, 53, 59 Hanging-drop preparations, 587
Glucoside-splitting enzyme, 59 Hansen, E. Chr., work of mentioned, 32,
Glycine hispida, figure of nodules on 141
roots, 29 Haploid chromosomes in slime moulds,
Glycocol, S3, 16
Glycogen in fungi, 53 Happy white elm, figure of, 286
Gnomonia veneta, 163, 264, 549; on Hard pan, influence of, 281
plane, 85 Harper, R. A., cited,
13, 15, 112, 165
Graft hj^brids, 329 Harshberger, John W., observations on
Grape, 512 acid spotting of morning-glories, 293;
Graphis scripta on bark, 83 work of, 308; on pine-barrens, 281; on
Gram's stain, 590 white cedar fungi, 394
764
Haustoria, 48; of Erysiphaceaj, 155 Hydnoraceae, parasites of, 299

Hay bacillus, 36 Hydnum erinaceus, figure of, 224, 556
Heald, F. D., work of, 342 Hydnum, species of, 223
Heart-rot of ash, 481, 483; of hemlock, Hydrocyanic acid, 59
517 Hymenium, 232

Heat as factor in plant disease, 282 Hymenogastraceae, character of family,
Helotiacese, characters of, 168 240
Helvellaceas, characters of, 170 Hymenomycetes, characters of, 219
Helvella crispa, 171; esculenta, fat con- Hyperchimaeras, 330
tent, 56 Hyperhydric tissues, 366
Helvelliineae, a suborder, 169 Hyperplasia, 355, 373
Hemibasidii, 178 Hypertrophy, 337, 347, 354, 364 et seq.;
Hemileia vastatrix, 503 kinds of, 364
Hemlock, 517 Hyphas, 42
Hemisyncotylous races, 329 Hyphomycetales, characters of, 266
Hemitery, 336 Hypochnaceae, characters of, 220
Hemitrichia vesparum, plasmodium of, Hypocreaceae, characters of, 160
12 Hypomyces, range of species of, 85;
Hepatica triloba attacked by Tranzs- lactifluorum parasitic on Lactarius, 1 60
chelia punctate, 348; figure of, 349 Hypoplasia, 354, 357; and cell contents,
Hepburn's definition of enzyme, 56 359, and tissue differentiation, 360;
Herbivores and spore distribution, 66 number of cells, 357; size of cells, 358
Heterodera radicicola, 391, 651 Hypothallus, 13
Heterogamy, 336 Hypoxylon, 164
Heteromorphy, 336 Hysteriaceae, characters of, 165
Heteroplasia, 374, 375
Heteroplasm, correlation, 376
Heterotaxy, 336
Heterothallic moulds, 93 Ice action, 295
Hill, T. G., book of, 57 Ice fringes, their formation, 283, 284
Histology emphasized, 271 Ice load of, figured, 285
Histology of callus, 379; of cataplasms, Ice storm and trees, 284, 285, 286
391; of fungi, 52; of galls, 396, 397 Iceland moss on ground, 83
Histozyme, 58 Icterus, 343
Hollyhock, 517; rust, 203, 206, 517; Idiotery, 337
518
figure of, Illuminating gas, effect of, 291, 292
Homooplasia, 374, 375 Immunity, 272, 325; to plant disease,
Homothallic moulds, 93 274
Homotypy, 336 Impregnation of wood with preserva-
Horses, injury by, 310 tives, 692
Host list of oomycetous fungi, 115 Indol, 33
Humphrey, C. T., mentioned, 75 Incubation, 312
Humus, influence on plants of, 281 Incubator, copper, 612
Hydnaceae, characters of family, 223 Infection by fungi, 307
Hydnocystis arenaria and black beetle, Infusions of plants, 600

71 Injured tree, figure of, 309
765
Injuries by acid, 649; by gas, 649; by fleshy fungi, list of, 729-732; to genera
smoke, 649 of family Exoascaceae, 133; to genera
Inoculation experiments, 643 et seq. of Peronosporaceae, 114; to Myxogas-
Inorganic elements in fungi, 55 trales, 693-695; to Nidulariaceae,
Insecticides, 678, 679 244, 245; to species of Penicillium
Insects as cause of disease, 275; as gall on agar and gelatin, 712-719; to
producers, 396; biting, 296; sucking, suborders of Basidiomycetales, 177
296; wood-boring, 310 Kiln-drying, 693
Intercellular hyphae, 48 Kinase, 57
Internal causes of disease, 326 Kinds of lichen thalli, 79
Intucellular hyphae, 48 Koernicke, Max, experiments with
Intumescences, 366 Roentgen rays, 62
Inulin, 58 Kohlhernie, 487
Inulase, 58 Koji fungus, 58
Invertase, 58 Kolkwitz, experiments of, 62
Involution forms of Bacillus radicicola, Knauers, 348
30; of bacteria, 364; of Pseudomonas Knife punch, figure of, 597
tumefaciens, 365 Knot of citrus trees, experiments with,
Iron indispensable to fungi, 54; in- 647
fluence of, 277 Kuehneola gossypii, 508
Irpex, species of, 223, 224 Kiihne, mentioned with enzymes, 56
Isolation of fungi in pure culture, 624 Kurssanow, work on rusts, 191
Ithyphallus impudicus, 252; and flies, 67
J
Laboratory exercises, 581; with slime
Jahn, E., cited, 13 moulds, 18
Jeffer's counting plate, 628; figure of, 629 Laboulbeniaceae, 172; hosts of, 86;
Jew's ear fungus, 216 work of Faull on, 173
Jones, L. R., work on cabbage immunity, Laboulbeniineae, 171
274 Labyrinthula Cienkowskii, 11
Juniperus virginiana, cedar apples 'on, Lachnea description of several species,
394 169; scutellata, figure of, 166

Lactarius, 731; chelidonium, description
K of, 740; deceptivus, description of,
740; deliciosus, description of, 740;

Kapoustnaja kila, 487 description of genus, 740; fumosus,
Karyokinesis, in fungi, 53 description of, 741; piperatus, de-
Kephir, 58, 140 scription of, 741; indigo, description
Kerner, Anton, work of, 385 of, 741; volemus, description of, 742
Key to determine species of Mucor, Lactic acid fermentation, 32, 59
695-702 Lactase, 58
Keys to Erysiphaceas, mentioned, 157 Lactose, 58, 59
Key to families of Oomycetales, 109; Lamium orvala, figure of, callus, 378
to families of Perisporiineae, 154; to Lamprocystis, 39
families of Zygomycetales, 97; to Lantz, Cyrus W., bibliography by, 564
766 INDEX
Larch, 519; canker, 168, 519; dry-rot, Lilac, 522; mildew, 522
519 Lime-sulphur, 675-677

Late-blight of potato, 542 Linaria vulgaris, peloria of, 329
Lathraea squamaria as a root parasite, Lindau, G., mentioned, 171
299 Lipase, 58, 59
Lathrop, Elbert C, worii of, 33 Liquid nutrient solutions, 592-595
Laticiferous hyphae, 48 List of common plant diseases, 414-473;
Laudatea, a lichen, 81 of keys to fleshy fungi, 729-732
Leaf-blotch of maple, 523 Lister, A., work of mentioned, 18
Leaf-casting, 575 Literature of plant diseases, exercises
Leaf-curl of cherry, 491; of peach, 534 in compiling, 642;on tree surgery, 324
Leaf-mildew of chestnut, 502 Litmus milk, 600; whey, 600
Leaf-spot of alfalfa, 476, 477; of apple, Living organisms as cause of disease, 275
figure of, 344; of beet, 484, 485; of Locomotion of bacteria, 23
coffee, 503; of violet, figure of, 559 Lohden wedge, 379
Leaves, skeleton, 294 Long, W. H., mentioned, 75
Leguminous tubercles, 387 Lophotrichous, 23
Leocarpus fragilis, figure of, 17 Loranthaceae, parasites of, 301
Leotia chlorocephala, 170; lubrica, 170 Lotsy, P., work on sexuality of As-
Lepidophyton, 147 comycetales, 122
Lepidosaphes ulmi, figure of, 276 Luminosity of fungi, 62
Leptothrix, 37; ochracea, 29 Lumpjaw of cattle, 39
Lepyrophylly, 337 Lupinus angustifolius, figure of cross-
Lemon, 520 section of tubercle, 30, 387, 388
Lenticels, hypertrophied, 366 Lycogala epidendrum, plasmodium of, 17
Lenzites betulina, 64; occurrence of, Lycoperdaceae, character of family, 241
230; sepiaria and rotting of slash, 75 Lycoperdon, species of, 241, 242
Lettuce, 522; drop, 522; experiments
with, 644 M
Leucin, 33
Leuconostoc mesenterioides, 34 MacBride, Thomas H., work of men-
Levulose, 58, 59 tioned, 18
Liberation of spores, 62; in Coprinus MacDougal, D. T., experiments with
comatus, 65 fungi in dark, 61
Lichen thalli, 79; algae, 78; as fungi, 79; Macrodactylis subspinosus, figure of, 275
parasitism of fungi, 79; nature of, 78; Macrosporium solani, 266, 267
structure of thallus, 81 Magnesium, influence of, 277
Life cycle of Oomycetales, diagram of, Magnification values, tables of, 663
108; of Pyronema contrasted with Maladie digitorie, 487
fern, 126 Malaria, 18
Life histories, description of, 641 Malformations, 329, 342, 348
Light and pathologic conditions, ex- Malpighi, 385
periments, 652 Maltase, 58
Light and red pigment, 360; influence Maltose, 58
of, 61, 281; action of, 288, 289 Manihot, oedema of, 567
Lightning, injury by, 311 Mannite, 53
INDEX 767
Mannose, 58, 59 Metatrophic bacteria, 31; organisms,- 28

Manual of American boletes, 227; of Meteorologic factors of disease, 281
polypores, 227 Methods of teaching, 407-410
Maple, 523; decay, 523; leaf-blotch, 523 Methylene blue, alkaline, 589
Map of chestnut blight fungus, 84 Meyer, mentioned, 54
Marasmium oreades, 74; figures of, 75, Micrococcus, 34; aurantiacus, 34; cinna-
735 bareus, 34; gonorrhoeae, 34; luteus,
Massee, George A., book of, 91; men- 34; progrediens, diameter of, 21, 22;
tioned, 169 pyogenes aureus, 34; ureae, diameter
Masters, Maxwell T., 331, 340 of, 22; with urease, 59
Matzoon, 141 Micrometer, eyepiece, 582; filar, 583;
Mazum, 141 figure of, 583; stage, 582; step, 585;
McAlpine, D., work of, 570, 571 figure of, 586; tables of values, 584,
Mechanic development of pathologic 585
tissues, 403, 404, 405 Micrometry, 582
Mechanic injury, 294 Microsphaera alni, 522; figure of, 157;
Mechanic tissue in galls, 398 key to species of, 724, 725
Mechanics of pathologic tissues, bibliog- Microspira, 37
raphy of, 405, 406 407 j
Microtome, figure of sliding, 654; with
Meiophylly, 337 freezing attachment, figure of, 658
Meiotaxy, 337 Microthyriaceae, characters of family,
Melampsoraceae, characters of family, 158
198 Mildew of grape, figure of, 515
Melampsora, species of, 199 Milk, 600; litmus, 600
Melampsoropsis, species of, 199 Mischomany, 337
Melampyrum as a root parasite, 299 Miso sauce, 146
Melanconiales, characters of, 264 Mistletoe diagram of habit, 304; figure
Meliola camelliae, 54; distribution of, of, 302, 303; references to literature,
85; Penzigi, 521 304; study of, 651
Melitiose, 58 Mites, 296
Melogrammataceae, characters of, 164 Mixing of cement, 321
Melon anthracnose, 525; wilt, 525 Miyoshi, M., experiments with chemo-
Merulius lacrymans, 553; description of, taxis, 60
224, 225, 226; figure of, 225, 226 Mnium hornum and underground
Meruloideae, 224 truffles, 71
Meschinelli, L., work on fossil fungi, 82 Molisch, Hans, experiments of, 62;
Mesospores, 188 mentioned, 54
Mespilodaphne sassafras, section of old- Mollisiaceae, characters of, 169
wood, figure of, 369 Monoblepharidaceae, characters of, 109
Metal-covered cavities, 323 Monoblepharis sphaerica, structure of,
Metamorphosis, 337 109
Metaphery, 337 .
Monospora, 141
Metaplasia, 354, 362 et seq.; and cell Monosy, 337
contents, 362; and cell membranes, Monotrichous, 23
363 Monstrosities, 329
Metastasis, 337 Moore, Geo. T., work of, 31
768
Morchella, 170, 171; esculenta, analysis Mycorhiza, 49

of, 55 Mylitta australis, sclerotium of, 71
Morel, 170, 171 Myriangiaceas, 153
Morphology emphasized, 271; of bac- Myrica carolinensis, tubercles on roots,
teria, descriptive terms, 633; of 39
chestnut blight fungus, 497 Myxamoebae, 15
Mortierellaceae, characters of family, 103 Myxobacter, 40
Mortification of tissues, 346 Myxobacteriaceae, 21, 39
Mosaic diseases, 327 Myxococcus, 40
Mosaic of bean, 577; of tobacco, 578 Myxogastrales characters of, 11; key to,
Mottle-leaf, 573 693-695
Mould fungi, 92; sexual reproduction, 93 Myxogastres, 7
Mounting bacteria, 588 Myxomycetes i, 7
Movement of Plasmodium, 12
Mucoraceae, character of, 97
Mucor, figure of, 42; key to species, N
695-702; mucedo, chitin in, 52; de-
scribed, 45; figure of, 44; occurrence Nanism, 346
of, 83; sporangia of, 96; structure of, Nature of tree surgery, 320
98; racemosus, chitin in, 52; Rouxii as Necrosis, 342, 346; frost, of potato
Chinese yeast, 99; various species tubers, 569
of, 98 Nectria cinnabarina, description of, 160
Multinucleate cells, 372; giant cells, Nectria, figures of various species, 159
371; spores of Rhizopus nigricans, 96 Neisser's counting apparatus, 629
Multiplication, 337 Nematode infection, 651; worms as gall
Mummification, 342, 347 formers, 391
Miinch, E., experiments on water and Neocosmospora vasinfecta, 646
air content of tissues, 280 Neoepigenesis, 404
Murrill's arrangement of fleshy fungi, Neoevolution, 404
228 Nidularia, 244
Muscaria, 56, 238 Nidulariaceae, character of family, 244;
Mushrooms, 231; chemistry of, 237; key to, 244, 245
cultivation of, 236, 237, 693; develop- Nitric organism, isolation of, 611
ment, 235, 236; figures of, 234, 746; Nitrifying bacteria, 29
toxicology, 237, 238, 239 Nitrobacter, 29
Mutations, 328 Nitrogen cycle, ^y, deficiency, influence
Mutinus caninus, development of, 248, of, 278; fixation, 612; influence of, 278;
249; and flies, 67 source of in fungi, 55
MyceHum, 42; of Endothia parasitica, Nitrococcus, 29
figure of, 496 Nitrosomonas, 29; javanensis, 29
Mycetozoa, 7 Nodule-forming bacteria, 29
Mycocecidia, 393 Nodules of roots, 387
Mycodendron paradoxum, 226 Non-parasitic diseases, 564; bibliography
Mycoderma aceti, 59; nature of, 142 of, 580
Mycomycetes, 46, 120 Normal solutions, 613
Mycoplasm, 49; Eriksson's, 190 Nothofagus with Cyttaria, 74
769
Nuclear apparatus of yeasts, 135; divi- Orobanche, as a parasite, 299; minor,

sion in yeasts, 136; phenomena in figure of, 300
fleshy fungi, 218; in rusts, 192, 193; Organized ferments, 56

phenomena of fleshy fungi, students Orton,W. A., on quarantine, 317
of problems, 218, 219 Osmomorphosis, 404
Nuclease, 58, 59 Ostwald, mentioned, 57
Nuclei in fungous cells, 53 Oyster mushroom, 738
Nucleus in bacteria, 23 Oyster-shell scale, figure of, 276
Number of spores produced, 63 Oxidizing enzj'mes, 59
Nummularia BuUardi on beech branches,
164
Nutrient solutions, 592-595 Pachyma cocos, 72; malacense, sclero-
Nutrition of bacteria, classification ac- tium of, 72
cording to, 28 Pallor, 342
Nutritive disturbance as cause of disease, Panaschiering, 326
328; tissues in galls, 398 Parachromatophorous, 26
Nyctalis asterophora, parasitic, on Paraffin method, 656
Russula nigricans, figure of, 43 Paralyzers, 57
Parasite, 42; chlorophylless, 298; green,
298; on roots, 299
Parasitic algae, 391
Oak, 526; decay, 526; root-rot, 530 Parasitism of lichen fungi, 79
Oat, 531; rust, 531; crown rust of, 202 Paratrophic bacteria, 33; organisms, 28
Obligate parasite, 42; saprophyte, 42 Parmelia perlata, figure of, 80; on
Qidema, 352; of manihot, 567; figure of, trunks of trees, 83
568 Pasteurization, 625
Oenothera Lamarckiana, 328 Pasteur mentioned, 56
Oidiospores, 50 Patellariacese, 169
Oidium lactis in Matzoon, 141 Pathogenic fungi, study of, 639
Oils in fungi, 53, 56 Pathologic plant anatomy, 354; tissues,

Olive, Edgar W., work on rusts, 191; mechanic development of, 403, 404,
cited, 13 405
Onion, 531; smut, 531 Pathologist, character of work of, 341
Oochytrieae, 117 Pathology, special plant, 411 et seq.
Oolysis, 337 Patterson, Flora W., bulletin of, 244
Oomycetales, 43, 50; bibliography of, Pea, 534; pod-spot, 534
118, 119; characters of, 107; key to Peach leaf cure, 534; yellows, 315, 573
families, 109; motile cells in, 52; Pear, 536; blight, experiments with,
occurrence of, 108; sexual reproduc- 644, figure of experiment, 645
tion in, 107 Peloria, 329, 337
Oomycetous fungi, host list, 115 Peltigera canina on ground, 83
Oospora scabies, occurrence of, 83 Penicillium atramentosum, figure of,
Oospores, 50 713; bif orme, figure of , 71 6 brevicaule,

;
Orange, 533; black-rot, 533^ fruit-rot, description of, 709; figure of, 709;
533; juice, 598 Camemberti, description of, 706, 707;
Orobanchaceae, parasites of, 299 figure of, 706; chrysogenum, 711;
770
claviforme, figure of, 710; commune, Phacidiaceae, characters of, 165

figure of, 717; decumbens, figure of, Pholiota adiposa, figure of, 76;*on living
715; digitatum, figure of, 720; Du- trees, 74
clauxii, figure of, 711; expansum, Phallaceae, character of family, 252
figure of, 704; funiculosum, figure of, Phallin, 56, 238, 239
714; general characters of, 703; Phallomycetes, 246-252
glaucum, 61; chitin in, 52; described, Phanerogamic parasites, 298
45; figure of, 46; with lipase, 59; Phoma, species of, 262
key for
italicum, 533; figure of, 708; Phoradendron flavescens, as a parasite,
on various substrata, 719, 720;
species 303; figure 302
of,
key to species grown on agar and Phosphorescent fungi, 62
gelatin, 712, 719; lilacinum, figure of, Phosphorus, influence of, 278
713; purpurogenum, figure of, 719; Photogens, 25
Roqueforti, description of, 704; figure Photographic prints, drawings
of, 666
of, 705 roseum, figure
; of, 712; rubrum, Photomicrographic attachment to Edin-
figure 718; rugulosum, figure of,
of, ger's apparatus, figure of, 662
721; spinulosum, figure of, 718; Photomicrography, method of, 666
stoloniferum, description of, 708; Phototropism, 61
figure of, 707 Phragmidiothrix, 38; multisepta, 38
Penzig, O., work of, 331 Phragmidium violaceum, fusion of ad-
Pepsin, 59 joining cell nuclei, 191, 192
Periclinal chimjeras, 330 Phycobacteriaceae, 37
Peridermium, 188; species of, 201; Phycomyces nitens, 61; structure of, 100
strobi, 537 Phycomycetes, 45, 46, 50, 92
Periphyllogeny, 337 Phyllactinia corylei, figure of, 53
Perisporiaceas, characters of family, 158 Phylloclady, 337
Perisporiineae, characters of, 154; key Phyllody, 337
to families, 154 Phyllomania, 338
Perithecium, structure of, 121 Phyllosticta paviae, figure of, 259; soli-
Peritrichous, 23 taria, figure of section, 262; on apples,
Permutation, 337 figure of, 261; species of, 261 et seq.
Peronosporacea?, cellulose in, 52; charac- Phylloxera mentioned, 295; vastatrix,
ters of family, in; generic key, 114 391
Peroxidase, 59 Phylogeny of Ascomycetales, 173, 174;
Pestalozzia Guepini var. vaccinii, 266 of fungi, 89, 90, 01; of Uredinales, 197
Petalody, 329, 337 Physarum sinuosum, figure of, 17;
Petalomania, 337 ellipsoideum, plasmodium of, 12;
Petersen, Henning E., work of, in psittacinum, 13
Petri dish, figure of, 622 Physcia parietina on rocks, 53
Peyritsch, J., mentioned, 171 Physical character of soil as determining
Peziza aeruginosa, uses, 168; aurantiaca, cause of disease, 279
color of, 53; described, 167; badia, Physical features of bacteria, 636
occurrence of, 167; coccinea, color of, Physics emphasized, 271
S3, on dead twigs, 83 ; described, 67; Physiologic diseases, 564
Fuckeliana, 61; repanda, figure *of, Physiology emphasized, 271
167; Willkommii or larch canker,"i68 Physiology of fungi, 54, 61
771
Phytase, 58 Pleurotus, 'description of genus, 737;

Phytocecidia, 385 olearius, 62; ostreatus, description of,
Phytomyxa, 9; leguminosarum, 11 738; figure of, 738; serotinus, de-
Phytomyxales, 8 scription of, 738, 739; sapidus, de-
Phytopathological Society, American, scription of, 738; ulmarius, descrip-
411 tion of, 739
Phytopathology, 411; definition of, Plowrightia, description of several spe-
272 cies, 162; raorbosa, 74, 540; distribu-
Phytophthora infestans, 315, 542; es tion of, 84; figure of, 73
cape of zoospores, 67; infection by, 273 Plugging test-tubes, 586
Pichia, 141 Plum, black-knot of, 540; pockets, 74,541
Pilacraceae, characters of family, 217 Pod-spot of pea, 534
Pilobolus, figures of species, 102; crys- Podospheera, key to species, 722
tallinus and horses, 68; occurrence Poisoning by fungi, symptoms, 238, 239
of, 83 Poisonous substances in fungi, 238
Pineapple chlorosis, 650 Pollaplasy, 338
Pink disease of cacao, 490 Polyangium, 40
Piptocephalidacea;, characters of, 103 Polychrome methylene blue, 591
Piptocephalis parasitic on Mucor, 83 Polyclady, 338
Pistillody, 338 Polyphagus euglenae, occurrence of, 117
Pith flecks,
294 Polyphylly, 338
Placing of cement, 321 Polyporaceae, characters of family, 224
Plague, 35 Polypores, manual of, 227
Planococcus, 35; citreus, 35 Polyporoideas, characters of, 226
Planosarcina, 35 Polyporus borealis, 517; mylittae, sclero-
Plant juices, 598 tium of, 71; ofiicinalis, analysis of,
Plant pathology, growth of, 410; 55 ; ponderosus, 539; sapurema, sclero-
special, 411 et seq. tium of, 71; sulphureus, 526; figure
Plants as disease producers, 298 of fruit, 527; figure of decaying oak,

Plasmodiocarps, 13 528; on with trehalase, 58;
trees, 83;
Plasmodiophora, 9; alni, 9; brassicas, tuberaster, sclerotium of, 71
9, 387, 487, 488; on cabbage roots, Polysphondylium violaceum, 8
figure of, 488; figure of, 10; eleagni, 9 Polystictus abietinus and rotting of
Plasmodium, aggregate, 8; colors of, 12; slash, 75; socer, sclerotium of, 72;
malarias, 18; movement of, 12 versicolor, 64, 545
Plasmopara viticola, 513; distribution Poppy, fasciated, figure of, 336
of, 84; figure of, 113 Poplar cutting, figure of, 378
Plasmolysis, experiments with, 653 Populin, 59
Plate counter, 628 Populus pyramidalis, cuttings of, 379
Plectenchymatous, 258 Potassium hunger, 277
Plectasciinea?, characters of, 143 Potato, 542; as medium, 596; broth, 597;
Plectridium, 25 curly-dwarf of, 576; glycerinated, 596;
Pleiomorphy, 338 juice, 596; late-blight, 542; rot,
Pleiophylly, 329, 338 experiments with, 643; scab, 544
Pleiotaxy, 338 Pouring plates, figure of, 622; method
Plesiasmy, 338 of, 622, 623
48
772
Powdery dry-rot of potato, 543 188; forms; of, 191, 201, 202; malva-
Powdery mildew of cherry, 491; of cearum, 206, 517; figure of, 518;
lilac,522 species of, 203, 204, 205, 206
Predisposing causes of disease, 272 PufT-balls, 239, 240
Preservation of wood, 692; of fungi, 726, Puffing of spores, 66
727 Pumps for spraying, figures of, 691
Prevention of disease, bibliography of, Punks, 342
318 Pustules, 342
Preventive measures, 319 Putrefaction, ^3
Prints of spores, 728 Pycnidial pustules of chestnut blight,
Prod'uction of spores, 63 499
Prolification, 338 Pycnidiospores, 50
Projection apparatus, 657 Pycnidium, 50
Prophylaxis, 298, 317 Pycnium, 188
Prosoplasms, 376, 395 Pycnoconidia, 50
Prosoplastic hypertrophy, 364 Pycnospores, 50; 188; germination of
Protease, 58, 59 chestnut blight, figures of, 501
Protective tissues in galls, 398 Pyrenomycetiinese, characters of, 159,
Proteins, splitting of, ^$, 59 1 60
Protista, 7 Pyronemaceae, characters of, 165
Protoasciineae, 131 Pyronema, life cycle contrasted with
Protomyces, occurrence of species, 121 fern, 126; confluens and sexuality,
Protomycetacese, 121 165; reinvestigation of , by P. Claussen,
Protophyta, 7 123; work on by R. A. Harper, 122
Protoplasm of fungi, 53' Pythiacystis citriophora, 520; on lemon,
Prototrophic organisms, 28 85
Protozoa, 7 Pythium de Baryanum, distribution of,
Pruning careless, 310; unskillful, figure 84

of, 310
Pseudomonas, 31?, 36; brassicse, 485,
486, 487; campestris, 36, 645; europaeus
Quarantine to prevent disease, 317
37; hyacinthi, 36; indigofera, length
Quercus reticulata parasitized by Cono-
and breadth of, 22; putida, 37;
pholia mexicana, 299
pyocyanea, 37; Stewarti, 644; syn-
Quercus Wislizeni, figure of section of
cyanea, 37; tumefaciens, 34, 388, 643;
gall, 399; gall on, 398
involution forms of, 365; vascularum,
Quick-drying varieties of plants, 273
.36
Pseudopeziza medicaginis, 169; on

alfalfa, 476, 477
Ptomaines, 23 Races of moulds, 95
Pucciniaceae, characters of family, 201 Rachitism, 338
Puccinia asparagi, 483, 484; character Raffinase, 58
of, 191; coronifera, 531; forms of, Ralfinose, 58
202; forms of, 202, 203;
coronata, Rafflesiaceae, parasites of, 301
glumarum, forms of, 203; graminis, Raspberry anthracnose, 544
560; distribution of, 84; figure of, Rate of spore fall, 64
773
Razoumofskya Douglasii laricis as a Root-rot of oak, 530; of tobacco, 550;

parasite, 304 figure of, 551
Recrudescence, 338 Roquefort cheese, 704, 705
Red clover, figure of tubercle section, 389 Rose chafer, figure of, 275
Red gum, 545 Rosellinia quercina on oak seedlings, 163
Red-rot of pine, 539 Rosettes, 342
Red spider, 296 Rostafinski mentioned, 7
Reduction in size, 342 Rotation of crops to prevent disease, 317
Regeneration, 355 Rottenness, 352
Reindeer lichen on ground, 83 Rotten wood, 307
Rennin, 59 Rotting, 343, 352; of brush, 75
Replacement, 342, 347 Rozites gongylophora and the tugging-
Reproduction in bacteria, 24 ant, 365; as food for tropic ants, 71
Resin in fungi, 56 Ruppia rostellata, 11
Resinosis, 343, 350 Russula, 48; description of genus, 742;
Resin wash, 521 emetica, 742; in forest litter, 83;
Resistance to disease, 325 nigricans parasitized by Nyctalis
Restitution, 355, 356, 357; meaning of asterophora, figure of, 43, with tyro-
word, 355; process of, 355 sinase; ochrophylla, description of,
Reticularia lycoperdon withajthalium,i7 743; roseipes, description of, 743;
Reticularia, spores of, 16 rubra, description of, 743; virescens,
Retting of fibers, li:^ color of, 53; description of, 743; in
Reynolds, Ernest Shaw, mentioned, 271 forest litter, 83
Rhabdochromatium, 39 Rust fungi, 187; occurrence of, 86;
Rhizinaceae, 171 lesion on apple leaf, section of, 213;
Rhizobium leguminosarum, 29, 36 life cycles, forms of, 189; of alfalfa,
Rhizocallesy, 338 477; of asparagus, 191, 483, 484;
Rhizoctonia solani, 269 of beet, 485; of clover, 502; of coffee,
Rhizomorph, figure of, 47 503; of cotton, 508; of hollyhocks,
Rhizomorpha subterranea, figure of, 47 203, 517; of oat, 531
Rhizopus nigricans, chitin in, 52; Rust, spore relations, diagram of, 190
conjugation of, 94; figure of, 100; Rusts, bibliography of, 214, 215, 216;
occurrence of, 82; structure of, 99 cytology of, 191; life cycle, 195
R.hodobacteriaceae, 38 Rye, 546
Rhodomyces Kochii, 61
Rhytisma acerinum, 523; on maple, 165
Ribes aureum, figure of hypertrophied Saccharomyces anomalus, 40; aquifolii,
bark, 367 140; cartilaginosus in Kefir, 104;
Ringing of trees, 295 cerevisise, 52; description of, 138;
Roentgen rays and fungi, 62 figure of, 135; ellipsoideus, descrip-
Rcestelia, 188; aurantiaca, figure of, tion of, 139, 140; figure of, 139; of
204; on apple, diagram of, 212; on nuclei and division, 136; exiguus,
apple leaf, figure of, 210; on apple, 140; fragilis in Kefir, 140; ilicis, 140;
magnified view, 211 Ludwigii, 141; octosporus with mal-
Rodents and truffles, 68; injury by, 294 tase, 58; Pastorianus I, 140; pyri-
Root asphyxiation, 565; parasites, 299 formis, 150; Vordemanni, 140
774 INDEX
Saccharomycetaceae, characters of, 134 Scorias spongiosa, 158; life history of, 72
Saccharomycetiinese, 134 Scrophulariacese, parasites of family, 299
Saccharomycodes, 141 Scyphogeny, 338
Saccharomycopsis, 141 Sectioning methods, 633, 654
Sake, 146 Sectorial chimseras, 330
Salicin, 59 Sepalody, 338
Salmon, Ernest S., monograph of, 157 Septoria leaf-spot, figures of, 263;
Salpinganthy, 338 species of, 264
Sandalwood, parasitism of, 298 Sequoia gigantea, annual rings of, 358
Santalum album, parasitic on Acacia Serum of blood,

604
leucophsa, 298; on roots of Melia Sexual act in slime moulds, 16
azidarachta, 298 Sexual reproduction in Oomycetales,
Saponaria officinalis, anther smut of, 72 107; in Sphserotheca Castagnei, 155;
Saprogenic organism, 33 in moulds, 93; in Ascomycetales,
Saprogens, 25 bibliography of, 129, 130; of As-
Saprolegnia, 44; ferax on fishes no, in; comycetales, 122, 123
structure of various species, no; Shaggymane, figure of, 749
escape of zoospores, 67 Shot-holes, 342, 345; of plum leaves,
Saprolegniacec-e, cellulose inj 52; charac- figure of, 345
ter of family, no Silene inflata, anther smut of, 72
Saprophyte, 42 Silverberry, 9
Sap-rot of red gum, 545; of timber, 558 Size of bacterial cells, 21, 22
Sarcina, 35; aurantiaca, 35; fiava, 35; Skatol, î
lutea, 35; maxima, diameter of, 22; Skeleton leaves, 294
rosea, 35; ventriculi, 35 Slant of vegetables, figureof, 597
Sarcosphaera, figures of several species, Sleeping disease of tomatoes, 646

166 Sliding microtome, figure of, 654
Scab of apple, 479, 480, 481; figures of, Slime flux, 343
480; of potatoes, 544 Slime moulds, bibliography of, 18, 19,
Scald of cranberry, 509 20; distribution of, 18; laboratory
Scarification of trees, 295 exercises with; in general, 7
Schizomycetes, i; origin of name, 21 Smelter fumes, effect of, 289, 290, 291
Schizonema imbricator, a scale insect Smith, Erwin F., quoted on peach
and Scorias spongiosa, 72 yellows, 315; work of, 34, 387
Schizophyllum commune, 64; figure of, Smoke, effect of, 289, 649
77; xerophytic habits of, 78 Smut boil of corn, figure of, 504, 505, 506
Schizosaccharomyces, 141 Smut explosions, 182
Schmitz, J., mentioned, 61 Smut of oats, figures of, 532; of onion,
Sclerodermaceae, characters of family, S31; spores, germination of, 181
246 Smuts, 178-186; bibliography of, 185,
Scleroderma vulgare on old stumps, 83 186; genera of, 182; of anthers, 72;
Sclerotia, 69; fungi bearing, 71 modes of infection, 181
Sclerotinia libertiana, 522, 644; descrip- Snow action, 295; influence of, 284
tion of several species, 168; sclerotia Soft rot, 343
of, 69; figure of, 168 Soja sauce, 146
Sclerotium, 48 Solenoidy, 338
INDEX 775
Solid vegetable substance, 598 Spot disease of violet, 558

Solution 338; normal, 613 Spots, colored, 342
Soot, effect of, 289 Spray calendar, 680-690
Sooty mould of orange, 521 Spray pumps, figures of, 691
Sorauer, P., book of, 564 Spraying for plant protection, 318
Sordariaceae, characters of, 162, 163 Sprays, 669 et seq.
Sorosphaera, 9; veronicae, ii Spruce gum, collection of, 352
Soy bean, figure of nodules on roots, 29 Squared cover-glasses, 616; figures of,
Sparassis crispa, 223 617
Special plant pathology, 411 et seq. Squashes, 525
Speiranthy, 339 Stab cultures, types of, 627; in figure>
Spermogonium, 187 627
Sphaeria carpophila, 61 Stage micrometer, 582
Sphaeriaceae, characters of, 163 Stag-head, 395, 565
Sphasrobolaceae, characters of family, 246 Staining bacteria, 588
Sphasrochorisis, 339 Stains, 589-592
Sphajronema fimbriata, 548 Staminody, 339
Sphaeropsidales, 260 Standardization of culture media, 613
Sphaeropsis malorum, 262; figures of Stasimorphy, 339
spots due to, 344; on apple, 478, 479; Statement, general, i
tumefaciens, 647 Steeps, 677-678

Sphaerotheca Castagnei, sexual repro- Stemonitis ferruginea, spores of, 16;
duction in, 155; key to species of, 722 flaccida, spores of, 16; fusca, figure of,
Sphaerotilus, 38 14
Spirillaceae, 37 Step micrometer, 585; figure of, 586
Spirillum, 37; berolinense, 37; comma, Stereonemata, 15
37; danubicum, 37; parvum, thickness Stereum, 221, 222; fasciatum, and
of, 21; rufum, 37 rotting of slash, 75; frustulosum, 553;
Spirochaeta, 37; dentium, 37; Ober- rameale and rotting of slash, 75; ura-
meieri, 37; pallida, 37 brinum and rotting of slash, 75;
Spiroism, 339 versiforme and rotting of slash, 75
Spirosoma, 37 Sterigmatocystisniger, character of, 147;
Spontaneous chimaeras, 330 figure of, 149
Sporabola, 234 Sterilization, 625
Sporangiospores, 50 Stesomy, 339
Spore discharge in mushrooms, 233, Stevens, Neil E., mentioned, 84
234; figure of, 64 Stigmatomyces Baeri, structure of, 172
Spore fall in Amanitopsis vaginata, Stippen, 570
figure of, 65; rate of, 64 Strains of moulds, 95
Spore formation in moulds, 96; germina- Strangulation, 294
tion, 61; prints, 728; production, 63 Strasburger, Ed., cited, 15
Spores of yeasts, 622; of rusts, nuclear Streak cultures, t3T)es of in fungi, 634
phenomena in, 192 Streak method of Bergey, 623
Sporodinia grandis, conjugation of, 94; Streak of sweet pea, 547
occurrence of, loi Streptococcus, 34; erysipelatos, 34;
Sporulation in yeasts, 137 mesenterioides, 34; pyogenes, 34
776
Streptothrix, 37; fluitans, 37 Synspermy, 339

Strobilomyces strobilaceus, 230 Syphilis, 37
Strophomany, 339 Systematic account of bacteria, 34
Structure of lichen thallus, 81 Systematic bacteriology, 630, 631;
Stub, figure of, 311 botany emphasized, 271; position of
Students of nuclear phenomena in fungi imperfecti, 260
fleshy fungi, 218, 219
Students, suggestions to, 407
Sturgis, W. C, literature of plant Taka-diastase, 58, 146
diseases, 411 Tannin as a protective substance, 274
Stylospores, 50 Taphrina cserulescens on oaks, 85
Succulence, abnormal, 368 Taphrina, description of various species,
Sucrose, 58 134; of figures of, 132
Sucking insects, 565 Tas Gu of Java, 146
Suffocation, 565 Taubenhaus, J. J., work of, 274
Suffulcra of Erysiphaceae, 155 Taxitery, 339
Sugar beets, curly-topof, 573 Teachers, suggestions to, 407
Suggestions to teachers and students, Teaching methods, 407-410
407-410 Telegraph wires, injury by, 310
Sulphur bacteria, 28; influence of, 278 Teliosorus of cedar apple, figures of
Sunscald, 282 section, 193, 194, 207

Sunscorch, 282 Teliospores, 187; of cedar apple rust,
Suppression, 339 figures of, 208
Surgery of trees, 319 et seq.; figures of, Telium, 187
320 Teleutospore, 187
Susceptibility to disease, 325; to infec- Teratology, 331; book on, 340
tion, 273 Terfas as food of Arabs, 151
Sweet pea diseases, experiments with, Terfeziacese, character of, 151
647; streak, 547 Terfezia, character and occurrence of

Sweet potato black-rot, 548 various species, 151
Swingle, Dean B., studies on columella Test-tube plugging, 586
formation, 96 Tetramyxa, 9; parasitica, 11
Sycamore, 549; blight, 549 Tetranychus mytilaspidis, 296
Symbiotic, 49 Thalloid shoot of Lunularia, figures of,
Symptomatology, 341 361

Symptoms, description of, 640; of Thallophytes, i
disease, 341; of poisoning, 238, 239 Thamnidium chffitocladioides, loi, char-
Synandry, 339 acter of species of, 102; elegans,
Synanthody, 339 figure of, loi
Synanthy, 339 Thaxter, Roland, work of, 172

Syncarpy, 339 Thelephoraceae, characters of family,
Synchytrieae, 117 221

Synchytrium, parasitism of various Thermogens, 25
species, 117; vaccinii, 509 Thesium alpinum, 298
Syncotylous races, 329 Thielavia basicola, 550; figures of, 551,
Synophthy, 339 552; pathogenicity, 149, 150

777
Thiobacteriaceae, 38 Traumatism, 294

Thiocapsa, 39 Treatment of cavities, 321
Thiocystis, 39 Tree surgery, figures of, 320; literature
Thiodictyoiij 39 on, 324; in general, 319
Thiogens, 25 Trehalase, 58
Thiopedia, 39 Trehalose, 53, 58
Thiophysa volutans, diameter of, 22 Trembling fungi, 217
Thiopolycoccus, 39 TremellacesB, characters of family, 217;
Thiosarcina, 39 mucilage in, 52
Thiospirillum, 39 Trichia, 15; chrysosperma with yellow
Thiothece, 39 elaters, 17; fallax, 15; scabra, Plas-
Thiothrix, 38; nivea, 38 modium of, 12; varia with yellow
Thoma's haematimeter, 617; details of, sporangia, 17
618, 620 Trichothecium roseum, 61; chitin in, 52
Threshing machine active in spread of Tricotylous races, 329
smuts, 179 Trimethylamin in spores of Tilletia
Thyridaria tarda, 490 caries, 56
Tillet, Matthieu, mentioned, 182 Tripe de roche, 83

Tilletiaceae, characters of, 182 Triplasy, 339
Tilletia, descriptions of various species, Trophic correlation, 404
184, 185 Trophomorphosis, 404
Tilletia foetans, chlamydospores of, 561; Tropisms of plasmodia, 12
tritici, description of, 184; figure of, Trommelschlagel, 25
183 Truflles and rodents, 68
Tilmadoche mutabilis, figure of, 17 Truffles, occurrence, 151, 153
Timber decay, 553 Trypsin, 58, 59
Timber sap-rot, 558 Tuberaceae, characters of, 151
Tip-burn of potato, 575 Tubercles of velvet bean, figure of, 386
Tissue forms of cecidia, 397 Tuber, characters of various species,
Toodstools, 231 et seq.; guide to de- 153; figures of, 152; Requenii and
scription of, 728, 729 black beetles, 71
Tobacco, 550; mosaic disease of, 578; Tubeuf, Carl von, quoted, 553
root-rot, 550; section of tumor, 392 Tubifera Casparyi, plasmodium of, 12;
Toothwort as a root parasite, 299 ferruginea red plasmodium of, 12
Top-dry, 565 Tuckahoe, 72
Tornadoes, injury by, 311 Tugging-ant and Rozites gongylophora,
Torsion, 339 365
Toxicology of mushrooms, 237, 238, 239 Tumescence, 352
Trama, 232 Tumor on apple stem, figure of, 390
Trametes pini, 519; radiciperda, injury Tumor, figure of section of tobacco, 392
by, 311; robiniophila, occurrence of, Tumors in plants, 34, 342
228; species of, 229; suaveolens, Turnips, brown- rot, figure of, 486
occurrence of, 228 Tyloses, 370; figure of, 369
Transfer of fungi, 624 Tylostomacese, 241
Tranzschelia punctata attack on Hepat- Types of colonies, 626, 627; of stab
ica triloba, 348 cultures, 627
778 INDEX
Tyrosin, 33 Velvet bean tubercles, figure of, 386

Tyrosinase, 58, 59 Venturia inequalis, 479, 480, 481;
Twin cherries, figures of, 334 figures of, 480; pomi, 163
Verpa digitaliformis, 171
U Verticillium albo-atrum, 646
Vibio cholera, rapidity of cell division, 24
Ultramicroscopic organisms, 21
Villia, 141
Umbilicaria on Octorara schist, 83
Violet leaf-spot, figure of, 559
Uncinula, key to species of, 725, 726
Violet spot diseases, 558
Unhappy white elm, figure of, 287
Virescence, 339
Unorganized ferments, 56
Volutin in fungi, 53
Urease, 59
Volva, 232
Urea-splitting enzymes, 59
Uredinales, 187; phylogeny
Von Tavel, Dr. F., cited, 89
of, 197
Uredineae, 187; characters of, 187
Von Wettstein, R., mentioned, 61
Urediniospores, 188
Uredinium, 188 W
Uredo gossypii, 508
Uredospores, 49, 188
Wager, Harold, work of, 135
Wallroth mentioned, 7
Urobacillus Duclauxii, length and
Walter, H., work of, 271
breadth of, 22
Ward, H. M., and ginger beer organisms,
Urocystis cepulas, 531; several species,
140
185
Uromyces Water analysis, 626
betae on beets, 485; figure cf
species, 200; species of, 201; striatus
Water content of tissues and disease, 280
Water-core of apple, 571
on alfalfa, 477; trifolii, 502
Water, influence of, 279
Usnea barbata, mechanic tissues of, 81;
Water-logging, 567
the beard lichen, 83
Watermelons, 525; wilt of, 646
Ustilaginaceas, characters of family, 178
Ustilago
Water requirements of plants, 279, 280
avenae, figures of, 532; of
Wettstein, R. von, 2
several species, 180; levis, figures of,-
Wheat, 560; broth, 599; rust, 188;
532; maydis on maize and teosinte,
forms of, 201, 202; smut, figures of, 562
86; origin of name, 178; zeae, 504, 505,
White pine blister-rust, 537
5c6; figure of, 505; tritici, figures of,
White rust of cruciferous plants, 74
562
Whey, litmus, 600
Will, Dr. H., 142
Wilt, 342
Vaccination, 314 Wilting, experiments
342, 345, 346;
Vaccinium vitis-idasa, gall on, 389 with, 652, 653
Valsaceae, characters of, 163 Wilt of corn, 507; figure of experiment
Van Wisselingh, C, work of, 52 with, 646; of cotton, experiments
Variegation, 343 with, 646; of cowpeas, of egg
646;
Vaucheria, 44 plant, 646; of melons, 525; of sweet
Vegetable slant, "figure of, 597 corn, 644; of watermelon, 646
Velum partiale, 232 Wilson, Lucy L. W. on Conopholis
Velum universale, 232 americana, 301
INDEX 779
Wind action, 295; dissemination of smut

spores, 179; distribution of spores,
Yeasts, 52; 134 et seq.; counting cells
66; its influence on plants, 286
of, 617; character of fermentation,
Wind-swept white poplar, figure of, 287
i37i 595! filrn formation, 137; on
Winkler, H., work of on graft hybrids,
gypsum blocks, 622; spores, 622;
330
sporulation, 137, with zymase, 59
Winogradsky, mentioned, 54
Yellow rust of wheat, 203
Winter, G., mentioned, 61
Yellows of peach, 573
Winterstein, research of, 52
Yolk of eggs, 603
Wire basket, figure of, 624
Youngken, H. \V., 39
Wire worms, 651
Witches' brooms, 72, 342, 348, 395;
on hackberry, figure of, 351
Withering, 652 Zeiller, work on fossil fungi, 82
Wood-boring insects, 310 Zdocecidia, 385
Woody fungi, 218 et seq. Zoogloea, 23
Worsdell, W. C, book of, 340 Zoology emphasized, 271
Wound-cork, 376; description of, 383 Zopf, W., cited, 7, 53, 56; handbook of,
Wounding of plants, artificial, 648 55

Wound- wood, 376, 381, 382 Zygo my ce tales, 5c; absence of cellulose,
52; bibliography of, 105; character
of order, 92, 93; key to families, 97
Zygosaccharomyces, 141
Xylaria Cookei, 62; digitata on old wood, Zygospores, 50
164; hypoxylon, 62; polymorpha on Zymase, 56, 59
old tree stumps, 164 Zymogen, 57
Xylariaccae, characters of family, 164 Zj^mogens. 25
^BOPtRfY UBRARY
W. C. State CMtge
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Qltf^ i. 1. Htll ffithrarg

5JortI| (Earolina ^tuU Iniupraitg

NOfTH CAROLINA STATE

MYCOLOGY AND PLANT

MYCOLOGY AND PLANT

JOHN W. HARSHBERGER, Ph.D.

WITH 271 ILLUSTRATIONS

THE MAPLE PRESS YORK PA

body. The which are given in detailed form are designed to

uniform courtesy of members of the firm of P. Blakiston's Son & Co.,

CHAPTER IV.— Classification of Bacteria • 28

CHAPTER V. Characteristics of the True Fungi 42

CHAPTER VI. Histology and Chemistry of Fungi 52

Histology; Cell Contents; Colors; Physiology; Enzymes;

Enzymes in Fungi; Chemotaxis.

CHAPTER VIII.— Ecology of Fungi 69

CHAPTER XII.— Oospore-producing Algal Fungi 107

dermacese; Sphserobolaceae; Phallomycetes; Development of Carrion Fungi;

PART II. GENERAL PLANT PATHOLOGY

CHAPTER XXIII. General Consideration of Plant Diseases . . .271

Enzymes; Panaschiering; Calico; Nutritive Disturbances; Mutations; Mal-

PART III. SPECIAL PLANT PATHOLOGY

CHAPTER XXXIIL— Specific Diseases OF Plants 411

PART IV. LABORATORY EXERCISES IN THE

tubes, etc.; Lesson 3, Microscopic Study of Culture Material; Stains;

GENERAL STATEMENT AND CLASSIFICATION

Class Myxomycetes (slime moulds).

cation win be an accompHshed fact. The choice of a classification by

treatise is based on that of Engler and Gilg, as published in the seventh

ORDER II. PHYTOMYXALES.

ORDER III. MYXOGASTRALES.

CLASS II. SCHIZOMYCETES.

Family 4. Phycobacteriaceae (Chlamydobacteriaceae).

ORDER II. MYXOBACTERIALES.

CLASS III. EUMYCETES.

ORDER III. ASCOMYCETALES.

ORDER IV. BASIDIOMYCETALES.

Suborder. Uredine^. (Usually Order Uredinales).

Fungi Imperfecti (Deuteromycetes).

SLIME MOULDS (MYXOMYCETES)

should be used. One of the strongest arguments is thai if we consider

name Myxomycetes aligns itself with Schizomycetes and Eumycetes

the Plasmodium, or develop directly into the plasmodium.

the excrements of animals and on the decaying parts of plants. They

rable, though closely packed together until the fructification forms,

of the true shme moulds, but there is no distinct sporangial wall,

stimulation of the host to form gall-hke swelUngs. The whole Plas-

of parts, which become surrounded by membranes to form spores.

A. Spores distinct from each other, irregularly aggregated and fiUing

{a) Spores regular in shape, spheric, (i) Plasmodiophora.

The genus Plasmodiophora comprises possibly three species found

Yellow Fiiligo seplica.

Orange TricJiia scabra.

White Pliysai'um. cUipsoidcum.

The movement of the plasmodium is associated with the incorpora-

extends, devouring by enzyme action the more delicate hyphae.

moves toward a more abundant water supply. It is hydrotropic.

nine months on a plate of glass placed inside of a laboratory case

SLIME MOULDS (m\^OMYCETES) 1

kept a Plasmodium in a streaming condition for over a month be-

globule of protoplasm. A pink flush now begins to appear in the

dome-shaped columellar cavity to the sporangial wall. The threads

Vacuoles are formed in a still further condensation of the sporangial