Untitled

PHARMACEUTICAL BOTANY 

YOUNGKEN

Dryopleris marginalis, one of the ferns whose rhizome and stipes constitute 

the drug, Aspidium. 

(Frontispiece)

PHARMACEUTICAL 

BOTANY 

A TEXT-BOOK FOR STUDENTS OF 

PHARMACY AND SCIENCE 

BY 

HEBER. W. YOUNGKEN, A.M., M.S., PH.M., PH.D. 

PROFESSOR OF BOTANY AND PHARMACOGNOSY AND DIRECTOR OF THE MICROSCOPICAL 

LABORATORIES IN THE PHILADELPHIA COLLEGE OF PHARMACY AND SCIENCE 

THIRD EDITION, REVISED AND ENLARGED 

WITH 238 ILLUSTRATIONS 

AND GLOSSARY OF BOTANICAL TERMS 

PHILADELPHIA 

P. BLAKISTON'S SON & CO. 

1012 WALNUT STREET

A V 

' 

' 

VU 

^-v < vYM " 

COPYRIGHT, 1921, BY P. BLAKISTON'S SON & Co. 

"The use in this volume of certain portions of the text of the United 

States Pharmacopreia is by virtue of permission received from the Board 

of Trustees of the United States Pharmacopoeial Convention. The said 

Board of Trustees is not responsible for any inaccuracy of quotation nor 

for any errors in the statement of quantities or percentage strengths." 

THE MAPLE PRESS YORK PA

PREFACE TO THE THIRD EDITION 

The author has first endeavored to present in a clear, systematic 

way those fundamental principles of structural and taxonomic 

botany which serve as a key to the approach of pharmacognic 

problems. But he has not been unmindful that the use of the work 

has extended to academic institutions and, so, in this edition, has 

broadened the scope of the former text. To this end about ninety 

additional pages of subject matter have been introduced. Several 

old cuts have been removed. Forty-three new ones have been in- 

serted. Hypothetical discussions have been avoided which saves 

time for the reader. 

The arrangement and plan of the chapters are similar to that of 

the former edition, in order to adapt the work to several methods of 

approach. Chapter I on " Fundamental Considerations" has been 

augmented by treatises on Botanical Nomenclature, Farafrme and 

Celloidin Imbedding, Sectioning, Staining and Mounting, Micro- 

tomes and other information dealing with the preparation of materials 

for microscopic examination. 

Ten pages have been added to Chapter V on Cytology. Under 

"Protoplasm and its Properties," six pages have been written on 

the subject of Irritability and Irritable Reactions. Under "Non- 

Protoplasmic Cell Contents" several additional commercial starches 

are discussed and two original plates on starch grains added. Addi- 

tional cuts on Collenchyma, Stone Cells, Sclerenchyma Fibers, 

Trichomes and Fibrovascular Bundles have been inserted in Chapter 

VI. 

Nine additional pages of subject matter and illustrations have been 

added to Chapter VII. Original figures of all of the important 

types of fruits appear here for the first .time. 

vii

Viii PREFACE TO THE THIRD EDITION 

Chapter VIII on "Taxonomy" has been increased by seven pages 

of new data, and the whole former text carefully revised. 

Chapter IX on "Ecology" has been newly introduced as has also 

a Glossary of Botanical Terms. The index has been so planned as 

to make the information contained in this book readily accessible. 

To the authors of works from which cuts were borrowed the 

writer's thanks are due. 

H. W. Y.

PREFACE TO THE SECOND EDITION 

The appearance of the United States Pharmacopoeia IX and the 

National Formulary IV, with the many changes in the lists and 

definitions of officially recognized vegetable drugs made it necessary 

to revise the former edition of this work. 

In the course of revision, the writer has taken cognizance 

of the 

growing 'importance of Botany in* the curricula of pharmaceutical 

institutions and has accordingly expanded upon the subject matter 

of the former text. 

With the adoption for the first time by the new United States 

Pharmacopoeia of pharmacognic standards for numerous drugs, 

Pharmacognosy has risen to the forefront in this country as a science. 

While its proper comprehension requires laboratory instruction in 

chemistry, physics, and crystallography as well as botany, neverthe- 

less a rather extended foundation in structural botany 

stands out 

preeminently as the most needed requirement. 

The work has been for the most part remodeled. Chapter I deals 

with Fundamental Considerations. Chapter 

II is devoted to the 

life history of the Male Fern, a median type of plant, the considera- 

tion of which, after the students have received fundamental practice 

in the use of the microscope, the writer has found commendable, for 

it not only gives beginners a working knowledge of structures and 

functions, the homologies and analogies of which will be met in the 

later study of forms of higher and lower domain, but holds their 

interest on account of its economic importance. 

The life history of a type of Gymnosperm, White Pine, is next 

taken up in Chapter III. Chapter IV considers the life history of an 

Angiosperm as well as coordinates the resemblances and differences 

between Gymnosperms and Angiosperms. Chapters V, VI and 

VII are devoted respectively to Vegetable Cytology, Plant Tissues 

and Plant Organs and Organisms. Among the many additions ta

X PREFACE TO THE SECOND EDITION 

the topics included in these might be mentioned a treatise on Cell 

Formation and Reproduction including Indirect Nuclear Division, 

twenty pages on Non-Protoplasmic Cell Contents, the considera- 

tion of Woods, Root Tubercles, the gross structure and histology of 

different types of leaves, broad histologic differences between Mono- 

cotyl and Dicotyl leaves, the histology of floral parts and the his- 

tology of types of fruits and seeds. Chapter VIII on Taxonomy 

has been increased by the addition of 144 pages. Several new fami- 

lies of drug-yielding plants have been added and the treatment of 

family characteristics has in the majority of instances been broadened. 

The habitats of drug-yielding plants have been added. In 

that portion of the tables dealing with the names of official drugs, 

those official in the National Formulary have been so designated by 

the abbreviation N. F., to distinguish them from others that may 

occur in the same portion of the table and which are official in the 

Pharmacopoeia. 

In keeping with the increased size of the book, many new illus- 

trations have been introduced. A number of these are original 

drawings, photographs and photomicrographs. To the authors of 

other works from which cuts were borrowed, the writer's thanks are 

due. 

The writer in conclusion desires to thank Dr. John M. Macfarlane, 

head of the Botanical Department of the University of Pennsylvania, 

for valuable suggestions during the preparation of portions of the 

text. 

H. W. Y.

CONTENTS 

CHAPTER I 

FUNDAMENTAL CONSIDERATIONS 

DEPARTMENTS OF BOTANICAL INQUIRY i. Morphology 

(Gross Anatomy, Histology, Cytology). 2. Physiology. 3. 

Taxonomy or Systematic Botany. 4. Ecology. 5. Genetics. 

6. Phytopathology. 7. Phytogeography. 8. Phytopalaeontology. 

9. Etiology. 10. Economic Botany and its sub- 

PAGES 

divisions 1-2 

PRINCIPLES OF CLASSIFICATION. Natural System: species^ 

genus; family; order; class; subdivision; division; variety; 

race; hybrid 2-3 

OUTLINE OF PLANT GROUPS 3~4 

BOTANICAL NOMENCLATURE 4-? 

THE MICROSCOPE. Definition. The simple microscope. The 

dissection microscope. The compound microscope: its construction 

and use. The binocular microscope. Rules for the 

care of the microscope 

7-14 

MAKING OF SECTIONS. Free hand sectioning. Kinds of sections. 

Microtomes (hand, sliding and rotary), their construction 

and use 15-20 

THE TECHNIQUE OF MAKING A TEMPORARY MOUNT 

THE TECHNIQUE OF MAKING PERMANENT MOUNTS. 

20 

The mounting medium; Staining. Method for the Prepara- 

tion of a Canada Balsam Mount. Method for the Prepara- 

tion of a Glycerin- Gelatin Mount. Technique of Fixing, 

Dehydrating, Hardening and Imbedding in Paraffine. Technique 

of Sectioning and Mounting Material Imbedded in 

Paraffine. Method for the Staining and Mounting of Material 

in Paraffine Ribbons affixed to slide. Imbedding in Celloidin. 

Sectioning Celloidin Material. Staining and Mounting 

Celloidin Sections 20-30 

DESILICIFICATION. SCHULZE'S MACERATION PROCESS. . 

MICROMETRY. Unit of microscopical measurement. Standardi- 

30-31 

zation of the Ocular Micrometer 31-32

Xll CONTENTS 

CHAPTER II 

LIFE HISTORY OF THE MALE FERN 

. . 

HISTORY OF THE SPOROPHYTE OR ASEXUAL GENERATION. 

PAGES 

Gross structure of stem. Histology of mature stem. His- 

tology of growing apex. Histology of mature root. Histology 

of root apex. Continuity of crude sap flow. Histology of 

stipe. Histology of lamina. Comparative physiology of 

root, stem and leaf. Gross structure and histology of the 

sori and sporangia. Rupture of sporangium and spore dissemination. 

HISTORY OF THE GAMETOPHYTE OR 

SEXUAL GENERATION. Origin of new sporophyte or 

diploid plant from fertilized egg. Growth of seedling into 

mature sporophyte. Alternation of generations 33~44 

% 

CHAPTER III 

LIFE HISTORY OF A GYMNOSPERM (PINUS STROBUS) 

DESCRIPTION OF THE WHITE PINE TREE (SPOROPHYTE). 

Staminate cones. Carpellate cones. DESCRIPTION OF 

THE GAMETOPHYTE GENERATION. The Male Gametophyte. 

The Female Gametophyte. Fertilization. Seed 

Formation and Distribution. GERMINATION OF THE 

SEED 45-52 

CHAPTER IV 

LIFE HISTORY OF AN ANGIOSPERM 

DESCRIPTION OF THE DOG'S TOOTH VIOLET. Development 

of Female Gametophyte. Maturation of the Pollen Grain 

and formation of the Male Gametophyte. Pollination and 

Fertilization. Ripening of the Ovule to form the Seed and 

of the Ovary to form the Fruit. Germination of the Seed and 

development of the Mature Sporophyte 53~58 

RESEMBLANCES BETWEEN GYMNOSPERMS AND ANGIO- 

SPERMS 58 

FUNDAMENTAL DIFFERENCES BETWEEN GYMNOSPERMS 

AND ANGIOSPERMS. 58-59

CONTENTS Xlll 

CHAPTER V 

VEGETABLE CYTOLOGY 

VEGETABLE CYTOLOGY. Definition. The Plant Cell as the 

Fundamental Unit. A typical Plant Cell 

PROTOPLASM AND ITS PROPERTIES. Structure. Metabolism. 

Irritability (Thermotropism, Chemotropism, Sitotropism, 

Hydrotropism, Heliotropism, Geotropism, Galvanotropism, 

Thigmotropism.) Reproduction 

PROTOPLASMIC CELL CONTENTS. Cytoplasm; nucleus; nu- 

cleolus; plastids (leucoplastids, chloroplastids, chromoplastids) 

CELL FORMATION AND REPRODUCTION. Asexual and sexual 

cells. Reproduction defined. Asexual Reproduction: Fis- 

sion; Gemmation; Free Cell Formation; Rejuvenescence. 

Sexual Reproduction: Conjugation and Fertilization 

INDIRECT NUCLEAR DIVISION. 

NON-PROTOPLASMIC CELL CONTENTS. Sugars. Starch. 

Inulin. Hesperidin. Strophanthin. Salicin. Saponin. Coni- 

ferin. Digitoxin. Characteristics of Glucosides. Alkaloids 

and their properties. The alkaloids, Strychnine, Veratrine, 

Nicotine, Caffeine, Cocaine, Aconitine, Colchicine. Gluco- 

alkaloids. Asparagine. Calcium Oxalate. Cystoliths. Silica. 

Tannins. Proteins. Aleurone Grains. Mucilages and Gums. 

Fixed Oils and Fats. Volatile Oils. Resins. Oleoresins. 

Gum Resins. Balsams. Pigments. Latex. Enzymes 

Classification of Enzyrnês 

CELL WALLS. Their formation and composition. Growth in area 

and thickness.- Various kinds of cell walls and behavior of 

each to micro-chemic reagents 

CHAPTER VI 

PLANT TISSUES 

PAGES 

60-6 1 

62-68 

68-70 

70-71 

71-74 

74-96 

96-98 

PLANT TISSUES. Tissue defined. The Tissues of Spermatophytes 

and Pteridophytes : Generative Tissues. List of Tissues 99-100 

MERISTEM. Definition. Primary and secondary meristems; their 

distribution.. 100

XIV 

CONTENTS 

PARENCHYMA. Definition; ordinary parenchyma; assimilation 

PAGES 

parenchyma; conducting parenchyma; reserve parenchyma; 

their structure, distribution and function 101-103 

COLLENCHYMA. Definition, function and distribution 103 

SCLERENCHYMA Definition; stone cells; sclerenchyma fibers; 

wood fibers; bast fibers; their distribution 103-106 

EPIDERMIS. Definition; transpiration and water stomata; epidermal 

papillae; trichomes; scales; their distribution and functions. 106-109 

ENDODERMIS. Definition, distribution and functions 109-110 

CORK. Definition; its derivation, function and distribution .... no 

LATICIFEROUS TISSUE. The structure, origin, 

latex cells, laticiferous vessels and secretory 

distribution of 

cells. Latex . . 110-112 

SIEVE (LEPTOME OR CRIBIFORM) TISSUE. Definition; distri- 

bution; function 112 

TRACHEARY TISSUE. Tracheae: Definition, function and classifi-* 

cation; Tracheids: Definition and function 112-116 

MEDULLARY RAYS. Primary and Secondary; their distribution 

and functions 

* 

116-117 

FIBRO-VASCULAR BUNDLES. Definition of the various types; 

the distribution of each type; Xylem and Phloem 117-119. 

- 

SECRETION SACS. Definition and distribution . 

INTERCELLULAR AIR SPACES. Definition; schizogenous and 

lysigenous air spaces; their relative site 119- 

SECRETION RESERVOIRS. Definition; structure; internal glands; 

secretion canals 119-120- 

CLASSIFICATION OF TISSUES ACCORDING TO FUNCTION . 

CHAPTER VII 

PLANT ORGANS AND ORGANISMS 

PLANT ORGANS AND ORGANISMS. Organ and organism defined. 

Vegetative Organs: Roots, stems and leaves. Reproductive 

Organs: Flower, fruit and seed. Embryo and its parts; function 

of the cotyledon 

119 

120

CONTENTS XV 

THE ROOT. Definition; functions; root hairs; root cap; generative 

PAGES 

tissues; differences between root and stem 121-123 

CLASSIFICATION OF ROOTS AS TO FORM. Primary root; tap 

root; secondary roots; fibrous and fleshy roots; anomalous 

roots; adventitious roots; epiphytic roots; haustoria 123-124 

CLASSIFICATION OF PLANTS ACCORDING TO DURATION 

OF ROOT. Annual; Biennial; Perennial 124 

ROOT HISTOLOGY. A. Monocotyledons. B. Dicotyledons; Histology 

and Development of the California Privet root; Abnormal 

structure of Dicotyl roots; Histology of a Dicotyl Tuberous 

Root, Aconitum 124-132 

ROOT TUBERCLES. Definition; occurrence on roots of Leguminosae, 

Myricaceae, etc.; their etiology 132-135 

THE BUD. Definition; plumule,, scaly buds, naked buds; Classifica- 

tion of Buds According to Position on Stem: terminal bud; 

axillary or lateral bud; adventitious bud; accessory bud. 

Classification of Buds According to Development: leaf bud; 

flower bud and mixed bud. Classification of Buds According 

to their Arrangement on the Stem: alternate; opposite; 

whorled 135-136 

THE STEM. Definition; direction of growth; functions; size; nodes 

and internodes; stem elongation; duration of stems; stem 

modifications; above ground stems; herb and tree defined; 

underground stems; exogenous and endogenous stems. . . . 136-140 

STEM HISTOLOGY. Annual Dicotyl; Perennial Dicotyl; Exceptional 

Types of Dicotyl Stems; Lenticels and Their Forma- 

tion; Annual Thickening of Stems; Method of formation; 

"Annual Ring." Bark: Definition; zones; Periderm; Phello- 

derm; Histology of Cascara Sagrada bark. Wood: alburnum; 

duramen; microscopic characteristics of Angiospermous and 

Gymnospermous woods; Histology of Typical Herbaceous 

Monocotyl Stems; Histology of Typical Woody Monocotyl 

Stem 140-154 

THE LEAF. Definition; leaf functions: photosynthesis; assimilation; 

respiration; transpiration 155 

TYPES OF LEAVES DEVELOPED IN ANGIOSPERMS. Cotyledons; 

Scale leaves; Foliage leaves; Bract leaves: bracts and 

bracteoles; Sepals; Petals; Microsporophylls; Megasporophylls 155-156

XVI CONTENTS 

ORIGIN AND DEVELOPMENT OF LEAVES. Primordial leaf. 

PAGES 

Its formation 156 

PHYLLOTAXY. Definition; spiral law of leaf arrangement; alternate; 

opposite, decussate; whorled; fascicled; leaf rank. ..... 156-157 

VERNATION. Definition; inflexed or reclinate; conduplicate; con- 

volute; circinate; plicate; involute; revolute 157-158 

THE COMPLETE LEAF. Its parts; sessile; petiolate; exstipulate; 

stipulate 158-159 

LEAF VENATION. Furchate; parallel; reticulate; pinni-veined; 

palmately veined; anastomosing veins 159 

LEAF INSERTION. Definition; radical; cauline; ramal; perfoliate; 

amplexicaul; connate-perfoliate; equitant 159-161 

FORMS OF LEAVES. Simple and Compound, (a) General Outline: 

ovate; linear; lanceolate; elliptical; oblong; inequilateral; or- 

bicular; peltate; filiform; oblanceolate; cuneate; spatulate; en- 

siform; acerose; deltoid, (b) Apex: acute; acuminate; obtuse; 

truncate; mucronate; cuspidate; aristate; emarginate; retuse; 

obcordate. (c) Base: cordate; reniform; hastate; auriculate; 

sagittate, (d) Margin: entire, serrate; dentate; ^crenate; 

repand; sinuate; incised; runcinate; lobed; cleft; parted; divided. 

Forms of Compound Leaves 161-166 

LEAF TEXTURE. Membranous; succulent; coriaceous 166 

LEAF COLOR. Variations in color 166 

LEAF SURFACE. Glabrous; glaucous; pellucid-punctate; scabrous; 

pubescent; villose; sericious; hispid; tomentose; spinose; 

rugose; verrucose 166 

DURATION OF LEAVES. Persistent or evergreen; deciduous; cadu- 

cous; fugacious 167 

GROSS STRUCTURE AND HISTOLOGY OF THE LEAF PETIOLE. 

In Monocotyledons. In Dicotyledons. Pulvinus. Peri- 

cladium 167-168 

STIPULES. Definition; lateral; free-lateral; lateral-adnate; lateral- 

connate; lateral interpetiolar. Axillary; ochrea. Modified 

Stipules 168-169

CONTENTS XV11 

THE LAMINA. Definition. Mode of Development of the Lamina 

of Leaves: Dorsoventral; convergent; centric; bifacial; re- 

versed; ob-dorso ventral. A. Dorsoventral: (a) Dorsoventral 

Umbrophytic; (&) Dorsoventral Mesophytic; (c) Dorsoventral 

Xerophytic; (d) Dorsoventral Hydrophytic. Gross Structure 

and Histology of Different Types of Dorsoventral Leaf Blades. 

Gross Structure and Histology of the following types: B. 

PAGES 

Convergent; C. Centric; D. Bifacial 160-173 

STRUCTURE AND DEVELOPMENT OF STOMATA 173-176 

HISTOLOGIC DIFFERENCES BETWEEN LEAVES OF DICOTY- 

LEDONS AND MONOCOTYLEDONS 176 

INFLORESCENCE. Definition; Parts of Inflorescences; Determinate; 

Indeterminate; Mixed Forms of Indeterminate and 

Determinate Inflorescences 177-180 

PRÊFLORATION. Convolute; involute; re volute; plicate; imbricate; 

valvate; vexillary; contorted 180 

THE FLOWER. Definition; floral parts; essential organs; complete; 

perfect; hermaphrodite; regular; symmetrical; imperfect; 

double; staminate; pistillate; neutral; connation; adnation. . 181-183 

THE RECEPTACLE, TORUS OR THALAMUS Definition; varia- 

tions in structure; anthophore; gonophore; gynophore; carpo- 

phore 183 

THE PERIGONE. Definition; dichlamydeous; monochlamydeous . 

THE CALYX. Definition; parts; physical characteristics; connation; 

kinds and forms; persistence; adnation; sepaline spurs; sepaline 

183 

stipules; sepaline position 183-185 

THE COROLLA. Definition; parts; physical characteristics; func- 

*. tions. Forms of the Corolla and Perianth 185-188 

THE ANDRCECIUM OR STAMEN SYSTEM Definition; parts; 

terms denoting number of stamens in flower; insertion of sta- 

mens; proportions of stamens; connation of stamens; stamen 

color. Gross Structure and Histology of the Filament. Gross 

Structure and Histology of the Anther. Anther Dehiscence. 

Development of the Anther. Attachment of Anther. 

Pollen: description; forms 188-195

XV111 CONTENTS 

THE GYNCECIUM OR PISTIL SYSTEM. Definition; Gymno- 

spermous and Angiospermous; parts; the pistil a modified leaf; 

carpel; dehiscence; apocarpous and syncarpous pistils; terms 

denoting the number of carpels entering into the formation of 

the pistil; compound pistils; ovules or megasori as transformed 

buds; position of ovules in ovary; Gymnospermous 

and Angiospermous ovules; structure of Angiospermous ovule; 

PAGES 

shape of ovule 195-197 

THE PLACENTA. Definition; types of placenta arrangement . . . 197 

THE STYLE. Definition; style-arms; relation to carpels forming the 

gynoecium; variations from typical stylar development . . . 198 

THE STIGMA. Definition; forms in wind- and animal-pollinated 

flowers; stigmatic papillae 199 

POLLINATION. Definition; Close and Cross Pollination; terms 

applied to plants pollinated by various agencies 199-200 

MATURATION OF THE POLLEN GRAIN AND FORMATION OF 

MALE GAMETOPHYTE 200 

MATURATION OF THE EMBRYO SAC AND FORMATION OF 

THE FEMALE GAMETOPHYTE 200 

FERTILIZATION IN ANGIOSPERMS. Process; formation of 

embryo and endosperm 

200-201 

THE FRUIT. Definition; modifications .... 202 

FRUIT STRUCTURE. Pericarp; pseudocarp; anthocarp; epicarp; 

mesocarp; endocarp; sarcocarp; putamen; sutures; valves; 

dehiscence 202-204 

CLASSIFICATION OF FRUITS. Simple; Aggregate; Multiple; dry 

dehiscent; dry indehiscent; fleshy indehiscent. Forms of 

Simple Fruits: capsular; schizocarpic; achenial; baccate; 

drupaceous. Etaerio. Forms of Multiple Fruits; strobile or 

cone; sorosis; syconium; galbalus 204-211 

HISTOLOGY OF A CAPSULE, VANILLA 211-212 

HISTOLOGY OF A MERICARP, FCENICULUM 212-213 

THE SEED. Definition; structure; functions; appendages 213-214 

MODE OF FORMATION OF DIFFERENT TYPES OF ALBUMEN. 

Perispermic; endospermic; perispermic and endospermic; 

exalbuminous and albuminous seeds 214-215

CONTENTS XIX 

PAGES 

A MONOCOTYL SEED. Its gross structure and histology 215-216 

A MONOCOTYL SEEDLING. Germination and structure 216-217 

A DICOTYL SEED. Its gross structure and histology -. 217-218 

CHAPTER VIII 

TAXONOMY 

THALLOPHYTA. Definition. Characters of the Protophyta, Myxomycetes, 

Algae, Fungi and Lichenes. The Mounting and 

Staining of Bacteria. Life histories of representative types of 

Thallophytes. Plants of the group yielding drugs and economic 

products 

BRYOPHYTA. Definition. Characters of the Hepaticae and Musci. 

219-282 

Life history of a typical true moss 282-287 

PTERIDOPHYTA. Definition. Characters of the Lycopodineae, 

Equisetineae and Filicineae. Life history of a typical fern. 

Plants of the group yielding official drugs 287-292 

SPERMATOPHYTA (PHANEROGAMIA). Definition. Charac- 

teristics of the Gymnosperms and of the Order and Family 

yielding important drugs and economic products. Table of 

official and unofficial drugs yielded by Gymnosperms including 

part of plants used, botanical origins and habitats. Charac- 

teristics of the Angiosperms and of the classes and families 

yielding drugs. Tables of official and unofficial drugs with the 

names of the plants, parts constituting the drugs and habitats. 292-407 

CHAPTER IX 

ECOLOGY 

PLANT ASSOCIATIONS. Definition. Classification based on re- 

lation plant associations have assumed in regard to water. 

Characteristics of Hydrophytes, Helophytes, Halophytes, 

Xerophytes, Mesophytes and Tropophytes 408-412 

GLOSSARY 412-440 

INDEX 441-479

ERRATA 

Page 1 8 Last word of last line, for '9' read '10'. 

Page 81 Second column, second line, for 'ellipticolar' read 'elliptical or'. 

Page 125 Second line of legend to figure, for rendodermis' read 'endodermis'. 

Page 130 Second line of legend to figure, for 'in Fig. 61' read 'in Fig. 62'. 

Page 138 Ninth line from bottom, for 'suffruitoose' read 'suffruticose.' 

for 'if' read 'in'. 

Page 183 Thirteenth line from bottom, 

Page 183 Seventh line from bottom, for 'monochalmydeous' read 'monochlamydeous 

. 

Page 245 Second line from top, for i 

archegonia read i 

odgonium > 

Page 355 

read Fixed . 

Fifth and sixth lines from bottom of second column, for 'Volatile' 

Page 378 Eight line from top should be placed beneath Apii Fructus. 

Page 418 Tenth line from top, for 'nucleus' read 'nucellus'. 

YOUNGKEN PHARMACENTICAL BOTANY. 

.


CHAPTER I 


Botany 

is the Science which Treats of Plants 

DEPARTMENTS OF BOTANICAL INQUIRY 

1. Morphology treats of the parts, or structure of plants. It is 

divided into: 

(a) Macromorphology or Gross Anatomy which deals with the ex- 

ternal characters of plants or their parts; (b) Micromorphology or 

Histology which considers the minute or microscopical structure of 

plants and plant tissues; and (c) Cytology which treats of plant cells 

and their contents. 

2. Physiology deals with the study of the life processes or func- 

tions of plants. It explains how the various parts of plants perform 

their work of growth, reproduction, and the preparation of food for 

the support of animal life from substances not adapted to that use. 

3. Taxonomy or Systematic Botany considers the classification or 

arrangement of plants in groups or ranks in accordance with their 

relationships to one another. 

4. Ecology treats of plants and their parts 

environment. 

5. 

in relation to their 

Plant Genetics seeks to account for the resemblances and dif- 

ferences which are exhibited by plants related by descent. 

6. Phytopathology treats of diseases of plants. 

7. Phytogeography or Plant Geography treats of the distribution 

of plants upon the earth. The center of distribution for each species 

of plant is the habitat or the original source from which it spreads, 

often over widely distant regions. When plants grow in their 

native countries they are said to be indigenous to those regions. 

When they grow in a locality other than their original home they 

are said to be naturalized.

2 PHARMACEUTICAL BOTANY 

8. Phytopalaeontology or Geological Botany treats of plants of 

former ages of the earth's history traceable in their fossil remains. 

9. Etiology is the study of the causes of various phenomena ex- 

hibited by plants. 

10. Economic or Applied Botany deals with the science from a 

practical standpoint, showing the special adaptation of the vegetable 

kingdom to the needs of everyday life. It comprises a number of 

subdivisions, viz.: Agricultural Botany, Horticulture, Forestry, 

Plant Breeding, and Pharmaceutical Botany. Pharmaceutical 

Botany considers plants or their parts with reference to their use 

as drugs. It interlocks very closely with other departments of 

botanical science. 

PRINCIPLES OF CLASSIFICATION 

The classification of plants is an attempt to express the exact 

kinship which is believed to exist among them. By grouping to- 

gether those plants which are in some respects similar and combining 

these groups with others, it is possible to form something like an 

orderly system of classification. Such a system based upon natural 

resemblances is called a natural system. In a natural system of 

classification every individual plant belongs to a species, every species 

to a genus, every genus to a family, every family to an order, every 

order to a class, every class to a division. In many instances species 

may be subdivided into -varieties or races. The crossing of two 

varieties or species, rarely of two genera, gives rise to a hybrid. 

Thus, the species Papaver somniferum which yields the opium of the 

Pharmacopoeia belongs to the genus Papaver, being placed in this 

genus with other species which have one or more essential characteristics 

in common. The genera Papaver, Sanguinaria and Cheli- 

donium, while differing from each other in certain essential respects, 

nevertheless agree in other particulars such as having latex, perfect 

flowers, capsular fruits, etc., and so are placed in the Papaveraceae 

family. The Papaveracea family and the Fumariacea family are 

closely allied, the latter only differing from the former in having 

irregular petals, usually diadelphous stamens and non-oily albumen 

and so both of these families are placed in the order Papaverales. 

The orders Papaverales, Geraniales, Sapindales, Rhamnales, etc., are


related by a common structure namely, two seed leaves or cotyledons 

and so are grouped together under the class Dicotyledonea. The 

Dicotyledonece differ from the Monocotyledonece 

in that the latter 

group possess but one cotyledon; but both classes agree in having 

covered ovules and seeds, and so are placed in the subdivision 

Angiospermce. The Angiospermce differ from the Gymnospermce in 

that the latter possess naked ovules and seeds; but both of these 

subdivisions agree in producing real flowers and seeds. For these 

reasons they are placed in the division Spermatophyta of the Vege- 

table Kingdom. 

I. Thallophyta 

II. Bryophyta 

III. Pteridophyta 

OUTLINE OF PLANT GROUPS 

i. Protophyta 

2. Myxomycetes 

3. Algae 

4. Fungi 

5. Lichenes 

Hepaticae 

Musci 

Lycopodineae 

Equisetineae 

Filicineae 

[ 

Bacteria 

Cyanophyceae 

Acrasiales 

Phytomyxales 

Myxogastrales 

Chlorophyceae 

Phaeophyceae 

Rhodophyceae 

Phycomycetes 

Ascomycetes 

Basidiomycetes 

Fungi Imperfecti 

Crustaceous 

Foliaceous 

Fruticose 

Marchantiales 

Jungermanniales 

Anthocerotales 

Sphagnales 

Andreaeales 

Bryales 

Lycopodiales 

Selaginales 

I Isoetales 

{ Equisetales 

f Ophioglossales 

1 Filicales

IV. Spermatophyta 


Cycadales 

Ginkgoales 

1. Gymnospermae ~ .? 

Coniferales 

Gnetales 

f Monocotyledoneae 

2. Angiospermae ~. 

I Dicotyledoneae 

BOTANICAL NOMENCLATURE 

Before Carl von Linne, (Linnaeus) the great Sweedish naturalist, 

brought forth the binomial plan of nomenclature, no uniformity 

existed in the assignment of plant names. Among the pre-Linnean 

botanists there were some who designated plants by single names, 

others who employed sentences in naming them, some of which 

were quite lengthy, and a number who adhered to the practice of 

naming them in their own modern tongue. The result was quite 

obvious, a number of systems were employed and confusion pre: 

vailed among students. According to the binomial plan, which 

has been universally adopted, every plant belongs to a species which 

is given two Latin names. The first name is the name of the genus 

or generic name, the second, the name of the species or specific 

name. The generic name corresponds, in the naming of persons, 

to the surname or family name, while the specific name is analagous 

to the given name. Thus, the Wild Cherry is named Prunus 

serotina, Prunus representing the name of the genus, serotina the 

specific name or kind of Prunus. The name of the genus (pi. genera) 

is always a substantive in the singular number and must not be 

applied to more than one genus. Its spelling should begin with a 

capital letter. Genera names may be taken from any source whatever. 

Some, like Fagus for the Beech genus, and Acer for the Maple, 

are of Latin origin. Others have been latinized from other lan- 

guages. Some have been named after some therapeutic property, 

their roots, leaves, flowers or seeds were thought to possess; for 

example, Jafeorhiza, a latinized compound 'of two greek words, 

idreipo:, healing, + ptfa, root, because of the healing virtues of the 

root. A number have had names ascribed to them because of some 

peculiarity of structure, color, taste, odor, behavior, habit or appearance 

of the plant or portion thereof.

FUNDAMENTAL CONSIDERATIONS 5 

Thus, Eriodictyon (from Gr. epiov, wool + 5iKTuo*>, net) alludes to 

its wooly, netted veined leaves; Melaleuca (from Gr. /zeXas, black, 

-J-XevKos, white) alludes to the black bark of the trunk and white 

bark of the branches; Marrubium (from Hebrew marrob, bitter) 

refers to its bitter sap; Barosma (from Gr. fiapvs, heavy + 007*17, 

odor) in allusion to its strong smell; Epiphegus (from Gr. era, upon, 

+ #7776$, the beech) alludes to its growth on the roots of that tree; 

Impatiens (from Lat. in, not and patiens, enduring) refers to the 

sudden bursting of the capsules of this genus when touched; Ly co- 

podium (from Gr. XVKOS, a wolf, + Trot's, a foot) pertains to the 

appearance of the shoots of this genus. Many have been named in 

honor of eminent naturalists or friends of these, or other noted 

persons. For example, Collins onia was named in honor of Peter 

Collinson, an English botanist of the i8th century; Dioscorea in honor 

of Dioscorides, the Greek naturalist \Paullinia after Paullini, a German 

botanist of the i7th century; Cinchona in honor of the countess of 

Chinchon, who brought the bark to Europe in 1640 and Jeffersonia, 

in honor of Thomas Jefferson. 

The specific names are for the most part adjectives which agree with 

the names of genera to which they belong in case, gender, etc. They 

may, however, be nouns and in a few instances consist of two nouns 

or a noun and an adjective. If an adjective it should begin with a 

small letter, as in Rhus glabra and Euonymus atropurpureus. When 

the specific name is a noun, it may either be a proper noun in the 

genitive case when it should begin with a capital, as Garcinia Han- 

buryi] or it may be a common noun in the genitive, when it should 

begin with a small letter, as Grindelia camporum; or the noun may 

be in apposition to the generic name and so in the same case, as 

Cytisus scoparius. Names that had formerly been used for genera 

but since reduced to species are always capitalized, whether origi- 

nally proper nouns or not, as Aristolochia Serpentaria and Anacyclus 

Pyrethrum. In cases where two nouns make up the specific name, 

the first of these is in the nominative case, the second in the genetive, 

the two names being connected by a hyphen, as Capsella bursa-pastoris. 

The botanical name of the species yielding the drug, Aspidosperma, 

(Aspidosperma Quebracho-bianco} will serve as an example of the 

specific portion of the names being composed of a noun and an ad-


jective. Specific names taken from names of persons should always 

begin with a capital. 

Names of varieties are applied in three different ways. Either 

the name of the species is given and followed by the prefix var.' 

before the varietal name, as Chenopodium ambrosioides var. anthelminticum; 

or the varietal name may be appended to the name of the 

species, as Chenopodium ambrosioides anthelminticum\ or the varietal 

name may be placed immediately after the name of the genus and 

the specific name dropped, as Chenopodium anthelminticum. 

It frequently happens that a botanist is careless in naming a species, 

and, without ascertaining whether the same name has been assigned 

to another species, applies it to his, thus causing duplication. For 

example, there are two distinct species of plants named Prunus 

virginiana, one of these, the Wild Black Cherry, the other, the Choke 

Cherry. In this instance the name Prunus virginiana does not tell 

us which species the writer or speaker refers to. It might be the 

Choke Cherry named ft 

Prunus virginiana" by Linnaeus or the Wild 

Black Cherry named "Prunus virginiana" by Miller at a later date. 

Accordingly, it is necessary to add to the name of the species the 

author's name. Thus, Prunus virginiana Linne refers to the Choke 

Cherry while Prunus virginiana Miller refers to the Wild Black 

Cherry. In this connection it is customary to abbreviate the name 

of the author thus, L. for Linne, Mill, for Miller, Ait. for Aiton, 

Loisel. for Loiseleur-Deslongchamps, or Tourn. for Tournefort. 

Whenever a plant is transferred from one genus to another, it 

must retain its original specific name, unless the genus to which it 

is transferred already possesses a species with that name, in which 

case a new specific name must be given it. Moreover, 

the name of 

the botanist who assigned the original specific name but placed it 

under a different genus must be placed in parenthesis between the 

specific name and the name of the botanist who later connects it 

with another genus. For example, we read as the official definition 

for Purging Cassia in the National Formulary IV "The dried fruit 

of Cathartocarpus Fistula (Linne) Persoon." The significance of the 

name Linne in parenthesis is that he had previously given the specific 

name Fistula to the plant indicated but placed it under a different 

genus, which genus happened to be Cassia. Therefore Persoon,


in connecting it up with another genus Carthartocarpus, avoided 

binomial duplication by interjecting Linne parenthetically between 

his name and the specific name Fistula. 

The names of families are designated by the name of one of their 

principal genera or ancient generic names with the ending aceae, e.g., 

Ranunculaceae from Ranunculus, Malvaceae from Malva, etc. The 

following names, because of long usage, are exceptions to the rule: 

Palmae, Gramineae,Guttiferae, Umbelliferae, Labiatae, and Compositae. 

Orders are generally designated by the name of one of their prin- 

cipal families, with the ending ales, e.g., Rhamnales from Rhamnaceae, 

Resales from Rosaceae. Suborders are likewise designated, but with 

the ending ineae, e.g., Mahineae from Malvaceae. Other older 

endings may, however, be retained for these names providing they 

do not lead to confusion or error. 

Names of classes, subclasses, divisions and subdivisions are desig- 

nated from one of their characters by words of Latin or Greek origin, 

some similarity of form and ending being given to those that desig- 

nate groups of the same nature, as Monocotyledoneae, Dicotyle- 

doneae; Archichlamydeae, Metachlamydeae; Thallophyta, Sperma- 

tophyta; Gymnospermae; Angiospermae. 

In the case of Cryptogams the use of old family names as Algae, 

Fungi, Lichens, Musci, etc. is permissible for designating groups 

above the rank of family. 

THE MICROSCOPE 

A microscope is an optical instrument, consisting of a lens, or 

combination of lenses, for making an enlarged image of an object 

which is too minute to be viewed by the naked eye. 

Microscopes are of two kinds, viz. : simple and compound. 

THE SIMPLE MICROSCOPE 

This consists simply of a convex lens or several combined into a 

system and appropriately mounted. A good example of a simple 

microscope is a reading glass. This type of simple microscope is 

valuable in field work, in the examination of dried herbarium material 

or the external characters of crude drugs, where only a low 

magnification of the object is required.


But when flowers or other plant parts are to be dissected, it is 

necessary to have both hands free. To meet this need various 

forms of stands have been devised which have been combined with 

an arm and lens to constitute what are known as "Dissecting 

Microscopes." One of the simpler forms of these is shown in Figs. 

i and 2. It consists of a low wooden stand with inclined sides that 

furnish convenient hand rests for the operator. In the center of 

PlG. I. Front view of a dissecting microscope. Description in text. 

FIG. 2. Rear view of same. 

the upper surface of the stand is a glass plate on which the object 

to is 

be^dissected placed. Beneath this a mirror is set which reflects 

light to the object. On either side of the mirror is a hollow cut out 

which permits light to strike the mirror from various angles. A 

lens arm fits in an aperture just behind the center of the glass place. 

The carrier on the end of the horizontal portion of this accomo- 

and down or from 

dates the magnifier. 

The arm can be moved up 

side to side in securing a focus. The rear of the block is hollowed 

out, providing a convenient receptacle for dissecting tools.


THE COMPOUND MICROSCOPE 

A. Its Construction: 

The principal parts of a compound microscope are: 

1. The base, generally horseshoe shaped, which rests on the table. 

2. The pillar, an upright bar, which is attached to the base below 

and supports the rest of the instrument. 

3. The stage, a horizontal shelf upon which is placed the preparation 

or slide to be examined. The stage is perforated in the center 

for transmitting light reflected up by the mirror. On the stage are 

two clips for holding the glass slide. 

4. The mirror, situated below thq stage, by which the light is 

reflected upward through the opening in the stage. 

5. The diaphragm, inserted in the opening of the stage or attached 

to its lower face, and used to regulate the amount of light reflected 

by the mirror. 

6. The body tube, a cylinder which holds the draw tube and lenses 

and moves up and down perpendicularly above the opening in the 

it back and 

stage. The tube is raised or lowered either by sliding 

forth with a twisting movement or by a rack and pinion mechanism. 

The latter is called the coarse adjustment. 

7. The fine adjustment, a micrometer screw back of the tube, 

which, on being turned, produces a very small motion of the entire 

framework which holds the body tube. 

8. The oculars or eyepieces which slip into the upper end of the 

draw tube. Each of these consist of two plano-convex lenses, the 

lower one being the larger and known as the field lens because it 

increases the field of vision. The upper or smaller lens is called the 

eye lens. It magnifies the image formed by the objective. Midway 

between the field and eye lens is a perforated diaphragm, the object 

of which is to cut out edge rays from the image. 

According to the system adopted by the maker, oculars are designated 

by numbers, as i, 2, 3, 4, etc., or by figures which represent 

focal lengths. 

9. The objectives, which screw into the bottom of the body tube 

or nose piece. They consist of a' system of two, three or four lenses, 

some of which are simple, others compounded of a convex crown 

lens and a concave flint lens. Objectives like oculars are usually

IO PHARMACEUTICAL BOTANY 

designated by numbers or by figures, as 3^2, M> ?> etc -> 

or in milli- 

meters, as 2 mm., 4 mm., 16 mm., which represent focal lengths. 

The smaller the number or fraction representing the focal length 

of an objective, the greater is its magnifying power. 

NoSEPlECE 

Oextcnves -I.j^ 

GRADUATED SHORT SLIDE 

REVOLVING 

STAGE 

ADJUSTABLE 

SPRING FINGER 

CONDENSER; MOUNTING OK 

DROP -SWING ARM*-~ "" , 

LOWER IRIS DIAPHRAGM-'' 

TOR OBUQUE 

STAGE CENTERING 

MIRROR 

MIRROJR 

MIRROR. BAR 

RACK & PINION 

FIG. 3. Illustrating the parts of a compound microscope. 

Objectives are either dry 

CONCENTRIC Awuyr- 

KG BUTTONS 

GRADUATED LONG 

lenses or immersion lenses. If an air 

space be present between the objective and the object, 

the lens is 

called a dry one; if a liquid is present between the objective and the 

object, the lens is called an immersion lens. If this liquid be oil,

FUNDAMENTAL CONSIDERATIONS II 

FIG. 4. Diagram illustrating optics of a compound microscope in use. Fi, 

Upper focal plane of objective; F2, Lower focal plane of eyepiece; A, Optical tube 

length = distance between Fi and Fa; Oi, object; Oz, real image in F2, transposed 

by the collective lens, to O 3 , real image in eyepiece diaphragm; O-t, virtual 

image formed at the projection distance C, 250 mm. from EP, eyepoint; CD, 

condenser diaphragm; L, mechanical tube length (160 mm.); i, 2, 3, three pencils 

of parallel light coming from different points of a distant illuminant, for instance, 

a white cloud, which illuminate three different points of the object. (Courtesy 

of Bausch and Lomb Optical Co.)


the objective is called an oil immersion objective; if water, a water 

immersion objective. 

Some microscopes are fitted up with a nose piece, capable of carrying 

two or three objectives, which may be revolved into place at the 

FIG. 5. FIG. 6. 

FIG. 5. Microscope lamp useful in illuminating opaque objects. 

FIG. 6. Compound microscope with binocular body, designed to relieve 

eyestrain for those spending many hours with the microscope. Each oculai 

inclines at an angle of 4 degrees from the perpendicular, which results in their 

converging to a point about 17 inches from eyes having the average pupillary 

separation. Adjustment for pupillary distance is accomplished by turning 

a knurled ring on the right hand ocular tube which gives a horizontal sliding 

movement of the oculars. The knurled ring on the lelt hand ocular tube provides 

a means of focusing one eyepiece independently of the other. (Made by 

Spencer Lens Co.) 

lower end of the body tube. Others have a condenser which is 

employed to concentrate the light upon the object examined. 

B. Its use: 

1. Place the micrqscope on the table with the pillar nearest you. 

2. Screw the objectives into the nose piece and slip an ocular into 

the upper end, if not already on instrument. Turn the lowest power 

objective into position.


3. Find the light by looking into the ocular (eye piece) and at the 

same time turning the mirror at such an angle that it reflects light 

from the window or lamp up through the opening in the stage to 

the objective. When opaque objects are to be illuminated, a 

stronger illumination is required than that usually afforded by an 

from a window. For this 

ordinary laboratory lamp or by the light 

purpose a microscope lamp, such as the Spencer no. 374 (see Fig. 

5) is very satisfactory. Mirrors have two faces, a plane and a concave. 

Use the concave unless employing the condenser, when the 

plane mirror should always be used. 

4. Regulate the quantity of light by the diaphragm. If too bright 

it must be cut off somewhat. The higher powers require brighter 

light than the lower. 

with the 

5. Place the slide on the stage in a horizontal position 

object over the middle of the opening through which light is thrown 

from the mirror. 

6. With the lower power in position, move the coarse adjustment 

until either the object or small solid particles on the slide appear 

distinctly, which means that the lenses are in focus. The object, 

if not under the lens, may now be brought into the field by moving 

the slide back and forth very slowly while looking through the ocular. 

To improve the focus, slowly turn the fine adjustment screw. 

7. To focus with- the high-power objective, first find the object 

with the low power and arrange in the center of the -field. Put clips 

on slide without moving it. Raise the body tube by means of the 

coarse adjustment. Then turn the high-power objective into posi- 

tion. (If two objectives only accompany your instrument, the high- 

power is the longer one.) Lower the body tube carrying the objec- 

tive until the objective front lens nearly touches the cover glass. 

A slight movement of the fine adjustment should show the object 

clearly. Never focus down with the high-power objective while 

looking through the ocular because of the danger of pressing 

the cover glass and ruining the delicately mounted lenses. 

it into 

8. Accustom yourself to use both eyes indifferently and always 

keep both eyes open. If right handed, observe with the left eye, 

as it is more convenient in making drawings. 

9. When the oil immersion objective is to be used, a small drop of

14 


immersion oil (slightly evaporated cedar oil) should be placed on the 

cover glass directly above the object and the body tube should be 

run down with the coarse adjustment until the front lens of the 

immersion objective enters the drop and comes almost into contact 

with the cover glass. This should be done while watching the objective. 

Then look through the ocular and draw the objective up with 

the fine adjustment until the object comes into focus. 

RULES FOR THE CARE OF THE MICROSCOPE 

1. In carrying the microscope to or from your table, grasp it 

firmly by the pillar and hold in an erect position, so that the ocular 

which is fitted loosely into the draw tube may not fall out and its 

lenses become damaged. 

2. Never allow the objective to touch the cover glass or the liquid 

in which the object is mounted. 

3. Never touch the objective or ocular lenses with fingers or cloths. 

4. Never change from lower to higher power objective without 

first ascertaining that the body tube has been raised sufficiently to 

allow the high power objective to be slipped into place without 

injury to the objective or mounts. 

5. Never clean the microscope lenses or stand with cloths that 

have been used for removing surplus of alkali, acid or other reagent 

from slides. 

6. Note whether the front lens of the objective is clean before 

attempting to use it. If soiled, breathe on the lens and gently wipe 

with an old, clean, soft handkerchief or lens paper. If the lens be 

soiled with balsam or some other sticky substance, moisten the 

handkerchief or lens paper with a drop of xylol, taking care to wipe 

it perfectly dry as soon as possible. 

7. Do not let the objective remain long near corrosive liquids, 

such as strong solutions of iodine, corrosive sublimate, or mineral 

acids. Never examine objects lying in such fluids without putting 

on a cover glass. 

8. Never lift the slide from the stage, but, after raising the objective, 

slide it off the stage without upward movement. 

9. Never allow the stand (microscope without lenses) to be wetted 

with such substances as alcohol, soap, etc., which dissolve lacquer. 

10. Keep the microscope covered when not in use.


MAKING OF SECTIONS 

Free-hand Sectioning. Free-hand sections are usually satisfactory 

for the general examination of roots, stems, leaves, barks and many 

fruits and seeds. Material which is fresh may be sectioned at once, 

but dry material should be well soaked in warm water before using. 

Very hard material like heartwoods, the shells of nuts and seeds, 

may be softened in solution of caustic potash or ammonia water 

and then washed free of alkali before sectioning. 

The object to be sectioned is held between the thumb and finger 

of the left hand. If tender and flexible, such as a flat leaf, it must 

be placed between the two flat surfaces of elder pith before sectioning. 

A segment of pith about an inch long is halved lengthwise 

with a sharp knife and a portion of the leaf is held between the halves 

of pith while the section is cut through pith and leaf. The pith 

is later separated from the leaf section. Sections through other 

delicate parts of plants may be made in the same way, only a groove 

should be made in the pith of such size as is necessary to hold the 

material firmly enough without crushing it. In certain instances, 

when, because of the smallness of the object and its resistance to 

cutting, good sections can not readily be made with the aid of pith, 

a small sized cork stopper can be used with better results. A hole 

just large enough to prevent the object from slipping is made in the 

center of the smaller end and the object inserted preparatory to 

sectioning. The upper surface of the razor is wetted with water or 

50 per cent, alcohol. The razor, which should be real sharp, is held 

in the right hand and is drawn across the object with the edge toward 

the student and the blade sliding on the forefinger of the left hand. 

The sections should be cut as thin as possible. As soon as a number 

of sections have been cut, they can be transferred to a vessel of water 

with a camel's hair brush. 

Sectioning in Paraffine or Celloidin. When it is necessary to study 

the microscopic structure of very delicate plant parts, superior results 

can generally be obtained by imbedding the material in paraffine 

or celloidin, which is subsequently hardened, and sectioned by means 

of a sliding or rotary microtome.

i6 PHARMACEUTICAL BOTANY 

KINDS OF SECTIONS 

1. A transverse or cross-section is one made horizontally through 

the object, hence its plane lies at right angles to the long axis. 

2. A radial-longitudinal .section is one which is made parallel to 

FIG. 7. Showing the 

planes in which sections 

are cut, A, transversely; 

B. longitudinal radially; 

C, longitudinal tangentially. 

(After Stevens.) 

the long axis of the object in such a way 

that it lies in plane of the radius. 

3. A tangential-longitudinal section is one 

made parallel to a plane tangent to the 

cylinder. This type of section is therefore 

prepared by cutting parallel 

long surface. 

MICROTOMES 

to the outer 

Microtomes are instruments employed to 

facilitate the cutting of sections of organic 

tissues. The three most commonly used 

types are the hand, sliding and rotary 

microtomes. 

Hand Microtome.- This type is shown in 

Fig. 8. If the object is sufficiently hard to 

bear the strain, it is placed directly in a 

clamp at the upper end of the tube that is 

tightened by the screw seen on the side of 

the tube, or it may first be inclosed in elder 

pith or cork and then clamped in. The 

object to be sectioned is raised a little at a 

time through the hole in the glass plate at 

the top by turning the finely graduated feed 

near the base of the tube. The section razor 

is then laid flat on the glass plate and pulled 

across the object with a long sliding motion. 

The upper surface of the razor blade is kept wet with 50 per cent, 

alcohol and after several sections have been cut they can be swept 

by the finger or camels hair pencil to a dish of water. Each 

division of the feed represents 10 microns, so that the thickness of 

sections desired can be regulated by moving the feed, accordingly, 

just before each stroke of the razor.


Sliding Microtome. This type of microtome (see Fig. 9) is adapted 

for cutting all kinds of sections. It consists of an iron supporting 

fame of horizontal and upright portions. The horizontal base rests 

on the table and is hollowed out to accommodate a drip pan that 

can readily be removed and cleaned. 

The front of the upright portion exhibits a frame which accomo- 

dates a sliding feed mechanism to which is attached the object 

carrier. The top of the upright portion 

shows a V shaped bed which carries a 

solid iron block which can be readily 

slid along the bed when the latter is 

lubricated with paraffin oil. The upper 

surface of the block is grooved to accomodate 

the thumb screw. The 

microtome knife consists of a blade 

a.. 

FIG. 8. Hand microtome. PIG. 9. A sliding microtome. Blade (a) ; lever 

Description in text. (6). (From McJunkin.) 

portion, that is flat on its lower and hollow ground on its upper 

face, and a forked handle. The latter is slid into the stem of^the 

thumb screw which has . previously been slid into the groove of 

the block and its position adjusted. Sections of woody material 

can be cut directly on this microtome and placed in dilute alcohol. 

When paraffine sections are cut, the cutting edge of the knife 

should be parallel to the motion but when celloidin sections are 

desired the knife must be set at an oblique angle to the frame 

and drawn across the block with a long sliding motion. The knife

1 8 PHARMACEUTICAL BOTANY 

FlG. 10. Rotary microtome. The feed mechanism is covered to protect the 

wearing parts from dust. 

FIG. ii. Plan of construction of rotary microtome shown in Fig. 9.


and the top of the celloidin block must be constantly kept wet with 

80 per cent, alcohol. 

The object is placed in the object carrier and clamped in. By 

means of the graduated disk at the base of the feed mechanism the 

thickness, in terms of microns, is regulated after each stroke of the 

razor. 

Rotary Microtome. When paraffin ribbons are desired, especially 

for the study of serial sections of material, the rotary microtome 

surpasses by far the efficiency of the sliding type of instrument. The 

Spencer Rotary Microtome No. 820 is shown in Fig. 10 and its 

plan of construction illustrated in Fig. n. In this instrument the 

sliding part which carries the object clamp (SP) is carried up and 

down by the block (B). The feed mechanism consists of a rigid 

bearirfg, on which the feed block (FB) (of which the projection P is 

a part), is moved by the feed screw (FS). As this block travels to- 

ward the side on which the balance wheel (W) is located, the sliding 

is forced forward towards the knife one-half as much. The 

part (SP) 

polished surface set against the point (P) is arranged at the proper 

angle to accomplish this end. The screw, 

cut with two threads to 

the millimeter, is revolved by a ratchet feed wheel with 250 teeth. 

Each tooth represents a forward movement of the object of one mi- 

cron. The feed can be set for sections from i micron to 60 microns 

thick, by turning the button at the back of the case until the number, 

representing the desired thickness, appears opposite the indicator at 

the small opening in the side of the case near the balance wheel. 

The total excursion of the feed is 37 mm. This allows a sufficient 

range for cutting a complete series of sections of a large object without 

the necessity of a break due to resetting the knife and feed mechanism. 

The object, after being placed in the object clamp, may 

be oriented to any desired angle. The clamp is held at its upper 

limit for orienting or trimming the block by pushing in the pin (F.) 

The whole knife support may readily be adjusted to and from the 

a lever connected 

object, and is readily clamped in any location by 

with an eccentric cam. The knife is fastened by two clamps and 

may be turned to any desired angle. The clamps can also be moved 

toward each other to bring them as near to the ribbon as desired to 

gain additional rigidity. The groove in the balance wheel is de-


signed for a cord or strap when it 

by a motor. 

is desired to run the instrument 

THE TECHNIQUE OF MAKING A TEMPORARY MOUNT 

1. Place a drop or two of water (or reagent) in the center of a clean 

glass slide. 

2. With the aid of a forceps take the section or very small quantity 

of the material to be examined and spread it on the drop of water. 

3. Place a clean cover-glass over the material. In placing the 

cover-glass do not drop it flat upon the drop of water, but place one 

side of it down first and allow it to squeeze the water along under it. 

4. Keep the top of the cover-glass dry. 

When filamentous algae or molds are to be examined, the material 

tends to cling together and must be carefully separated, in the drop 

of water, with dissection needles before the cover glass is placed over 

the material. In case a coarse ground drug 

is to be mounted the 

coarser particles should be first crushed in the water on the slide 

and subsequently teased apart with dissection needles. 

Care should always be taken to see that the water or mounting 

medium used is not contaminated with foreign substances. This 

can best be practiced by examining the mounting medium under the 

microscope before the material to be examined is placed in it. 

THE TECHNIQUE OF MAKING PERMANENT MOUNTS 

i. The Mounting Medium. When a microscopic object is to be 

preserved permanently it must be kept from decaying and the fluid 

in which it is placed must be kept from evaporating. These condi- 

tions can be met by adding an antiseptic (2 per cent, acetic acid, or 

formaldehyde) to the water used in mounting and carefully sealing 

the cover glass with asphaltum or zinc white. As a rule, a better 

way is to use a mounting medium that will not evaporate, e.g., 

glycerine, glycerin gelatin or Canada balsam. These fluids have 

a high refractive index and so render the objects penetrated by them 

more transparent. This quality is generally an advantage, but for 

objects already almost transparent it is quite the reverse. Glycerine 

has the disadvantage of always remaining soft, so that the mount 

may at any time be spoiled by careless handling. "Glycerin-gelatin


' 

has the advantage of mixing readily with 50 per cent, glycerin in 

which the object should be placed before being mounted in this 

medium. It should be warmed on a water bath before using and the 

cover glass applied quickly after it is placed on the specimen. It 

cools rapidly and constitutes the quickest and simplest means of 

effecting a durable permanent mount. Its disadvantage 

is due 

mainly to its jelly like consistency which is frequently responsible 

for damaged mounts when the cover glass above the preparation is 

too greatly strained. Canada balsam, slowly becomes solid, so 

that the mount is exposed to no accident short of actual breakage. 

Balsam has the disadvantage of being non-miscible with water, so 

that before it can be used the object must be carefully dehydrated. 

Even after this is done, and the object lying in absolute alcohol, an 

oil must be used as an intermediate agent between alcohol and 

balsam. 

2. Staining. For two reasons it is generally better to stain plant 

tissues before mounting. Transparent tissues may become almost 

invisible in glycerine, glycerin-gelatin, or balsam, and different 

tissues take a stain differently. This being the case it becomes 

possible to stain one tissue and not another, 

or one tissue with one 

stain and another in the same section with a different stain, so that 

the different parts may be brought out like areas on a colored map. 

The most common stains are haematoxylin derived from logwood, and 

various anilin stains safranin, fuchsin, eosin, iodine green, methyl- 

green, malachite green, etc. 

METHOD FOR THE 'PREPARATION OF A CANADA BALSAM MOUNT 

1. Stain object with 0.5 per cent, solution of safranin or fuchsln 

in 50 per cent, alcohol for from three to five minutes. 

2. Wash out excess of stain and further dehydrate with 70 per 

cent, alcohol. 

3. Stain with 0.5 per cent, solution of methyl-green, or malachite 

green, or iodine-green in 70 per cent, alcohol for twenty seconds or 

longer, depending upon the nature of the material. 

4. Dehydrate and wash out excess of stain with 95 per cent, alcohol 

for two minutes.


5. Further dehydrate by placing 

one minute. 

6. Clear in cedar oil for i minute. Blot up 

edge of section. 

7. Mount in Canada balsam. 

8. Label slide. 

material in absolute alcohol for 

excess from around 

Should air-bubbles be detected in the balsam shortly after mount- 

ing, heat a dissection needle in a flame and touch each with its tip, 

when they will be found to disappear. 

If too much Canada balsam has been used, some of it usually 

spreads beyond the edge of the cover-glass, or on its surface. In 

this event wait until the balsam hardens, when it can be scratched 

off with a knife, and the surface of the glass cleaned with a rag 

moistened with turpentine oil or xylol. 

Should the Canada balsam become too thick, it can be thinned 

down with either xylol or benzol. 

METHOD FOR THE PREPARATION OF A GLYCERIN-GELATIN MOUNT 

1. Stain object with an aqueous solution of eosin. 

2. Wash out excess of stain by moving 

dish of water. 

the section about in a 

3. Transfer object to weak glycerin (glycerin 10 parts, water 

90 parts) for 3 to 5 minutes. 

4. Transfer object to 50 per cent, glycerin for 3 to 5 minutes. 

5. Transfer object to concentrated glycerin for 5 minutes. 

6. Remove excess of glycerin around object and mount in glycerin- 

gelatin. The slide and cover slip 

should be warmed before the 

glycerin-gelatin is dropped over the object and the cover slip quickly 

lowered. The preparation of Glycerin-Gelatin is as follows: Mac- 

erate 14 grams of gelatin in 84 mils of water for 2 hours, add 76 mils 

of glycerin and warm; add 2 mils of liquefied phenol, warm and 

stir for 15 minutes until clear. Filter while hot through glass- 

wool or filter paper and collect the filtrate in a wide mouthed bottle. 

Keep well stoppered so as to exclude dust. 

Glycerin- Gelatin becomes solid when cool. For use warm the 

bottle in a water ttath after first removing the stopper. A glass rod


sufficiently long to reach to the bottom of the bottle can be inserted 

in the cork and used for transferring the material to the slide. 

7. Ring mount with zinc white or asphaltum at the edge of cover 

slip. If the cover slip is circular, this can best be done by means 

of a centering turn-table. A camel's hairbrush is dipped into the zinc 

white or asphaltum and held to the margin of the cover slip while 

the slide fastened with clips to the turn table, is rotated with it. 


If the objects or sections are such as not to be liable to shrink they can 

be transferred from water directly to glycerin-gelatin. 

TECHNIQUE OF FIXING, DEHYDRATING, HARDENING AND IMBED- 

DING IN PARAFFIN 

When the intention is to study the protoplasts in their natural 

form or the processes of cell division, the fresh material must be 

put through the various stages of fixation, hardening and imbedding 

before it is sectioned. The steps will now be considered in the order 

in which they must be carried out. 

Fixation. This is the process of killing and coagulating the protoplast. 

The essence of good fixation is in rapid killing. It should be 

simultaneous with coagulation or hardening so that the protoplast 

will not be modified by later treatment. Fixing fluids are always 

substances unknown to protoplasm e.g. poisons. The coagulation 

of protoplasmic structures is due to the fact that these are alkaline 

in reaction whereas the fixing fluid is acid. Fixing fluids must be 

but also as to reaction 

judged not only as to killing and hardening 

of tissues to stains afterward. Fluids that are mixtures make the 

best fixing agents. Among the fixing agents employed are the fol- 

lowing: Osmic acid (OsO 4) comes in sealed glass tubes containing 

0.5 gm. or i gm. It has a very powerful odor and is easily affected 

by organic materials. It is used in i to 2 per cent, solutions and 

should be made up in distilled water. It fixes cytoplasm well but 

the nucleus not as good. Its disadvantage lies in its inability to 

penetrate rapidly. 

Chromic acid (CrOs) in 0.5 to i per cent, aqueous solution is very 

favorable for nuclear structure but like osmic acid penetrates rather 

slowly. 

Picric acid C6H 2 (OH)(NO 2)3 is one of the most penetrating


fixing fluids but has very little hardening power. It is employed 

in saturated aqueous solution. 

Corrosive sublimate (HgCl 2 ) in 0.2 per cent, aqueous 

or alcoholic 

solution penetrates and hardens rapidly but doesn't give as sharp opti- 

cal differentiation as the others considered. 

Absolute alcohol can be employed for very small objects that are 

dry. If the objects are moist shrinkage will follow. 

Carnoy fluid; consisting of 6 parts absolute alcohol, 3 parts 

formaldehyde and i part of glacial acetic acid, can also be used for 

fixing small objects. It has the advantage of fixing these in about 

10 minutes. Moreover the objects can be carried directly to abso- 

lute alcohol, thence to Paraffin and imbedded. 

For most materials the Flemming fluids have proven very satis- 

factory and are the most generally employed. They are of two 

involve chromic acid and acetic 

classes, viz; i. Those that simply 

acid (the chrome-acetic fluids) and 2. Those that involve chromic 

acid, acetic acid and osmic acid (the Chrome-Osmium-Acetic Fluids). 

The formulae follow: 

Strong , 

CHROME-ACETIC FLUIDS 

i per cent. Chromic acid solution. . .100 

mils 

. , . . . ., 

Glacial acetic acid i mil 

i per cent. Chromic acid solution 70 mils 

Medium i per cent. Glacial acetic acid i mil 

Distilled water 29 mils 

i per cent. Chromic acid solution . . 25 mils 

Weak i per cent. Glacial acetic acid 10 mils 

Distilled water 65 mils 

CHROME-OSMIUM-ACETIC FLUIDS 


Strong 2 per cent. Osmic acid solution 20 mils 

Glacial acetic acid 5 mils 

Weak . 


i per cent. Osmic acid solution 10 mils 

. , 

i per cent. Acetic acid solution 10 mils 

Distilled water 55 mils


The acetic acid in all of the Flemming fluids is of great advantage 

since it penetrates very rapidly, carrying the chromic acid or chromic 

and osmic acids into the tissue depths, thus insuring complete 

fixation. 

The material to be fixed should be cut into small pieces not longer 

than 5 mm. nor broader than 2 or 3 mm. The amount of fixation 

to be used should not be less than 15 times the bulk of the material 

to be fixed. The material should be placed in the fixing fluid im- 

mediately after it is gathered. One or two drachm homepathic 

phials are convenient for the process. The material is kept in the 

fixing fluid for from 12 to 24 hours and then washed in small cheese 

cloth bags which are placed in running tap water for from 6 to 12 

hours or over night. 

Dehydrating and Hardening. After washing the material, still 

kept in the bags, is placed in 10 per cent, alcohol for i hour and is 

then carried through a series of alcohols. Each of the series 10 

per cent stronger than the one before it, remaining in each grade for 

ij^ to 2 hours until 70 per cent, alcohol is reached. Take out of 

bag and place in phial in 70 per cent, alcohol. If the material is 

not to be imbedded in paraffine immediately, it can remain in 70 per 

cent, or 85 per cent, alcohol (if very delicate) until needed. It is 

not safe to leave very valuable material in a grade below 70 per cent, 

over night. From the 70 per cent, alcohol it is carried to 85 per 

cent, to 95 per cent, to absolute alcohol, remaining in each at least 

6 hours'with 2 or 3 changes of the last. 

Clearing and Imbedding. In order to get the material from ab- 

solute alcohol into paraffine, some medium must be used which mixes 

with absolute alcohol and which also dissolves paraffine. Either oils 

such as cedar, clove or bergamot or substances like xylol, chloroform 

or benzol satisfy this requirement. To clear with xylol transfer 

material from absolute alcohol to a mixture of âbsolute alcohol and 

y xylol for 12 hours, then to mixture of equal parts of absolute 

alcohol and xylol for 12 hours, then to % xylol and J^ absolute 

alcohol for 12 hours to pure xylol for 12 hours. To phial containing 

material in pure xylol add paraffine in small pieces and put on top of 

paraffine bath sufficiently long until paraffine is melted. Then add 

more paraffine and put phial in paraffine bath at 56C. over night.


Pour fluid off and add pure melted paraffine and repeat 2 or 3 times 

until rid of all trace of xylol. A tray is then prepared by taking a 

piece of paper and folding up its edges all around to the height of 

about a half inch. Half fill this on a cool surface with melted paraffine. 

Heat two dissection needles in bunsen flame and with these 

dispose pieces of material in orderly fashion over the crust which has 

by this time formed at the bottom of the tray. Blow upon the sur- 

face of the paraffine to harden it more quickly and as soon as the 

surface crust will bear it, plunge the tray into cold water. The 

material can now be left imbedded in paraffine until required for 

sectioning. 

If cutting is to be done in a cool room, softer grades of paraffine 

with 'melting points between 40 and 5oC. should be used for 

imbedding. If on the other hand cutting is to be done at summer 

temperatures, the harder grades melting at between 55 and 7oC. 

should be employed. 

TECHNIQUE OF SECTIONING AND MOUNTING MATERIAL IMBEDDED 

IN PARAFFINE 

Strip off the paper tray from the imbedded material and cut out a 

block of paraffine containing the object which is to be sectioned, 

taking care to include at least 2 or 3 mm. of paraffine on all sides 

beyond the specimen. Take a segment of pine wood about an inch 

long and with a surface at one end about % in. square and coat the 

square area with melted paraffine. Warm the paraffine on the piece 

of pine wood and quickly press the paraffine block containing the 

specimen into this melted paraffine in the desired position for cutting. 

Heat a dissecting needle and apply this all around the base so that 

the paraffine block is firmly sealed to the wood. Dip paraffine block 

in cold water to harden. Now trim the paraffine block with a sharp 

scalpel so that the faces form right angles with each other. Adjust 

the wood in the clamp of the microtome and the microtome blade 

so that the top of the paraffine block just touches the near surface 

of the microtome knife. Make certain that the knife edge and the 

two opposite faces of the paraffine block are perfectly parallel. 

Now 

trim the remaining two sides of the block close to the object. Adjust 

the automatic feed of the rotary microtome by moving dial to num-


her on scale representing thickness in microns desired of sections and 

turn wheel of microtome. It will be observed that the carrier moves 

up and down and with each downward movement slightly forward, 

causing the knife to cut sections which adhere in ribbons. 

Transfer the ribbons by means of a camel's hair pencil or dissecting 

needle to a piece of dust free paper with the side downward which was 

next to the knife. The ribbons are now ready to be mounted on 

slides. 

The slides to be used should only be those which are devoid of 

grease or dirt of any kind particularly on the surface upon which 

the ribbons are to be mounted. A very good plan is to keep a number 

of slides intended for this purpose submerged in a saturated solution 

of potassium dichromate in concentrated sulphuric acid. These 

can be taken out as needed and thoroughly rinsed with water. 

With a clean cloth stretched over the forefinger vigorously rub 

one surface of each slide until perfectly dry and free of lint. Then 

place a small drop of Mayer's albumin fixative on clean- surface and 

rub over the surface. (The formula for Mayer's Albumin Fixative 

Carbolic acid 

is as follows: Egg white and Glycerin, equal parts, 

i or 2 drops. Mix thoroughly.) Now flood the surface with water 

and cut the ribbons into segments of the desired length and arrange 

in rows on slide, being careful to have the segments somewhat 

shorter than the length of the cover slip because of tendency of 

parafrine to stretch when warmed. Warm slide gently by holding 

high above a bunsen flame or flame of an alcohol lamp until ribbons 

stretch out in smooth fashion. Absorb superflous water from beneath 

ribbons with blotting paper held to their edges and at same 

time push trie sections into even rows. Then leave the sections to 

dry for several hours or over night. 

METHOD FOR THE STAINING AND MOUNTING OF MATERIAL IN 

PARAFFINS RIBBONS AFFIXED TO SLIDE 

1. Gently heat the dry slides with parafrine ribbons adhering to 

the fixative, high above the Bunsen flame (with the ribbon side up). 

2. Place the slide upright in. a well of xylol or turpentine. The 

xylol or turpentine will dissolve the melted or two. 

parafrine in a minute


3. Take the slide out of the well, wipe off the under side and allow 

a stream of 95 per cent, alcohol to run over the upper side from a 

pipette. 

4. Place the slide upright in a well of safranin for from four to 

twenty-four hours. 

5. Take the slide out of the safranin well and extract excess of 

stain with 57 per cent, alcohol. 

6. Place the slide in a well of gentian violet or methyl-green for a 

second or more. The time varies for different objects and can only 

be determined by trial. 

7. Rinse slide with 70 per cent, alcohol from pipette. 

8. Pour absolute alcohol over sections, follow with a few drops of 

clove oil, replace clove oil with cedar oil. 

9. Mount in balsam. 


IMBEDDING IN CELLOIDIN 

Whenever material is unsuited for free hand sectioning and will 

not give good results when imbedded in paraffine on account of size, 

hardness, or brittleness, celloidin may be resorted to as an imbedding 

medium. 

The technique employed is similar to that of the paraffine method 

so far as the preliminary fixing, hardening and dehydrating are concerned 

up to and including the 95 per cent, alcohol stage. From this 

point the various succeeding steps in the procedure are as follows: 

1. Place material in equal parts of 95 per cent, alcohol and ether 

(known as ether-alcohol) for several hours. 

2. Transfer to a 2 per cent, solution of celloidin in ether-alcohol, 

for 2-5 days. 


for 2-5 days. 


for 3-10 days. 

5. Prepare a pine block sufficiently large in cross section to sup- 

in the 

port the material and otherwise adapted to its being clamped 

object carrier of the microtome. Soak one end of this block in


ether-alcohol for a while and then dip it in the 2 per cent, celloidin 

solution. 

6. Take the material from the thick celloidin and set it in proper 

position, for cutting the sections desired, on the prepared end of 

the block and allow the celloidin to thicken for a few seconds only. 

7. Dip the celloidin end into the thick solution; remove and hold 

upright so that the new coating may spread out over the end of 

block and solidify the union. 

the 

8. As soon as the celloidin has hardened a little to form a surface 

film, drop the preparation into a vessel of chloroform and allow to 

remain here i day. 

9. Transfer preparation to a vessel containing equal parts of 

glycerin and 95 per cent, alcohol until required for sectioning. 

SECTIONING CELLOIDIN MATERIAL 

Clamp the block in the sliding 

microtome and set the knife 

obliquely so that the sections can be cut with a long sliding stroke. 

Keep the knife and top of the block wet with the alcohol-glycerin 

mixture and as soon as the sections are cut, sweep them with a 

camels hair pencil into a dish of 70 per cent, alcohol. The sections 

can be attached to a slide by placing the slide in a closed chamber 

over ether. The ether vapor dissolves the celloidin and causes the 

sections to adhere to the slide. 

STAINING AND MOUNTING CELLOIDIN SECTIONS 

1. Place sections in safranin solution for i day. This safranin 

solution should be made by dissolving as much safranin in 95 per 

cent, alcohol as it will take up and then diluting with an equal 

quantity of water. 

2. Rinse sections in 50 per cent, alcohol to remove excess of stain. 

3. Transfer them to Delafield's haematoxylin (made by dissolving 

i gram of haematoxylin in 6 mils of absolute alcohol and adding 

this gradually to 100 mils of a saturated aqueous solution of ammonia 

alum. This is left exposed for a week, filtered, 25 

mils each of 

methyl alcohol and glycerin added, allowed to stand 6 hours, again 

filtered, and ripened about 2 months before using) for 10 minutes.

30 


3. Rinse sections thoroughly first in water, then in 35 per cent, 

alcohol, then in 50 per cent, alcohol. 

4. Put them quickly through acid alcohol (i drop of HC1 in 50 

mils of 70 per cent alcohol). 

5. Transfer to 70 per cent, alcohol for about 2 minutes. 



8. Transfer to absolute alcohol for about 2 minutes. 

9. Clear sections in a mixture of equal parts of cedar oil and phenol 

for at least 2 minutes. 

10. Remove excess of clearing solution and mount in balsam. 


DESILICIFICATION OF HARD WOODY MATERIALS 

It frequently happens, even after prolonged maceration or boiling 

in alkaline solutions, that thin sections of hard roots, stems, woods or 

fruits are difficult or impossible to procure. This is due to the pres- 

ence of deposits of silica and other mineral substances that usually 

occur in woody tissues. Therefore, it is of prime importance that 

these substances be removed as thoroughly as possible. 

For this 

purpose a 10 per cent, aqueous solution of commercial Hydrofluoric 

Acid (or stronger solutions up to the pure acid for very hard materials) 

is most useful. Small fruits or short segments of other hard 

materials are placed in this acid (which should be kept in a bottle 

coated internally with a thick layer of paraffine) for from 3 days to a 

week, depending on the size of the objects, with one or two changes 

of the acid. The acid is then washed out thoroughly with running 

water for 2 to 5 hours. This treatment completely frees the tissues 

of all mineral deposits without affecting the organic structure. 

SCHULZE'S MACERATION PROCESS 

This method is employed for the separation of cells. Radial- 

longitudinal sections, that may be cut with a pen knife, are placed in 

a beaker or test tube containing 50 mils of nitric acid of specific 

gravity 1.3 (about 2 volumes of nitric acid and i volume of water 

will serve purpose). To this add i gram of chlorate of potash 

crystals and heat gently until the reddish color which first appears


in the tissues has disappeared. Stop the action by pouring the 

whole 

' 

of the contents into a vessel containing water and wash 

well with water. The cells can now be readily separated with 

dissection needles and mounted in water for examination. Do not 

mount in glycerine, for it makes the already bleached elements too 

transparent. 

MICROMETRY 

The unit of length used in microscopic measurement is the micron 

(jit) which is one-thousandth part of a millimeter (o.ooi mm.) or one 

twenty-five thousandth part of an inch. 

In measuring microscopic objects it is necessary to make use of 

a micrometer of some kind. That pretty generally used is the 

ocular micrometer. It is a circle of glass suitable for insertion 

within the ocular with a scale etched on its surface. 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. 

STANDARDIZATION OF OCULAR MICROMETER 

The value of each division of the ocular micrometer scale must be 

ascertained for each optical combination (ocular, objective, and 

tube length) by the aid of a stage micrometer. 

The stage micrometer is a slide with a scale engraved on it divided 

to hundredths of a millimeter (o.oi mm.), in some cases to tenths 

of a millimeter (o.i mm.), every tenth line being made longer than 

intervening ones, to facilitate counting. 

METHOD : 

1. Insert the ocular 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 so that the 

Make 

lines of the two micrometers will appear in the same plane. 

the lines on the two micrometers parallel each other. This can 

often be done by turning the ocular to the right or left while looking 

into the microscope.

32 


3. Make two of the lines on the ocular micrometer coincide with 

two on the stage micrometer. Note the number of included divi- 

sions. 

4. Note the known value for each division of the stage micrometer 

scale which may either be etched on the stage micrometer or indi- 

cated on a label found pasted upon it. If the value indicated is o.oi 

mm. (Jfoo mm.) then each division of the stage micrometer scale 

has a value of 10 microns; if o.i mm. (Jfo mm.), 100 microns. 

5. Multiply'the number of included divisions of the stage microm- 

eter scale by the value in microns given for each division and divide 

the result by the number of included divisions of the ocular micrometer 

scale. The quotient represents the value of each division of 

the ocular micrometer scale. 

6. Note the optical combination (number of ocular, objective and 

tube length) used and keep a record of it with the calculated microm- 

eter value. Repeat for each of the combinations. 

To measure an object by this method read off the number of 

divisions it occupies of the ocular micrometer scale, and express the 

result in microns by looking up the recorded value for the optical 

combination used.

CHAPTER II 

LIFE HISTORY OF THE MALE FERN [DRYOPTERIS 

(ASPIDIUM OR NEPHRODIUM) FILIX-MAS] 

The Male Fern along with the Marginal Fern (Dryopteris margin- 

alis) have long been known to the pharmaceutical and medical professions 

as the source of the drug Aspidium, a most valuable remedy 

for the expulsion of tapeworm. The parts of these plants employed 

are the rhizome and stipes which are collected in autumn, freed of 

the roots and dead portions and dried at a temperature not exceeding 

7oC. 

HISTORY OF THE SPOROPHYTE OR ASEXUAL GENERATION 

Gross Structure of Stem. The main axis of Dryopteris Filix-mas 

is the creeping underground stem or rhizome which is oblique or 

ascending in habit. It gives off numerous roots from its lower and 

posterior portions and fronds from its upper and anterior portions. 

Behind the fronds of the present year are to be noted the persistent 

stalk bases of fronds of previous seasons. Lateral buds are frequently 

to be noted connected with these. The roots are slender and brown 

with semi-transparent apices. They are, inserted on the bases of 

the fronds, close to their junction with the stem. The growing end 

of the rhizome is called the anterior extremity and is marked by the 

presence of an apical bud overarched by young fronds. The opposite 

end is known as the posterior extremity and in the living plant is 

constantly decaying, as the anterior portion elongates. 

Histology of Mature Stem (Rhizome). Passing from periphery 

toward the center the following structures are to be observed : 

1. Epidermis, a protective outer covering tissue, composed of a 

single layer of brownish polyhedral cells from which are given off 

scaly hairs. 

2. Outer Cortex (hypodermis), a zone of several layers of thick- 

walled lignified cells separating the epidermis from the inner cortex 

beneath. Its main function is to support the epidermis. 

3 33

34 


3. Inner cortex of several layers of more or less isodiametric cells 

(cells of nearly the same length, breadth and thickness) with thin 

cellulose walls and containing stored starch surrounded by a proto- 

FIG. 12. Dryopteris filix-mai -Plant and section through pinnule and sorus. 

(Sayre.) 

plasmic investment. These cells conduct sap by osmosis and 

store food. Between the cells are" to be noted intercullular-air- 

spaces, many of which contain internal glandular hairs.

LIFE HISTORY OF THE MALE FERN 35 

4. Fundamental tissue, resembling the last in aspect and function. 

5. Vascular Bundles. These are of two kinds, viz.: stem bundles 

and leaf-trace bundles. Both are of elliptical outline, as seen in 

cross section, and are embedded in the parenchyma forming the 

broad central matrix. The stem bundles are comparatively broad 

and, as viewed in longitudinal sections, form a continuous network 

with good-sized meshes, each mesh being opposite the point of in- 

sertion of one of the leaves (See Fig. 13). 

In transverse section these bundles are 

seen to be usually ten in number and ar- 

ranged in an interrupted circle within the 

fundamental tissue. The leaf-trace bun- 

dles are comparatively narrow and are 

observed to come off of the stem-bundles 

and pass out through the cortex into the 

/leaves (fronds). When each bundle is 

examined under a high-power magnification 

it is seen to be composed of: (a) an 

endodermis or bundle sheath, a single layer 

of cells with yellowish walls and granular 

contents; (o) a per^cambûm or phloem 

FlG - I^~A 

B 

- Cylindrical 

network of vascular bundles 

in the stem of Dryopteris 

sheath of one to three layers of delicate Fiiix-mas. B. A portion of 

,, . n j n i i / \ the same more highly mag- 

thin-walled cells, rich in protoplasm; (c) a nified. At L are the interphloem, 

a broad zone stices 

of tissue formed of 

over which the leaves 

,,...., are inserted; at G are 

phloem cells, with thin cellulose walls and branches (leaf trace bunprotoplasmic 

contents, which convey sugar dles ) Passins into the leaves 

. . p from the main vascular 

involution irom the leaves to the roots and bundles. (Sayre.) 

broader sieve tubes which appear polygonal 

in transverse section and whose function is that of conveying soluble 

proteins in the same direction; (d) a xylem (wood) formed of thinwalled 

xylem cells which store food and scalariform tubes or tracheids 

which conduct crude sap (water with mineral salts in solution) from 

the roots to the leaves (fronds) . Since the xylem is surrounded by the 

phloem, the fibre-vascular bundle is of the concentric type. Strictly 

speaking the endodermis and pericambium are accessory regions, sur- 

rounding, but not part of the bundle proper.


Histology of Growing Apex. When the bases of the leaves of the 

current year, the circinate leaves of the following year and the large 

mass of brown scales have been removed from around the apical bud 

of a well-grown plant, tjie following structures may readily 

served with a hand lens : 

be ob- 

1. The apical cone (punctum vegetationis 4 

) , a rounded papilla, which 

occupies a terminal position in the apical region. 

2. The young fronds, arranged around the apical cone. 

Upon removing the extreme apex of the apical cone with a sharp 

razor, mounting in dilute glycerine or water and examining under 

low power, it will be noted 

FIG. 14. Apical cell of a fern rhizome in 

vertical longitudinal section. a.c., apical 

cell; h, hair; m, meristem. 

Sedgwick & Wilson's 

Henry Holt &" Co. 

(After Hofmeister.) 

General Biology, 

that a large pyramidal cell 

occupies 

the center of the 

apical cone. This is the 

apical cell (Fig. 14). The 

cells surrounding it have 

been derived by segmenta- 

tion (cell-division) from it, 

by means of walls parallel 

to its three sides; they are 

termed segment cells and in 

turn undergo 

further di- 

vision and redivision to . 

originate the entire stem 

tissue and leaf tissue. Step 

by step the tissue cells become modified into epidermal, cortical, 

bundle and fundamental cells. 

Histology pf Mature Root. Transverse sections cut some distance 

above the apex will present the following structures for 

examination : 

1. Epidermis, of epidermal cells whose outer walls are brown. 

Some of these cells have grown out as root hairs which surround soil 

particles and absorb water with mineral salts in solution. 

2. Cortex, of many layers of cortical parenchyma cells with brown 

walls. The outer layers of cells of this region are thin-walled, while 

the extreme inner ones are lignified and form a sclerenchymatous 

ring which surrounds the


3. Endodermis, a single layer of cells tangentially-elongated. 

4. Pericambium (Pericycle), usually of two layers of thin-walled 

cells containing protoplasm and large nuclei. This region surrounds 

the 

5. Radial fibro-vascular bundle, consisting of two phloem patches 

of phloem cells and sieve tubes on either side of two radial xylem 

arms of xylem cells, spiral tracheae and scalariform tubes. 

6. Lateral rootlets, which take origin in the pericambium. 

Histology of Root Apex. Microscopic examination shows this 

region to be composed of soft, pale, growing cells ending in the tri- 

angular apex-cell of the root. From the free base of the apex cell 

segment cells are cut off as calyptrogen cells. These by dividing 

form the root cap. The root cap or calyptra consists of a mass of 

loosely attached cells which forms a protective covering 

around the 

tip of the root. 

From the inner sloping sides of the apex cell the segment cells give 

origin to the dermatogen^ which by repeated division of its cells, 

the 

originates the epidermis (outer protective covering of the root) , 

periblem, originating cortex and the plerome originating the bundle 

and related tissue. 

Continuity of Crude Sap Flow. The crude sap (water with mineral 

salts in solution) penetrates the thin walls of the root hairs by osmosis 

and passes into the interior of hairs, thence into the root xylem and 

through this to stem xylem, thence through stem xylem 

leaves. 

into the 

, Histology of Stipe (Petiole). This, in transverse section, passing 

from periphery toward the center, presents the following structural 

characteristics: (see Fig. 15). 

1. Epidermis, a single layer of epidermal cells with dark brown 

outer walls. 

2 . Outer cortex (hypodermis) , a wide band of small cells with ligni- 

fied walls. 

3. Inner cortex, similar to inner cortex of stem but devoid of leaftrace 

bundle. 

4. Fundamental parenchyma, similar to same region of stem, in 

which are embedded a number of concentric fibro-vascular bundles 

arranged in an interrupted circle. Each of these shows a central


mass. Each bundle is 

xylem mass surrounded by an outer phloem 

enveloped by a pericambium and an endodermis or bundle sheath. 

Histology of Lamina. In transverse and surface sections the 

lamina or blade shows the following structural details: 

i. Upper epidermis, of wavy-walled, slightly chlorophylloid, flat 

upper epidermal cells, devoid of stomata, but with rather thick 

cuticle. 

FIG. 15. Transverse section of stipe of Dryopteris Filix-mas showing epidermis 

(e) ; hypodermis (h) ; inner cortex (ic) ; concentric fibrovascular bundles, 

one of which is shown at (&); endodermis (en); pericambium (p); xylem (#),< 

and phloem (p). (Photomicrograph.) X 50.4. 

2. Mesophyll, of irregular shaped chlorphylloid cells, containing 

abundant chloroplasts. Intercellular-air-spaces are found between 

various cells which are larger in the lower than in the upper region. 

Internal glandular-hairs are frequently to be discerned in many of 

these spaces. 

3. Concentric vascular bundles or laminar veins, that distribute 

sap to, and carry sap from the mesophyll. These are seen to be


embedded in the mesophyll. The xylem portion of each bundle is 

nearest to the upper surface of the leaf and so the bundles approach 

the collateral type. 

4. Endodermis, a continuous layer of mesophyll cells which surrounds 

each bundle and binds it in place. 

5. Lower epidermis of wavy-walled, flattened, chlorophylloid cells 

with thin cuticle and many stomata (breathing pores). Each stoma 

is surrounded by a pair of crescent-shaped guard cells which regulate 

its opening and closing. The upper and lower epidermis are con- 

tinuous around the laminar margin. 

Comparative Physiology of Root, Stem and Leaf (Frond). The 

primary function of the roots of the Male Fern is that of absorption 

of water with mineral salts in solution. The secondary function 

is that of support for the stem, the tertiary, that of storing food- 

stuffs to tide the plant over the season when vegetative activities 

are lessened. Water is the most essential of all materials absorbed 

by vegetable organisms. It is found in the soil surrounding the 

soil particles with certain mineral salts dissolved in it. The delicate 

root-hairs with thin cellulose walls, protoplasmic lining and sap 

denser than the soil water, are firmly adherent to these particles. 

The soil water diffuses through these walls by osmosis and comes into 

relation with the ectoplasm, a delicate protoplasmic membrane, 

which has the power of selecting what it wants and rejecting what it 

does not need. In this way only such solutes as are of value to the 

plant are admitted.' The water with mineral salts in solution, once 

within the root-hair protoplast, is called "crude sap." This passes 

cells which are in 

through the hair into the cortical parenchyma 

contact with the spiral ducts and scalariform tracheids. It passes 

from one cortex cell to another by osmosis and, under considerable 

root pressure, is forced into the spiral and scalariform tubes of the 

xylem. Therein it is conveyed upward by root pressure through the 

tracheids of the stem bundles into those of the leaves and finally 

osmoses into the leaf parenchyma cells (mesophyll) . 

Carbon dioxide, (CO 2), from the air, enters the leaf through the 

stomata. From the stomata it moves through the intercellular-air- 

spaces to the mesophyll cells which line these, whence it is absorbed . 

Within the mesophyll ceUs are found small chloroplasts composed

40 

of protoplasm and chlorophyll. 


The kinetic energy of the sun's rays 

is absorbed by the chlorophyll which j's thus energized to break up 

the CO 2 and H 2O into their component elements C,H and O, and 

rearrange them in such a way as to ultimately form sugar or starch. 

This process is called photosynthesis. According to von Baeyer, 

CO 2 is split into C and O 2 , the C being retained, the O 2 given off. 

The nascent C is linked with H 2O to form CH 2O (formic-aldehyde) ; 

six molecules of this are then united to form grape sugar (C 6Hi 2O 6 ). 

The formation of starch may be expressed by the following equation : 

6C0 2 + sH 2 = C 6Hi O 5 + 6O 2 . A portion of the grape sugar is 

removed from solution by the chloroplast and converted into starch 

which is stored up within it; another portion is used to nourish the 

protoplasm of the cell. But the greater portion of sugar manufactured 

descends in solution through the phloem cells of the bundles 

of the veins, mid-rib and stipe to the stem or roots, where it is removed 

from solution by the action of the leucoplasts which convert 

it into reserve starch. Sugar and starch, however, are not the only 

food materials manufactured in the leaf. Proteins are likewise 

formed. These are composed of carbon, hydrogen, oxygen, nitrogen, 

sulphur and sometimes phosphorus. They are formed from grape 

sugar with the addition of nitrogen and the other elements by the 

living protoplasm. The source of nitrogen, sulphur and phosphorus 

is the mineral salts which are found in the crude sap. These 

proteins descend through the sieve tubes of the veins, midrib and 

petiole to the stem and roots, nourishing all of these parts with protein 

material. 

Gross Structure and Histology of the Sori and Sporangia. The 

sporangia or spore cases are found clustered together in circular 

groups on the under surface of the pinnules nearer the mid-vein than 

the margin. Each group of sporangia is covered with a membranous 

expansion of the epidermis called the indusium. The whole is called 

a sorus (Fig. 12) (pi. sori) and contains many sporangia. Each 

sporangium is composed of: (a) the stalk of considerable length and 

usually comprising three rows of cells, outgrowths of the epidermis of 

the pinnule; and (b) the head, sub-globular and hollow, consisting for 

the most part of a covering of thin walled, flattened cells, within


which will be noted a marginal ring of cells, with walls having 

U-shaped thickenings, and called the annulus. 

Within the sporangium are found the spores. Each spore is a 

single cell composed of an outer brown wall with band-like markings 

called an exosporium, an inner thinner wall or endosporium, and with- 

in this a mass of protoplasm containing a nucleus. 

Rupture of Sporangium and Spore Dissemination. As was previously 

indicated, each sporangial head has a row of cells with 

U-shaped thickenings around the margin called an annulus. As the 

sporangium matures the water escapes from the cells, pulling them 

together and holding the annulus like a bent spring. The thinner 

PIG. 16. Sporangia of an undetermined species of fern; li, lip-cells; an, annulus; 

st, stalk; sp, mature spores. Each of the four nuclei in the upper cells of the 

stalk is in the terminal cell of one of the four rows of cells that compose the stalk. 

(Gager.) 

walled cells at the side of the spore case opposite the annulus, unable 

to stand the strain, are consequently torn; the annulus then 

straightens and a wide rent is made in the sporangium. The annulus 

then recoils and hurls the spores out of the sporangium. This 

closes the sporophyte generation. 

History of the Gametophyte or Sexual Generation. The fern 

spore, falling upon a moist surface, germinates, producing a delicate 

green septate filament called a protonema. One end of this structure 

shows larger cells, which, by the formation of oblique walls, cut out 

an apical cell of somewhat triangular shape. This is the growing 

point of what eventually becomes a mature, green, heart-shaped 

body called the "prothallium" or "prothallus." The prothallium, 

about the size of an infant's finger nail, develops on its under surface


PIG. 17. A, under surface of a fern prothallium showing archegonia (/)' 

antheridia (m) and rhizoids (r); B, immature archegonium showing binucleate 

neck canal cell (n.c.c.), ventral canal cell (v.c.c.), and egg (e); C, mature archegonium 

showing sperms (sp.) moving through neck canal (n.c.}' toward ovum 

(e); and venter (). All highly magnified.


antheridia, or male sexual organs, archegonia, or female sexual organs, 

and rhizoids or hair-like absorptive structures. The antheridia 

appear three to five weeks after spore germination. They are hemispherical 

in shape and are situated among the rhizoids toward the 

posterior end. Each antheridium consists of a three-celled wall 

which completely surrounds the spermatocytes or mother-cells of the 

spermatozoids. Within each spermatocyte the protoplasm arranges 

itself in a sphal fashion forming a spermatozoid, a spiral, many 

ciliated, male sexual cell. From two to four weeks after the matura- 

tion of the antheridia, the archegonia make their appearance toward 

B 

an: 

FIG. 18. A, median longitudinal section through immature antheridium, and 

cell of prothallium showing prothallial cell (p), and antheridial wall surrounding a 

number of spermatocytes; B, similar section through mature antheridium and cell 

of prothallium showing fully developed spermatozoids (sp.} enclosed by wall of 

antheridium. Both highly magnified. 

the indented apex of the lower prothallial surface as outgrowths of 

the prothallial cushion. Since they appear later than the antheridia 

they are not likely to be fertilized by spermatozoids from the antheridia 

of the same prothallium. Each archegonium is composed of a 

venter, neck, neck canal-cells, "ventral canal-cell, and ovum or egg-cell. 

The neck is composed of cells arranged in four rows, forming a cylin- 

der, one layer of cells thick. This protrudes from the surface of 

the prothallium and encloses the neck canal-cells and ventral canal- 

cell. The ovum is embedded in the prothallial cushion just beneath 

the ventral canal-cell. Upon the maturation of the archegonium, 

the canal cells are transformed into a mucilaginous substance which 

fills a canal extending from the outside opening (mouth )to the ovum.

44 . PHARMACEUTICAL BOTANY 

During wet weather the mature antheridial wall bursts open and 

the many ciliated spermatozoids escape into the water. These' moving 

in the water are drawn by the chemotactic malic acid to the 

mouths of the archegonia of another prothallus, and, passing down 

the canal of each of these, gather around the ovum. One, probably 

the best adapted, fuses with the ovum and fertilizes it forming an 

oospore or fertilized egg. 

Origin of New Sporophyte or Diploid Plant from Fertilized Egg. 

The fertilized egg now rapidly divides and redivides to form octant 

cells. The octant cells further divide to produce anteriorly a stem 

rudiment (one cell), first leaf (two cells), second leaf (one cell) and, 

posteriorly, root rudiment (one cell), foot rudiment (two cells) and 

hair rudiments (one cell) . 

Growth of Seedling into Mature Sporophyte. The foot rudiment 

develops into the foot which obtains nourishment from the prothal- 

The 

lium, upon which the young sporophyte is for a time parasitic. 

root rudiment becomes the first root which grows downward into 

the -soil. The stem and leaves turn upward. In a few weeks the 

prothallus decays and the sporophyte is established as an independent 

plant. More roots and leaves (fronds) are developed and ere 

long continued growth results in the formation of a mature sporophyte 

which presents for examination: (i) a subterranean stem bear- 

ing several roots; and (2) arid fronds, each of which consists of a 

stipe or petwle and a lamina or blade, the latter divisible into pinna 

or lobes and pinnules, upon which last sori are developed. 

Alternation of Generations. It will be observed that in the life 

cycle of the Male Fern there occur two distinct generations, one, a 

sporophyte or asexual generation which begins with the oospore and 

ends with the dispersion of asexual spores; a second, the gametophyte 

or sexual generation, beginning with the protonemal outgrowth of 

the spore and ending with the fertilization of the egg to form an 

oospore. The sporophyte gives rise to the gametophyte which in 

turn gives origin to the sporophyte.

CHAPTER III 

LIFE HISTORY OF A GYMNOSPERM (PINUS STROBUS) 

The White Pine frequently called the Weymouth Pine (Pinus 

Strobus), one of the principal 

timber trees of the Northern States 

PIG. 19. Transverse section of white pine stem ot four years' growth, showing 

cork (a), cortex (&), phloem (c), cambium (d), xylem (e), secretion reservoir (/), 

pith (g) and medullary ray (&). (Photomicrograph.) X34O. 

and Canada, is also of value in pharmacy and medicine. The inner 

bark of its trunk and branches is used because of its valuable ex- 

pectorant properties and is official in the N. F. IV. 

45


DESCRIPTION OF THE WHITE PINE TREE (MATURE 

SPOROPHYTE) 

From an underground spreading root system there arises an erect 

aerial trunk or stem that extends from the ground to the apex of the 

tree, ending in a terminal bud. The trunk rarely exceeds 3 feet in 

diameter and 125 feet in height and is averagely ij^ to 3 feet in 

PIG. 20. Transverse section of white pine needle (leaf) showing epidermis (a), 

infolded parenchyma cells of mesophyll (b, &') oil reservoir (c), endodermis (d), 

clear cellular area (e) surrounding fibrovascular tissue in center (/). X4OO. 

diameter and 50 to 90 feet high. 

At a varying distance above the 

soil, depending upon environal conditions as well as the age of plant, 

whorls of lateral branches (three to seven in a whorl) are seen emanating 

from the trunk in horizontal fashion at various levels up to 

near the apex. These become, under conditions prevalent when the 

tree is grown in the open, gradually shorter until the summit is

LIFE HISTORY OF A GYMNOSPERM 47 

reached, giving to the crown or upper part of the tree the appearance 

of a pyramid. These branches give rise to other branches which 

agree with the lateral branches in bearing, commonly, only scale like 

leaves as well as in ending in terminal buds. Another kind of 

branch, however, is found which is always shorter than the scaly 

branches. This type of branch is called a "spur shoot" and arises 

FIG. 21. Staminate cones of the Austrian pine (Pinus austriaca) . Below, before 

shedding pollen; above, after shedding. (Gagerj 

from the former branches. The spur shoots bear the needles or 

foliage leaves which are light-green, when young, and bluish-green, 

soft, flexible, 2% to 5 inches long, when mature. The "needles" 

occur in tufts (fascicles) of five, are triangular in cross-section, have 

finely serrate (saw-toothed) edges and are surrounded at the base by a 

deciduous sheath. These foliage leaves persist until the end of their 

second year, when they are shed with the spur shoot which bears 

them. 

The white pine, like most of its allies among the Coniferae, bears 

cones. These structures are of two kinds, viz. : staminate and car- 

pellate. 

Both kinds are produced on the same tree.


Staminate Cones. The yellow, ovate, staminate cones appear 

about May and are clustered at the base of the new growth of the 

current season. Each consists of a main axis (modified branch) 

which bears spirals of scales (microsporophylls or stamens). On the 

PIG. 22. Scotch pine (Pinus sylvestris}. A-D, stages in the development of the 

carpellate cone, and its carpotropic movements. E, very young carpellate cone 

much enlarged; F, ventral,. G, dorsal views of a scale from E; i, ovuliferous scale; 

2, ovule (in longitudinal section) ; 3, pollen chamber and micropyle leading to 

the apex of the nucellus {megasporangium) ; 4, integument of the ovule; G, i, tip 

of ovuliferous scale; 5, bract; 4, integument; H, longitudinal section at right 

angles to the surface of the ovuliferous scale (diagrammatic); 6, megaspore; 7, 

pollen chamber; /, longitudinal section of a mature cone; 6, ovule; J, scale from 

a mature cone; 6, seed; w, wing of seed; K, dissection of mature seed; h, hard seed 

coat; c, dry membraneous remains of the nucellus, here folded back to show the 

endosperm and embryo; e, embryo; p, remains of nucellus; L, embryo; c, cotyledons; 

e, hypocotyl; r, root-end. (Gager.) 

under surface of each scale are the spore-cases (microsporangia), 

which develop the micros pores (pollen grains). Each pollen grain 

when mature consists of a central fertile cell and a pair of air-sacs 

or wings, one on either side of the fertile cell. The purpose of the 

latter is to give greater buoyancy in the air to the microspore. 

Carpellate Cones. The young carpellate cones appear in May or 

early June as pinkish-purple structures arranged in solitary, fashion

LIFE HISTORY OF A GYMNOSPERM 49 

or in small groups, "lateral along the new growth. Each terminates 

a lateral axillary branch. A carpellate cone is composed of a main 

axis which bears spirals of scales, by some termed megasporophylls 

(carpels). Each scale is composed of an ovuliferous scale bearing 

two ovules or megasori and a bract. Each megasorus contains a 

FIG. 23. Mature carpellate cones of white pine showing separated scales. 

nucellus or megasporangium which is surrounded by an integument, 

except at the apex where an opening, the micropyle is evident. The 

micropyle is the gateway to the pollen chamber which lies below it. 

Within the nucellus occurs a megaspore or embryo sac. 

DESCRIPTION OF THE GAMETOPHYTE GENERATION 

The Gametophyte generation of the White Pine begins with the 

development of the male and female gametophytes and terminates 

with the fertilization of the egg. 

The Male Gametophyte. The male gametophyte commences to 

form in the ^nature pollen grain before the pollen is shed. A series 

4

50 


of three nuclear divisions takes place which cut off two small prothallial 

cells (traces of one of which may be seen pushed up against the 

wall of the fertile cell of the pollen grain), a tube nucleus and a 

generative cell. At this stage the pollen is shed and some of it is 

carried by air currents to the carpellate cones where it sifts in be- 

tween the ovule-bearing scales and accumulates at the scale bases. 

A number of the pollen grains are drawn close to the nucellus of the 

ovule by the drying up of the viscid fluid which fills the pollen 

chamber. In this fluid they germinate forming pollen tubes. The 

PIG. 24. The white pine (Pinus Strobus). Sections through mature pollen 

grains; at the left the remnants of two prothallial cells can be seen, while at the 

right all signs- of the first cell have disappeared. Pollen collected June 9, 1898. 

X about 600. (Gager, after Margaret C. Ferguson.) 

transfer of pollen grains from the pollen sac to the pollen chamber 

and consequent germination therein is called pollination. The con- 

tents of a mature pollen-grain constitutes the male gametophyte. 

The Female Gametophyte. If the embryo sac be examined at 

about the time of pollination, it will be found to consist of a single 

cell containing a single nucleus surrounded by cytoplasm. Very 

shortly afterward, however, the nucleus divides repeatedly to form a 

large number of nuclei which are scattered throughout the cytoplasm. 

Each nucleus accumulates around itself a portion of the 

cytoplasm and ultimately cell walls are laid down and the entire 

embryo sac contains endosperm (prothallial) tissue. Toward the 

micropylar end of the endosperm (prothallus) originate several 

archegonia. 

Each archegonium consists of a much-reduced neck of four cells 

and an egg (ovum) which lies embedded in the prothallus which 

forms a narrow layer of cells around it called the jacket. The con- 

tents of the mature embryo-sac constitutes the female gametophyte.

LIFE HISTORY OF A GYMNOSPERM 

PIG. 25. White pine (Pinus Sttobus). At left, megasporangium with megaspore 

in the center; above, pollen grains in the micropyle and pollen chamber. At 

right, pollen grains beginning to germinate; the cells of the integument have 

X enlarged and closed the micropyl< . (Gager, after Margaret C. Ferguson.) 

FIG. 26. White pine (Pinus Strobus). Vertical section through the upper part 

of an ovule, shortly before fertilization, s.n., sperm-nuclei; st. c., stalk-cell; t.n. 

tube-nucleus; arch, archegonium; e.n., egg-nucleus. (Gager, after Margaret C. 

Ferguson.) 

j 

e.n

52 


Fertilization. About a year after pollination the pollen tubes, 

lying within the pollen chamber show signs of renewed activity. 

The tube nucleus passes to the tip of the tube. The generative-cell 

divides to form a body and a stalk-cell which pass into the tube. 

The body-cell later forms two sperm nuclei. While these changes are 

taking place the tube is penetrating the nucellus and growing toward 

the embryo sac with its contained female gametophyte. It finally 

enters it, passing between the neck-cells of the archegonium. The 

tip of the tube then breaks and the entire tube contents is emptied 

into the egg. One of the sperm nuclei fuses with the egg nucleus 

and fertilizes it forming an oospore. 

Seed Formation and Distribution. The oopsore undergoes re- 

peated divisions and forms the embryo or young sporophyte plant 

and a suspensor to which it is attached. The embryo is nourished 

by a portion of the prothallus but the greater part of the prothallus 

forms the endosperm tissue of the seed surrounding the embryo. 

The thin nucellus persists as an endosperm covering. The integument 

becomes modified to form the hard protective seed coat. A 

portion of the scale of the cone directly above and adjacent to the 

ovule forms a membranous wing which separates from the scale as 

part of the seed. 

By this time (about two years after pollination) the scales of the 

cone, now quite woody, separate, the seeds are shaken out, and many 

are carried for a considerable distance by winds. 

Germination of the Seed. Under favorable conditions, the seeds 

absorb water and germinate in the spring following their dispersal. 

The hypocotyl of the embryo appears first, arching upward and 

downward, and, straightening out, draws the green cotyledons with 

it which spread out toward the light, while it grows into the soil to 

form the tap root and in time the remainder of the root system. 

Thus the seedling sporophyte is formed which in time develops into 

the mature White Pine tree.

CHAPTER IV 

LIFE HISTORY OF AN ANGIOSPERM (ERYTHRONIU1VL 

AMERICANUM) 

This attractive little plant, commonly called the Dog's Tooth . 

Violet but related to the Lily, is found in the hollows of woods and 

may be seen in flower during the month of April 

in the Middle 

FIG. 27. Dog's-tooth violet . (Erythronium americanum) Stages of 

development from the seed. 15 show the stage of development in each of five 

successive years k Pull explanation in the text. 6, Bulb showing a surface bud 

(the sprout has been destroyed). (Gager After F. H. Blodgett.) 

Atlantic States. It consists of an underground stem bearing scales 

(modified leaves) which is termed a bulb. From the lower surface 

of the bulb are given off numerous slender rootlets which penetrate 

the soil and from the upper surface, a pair of oblong lance-shaped 

53

54 


leaves of pale green color mottled with purple and white, and later, a 

flower stalk (scape), which bears upon its summit a single yellow, 

nodding flower, which is often marked with purple stripes. The 

flower consists of a torus or receptacle which will be observed as the 

upper swollen end of the flower stalk (scape). Inserted upon it, 

will be noted four whorls 

passing from periphery toward the center, 

of floral leaves which, in order, are calyx, corolla, 

gyncecium. The calyx is composed of three lance-shaped 

andrcecium and 

and re- 

curved yellow parts called sepals] the corolla of three similarly 

looking parts called petals which alternate in position with the sepals. 

Both of these whorls are collectively called the perianth or floral 

envelope. The androecium or male system of organs is composed 

of two whorls or circles of structures called microsporophylls or 

stamens. There are three stamens in each whorl. The outer whorl 

of stamens will be found opposite the sepals while the inner will be 

observed opposite the petals. Each stamen (microsporophyll) consists 

of an awl-shaped stalk or filament bearing upon its summit an 

oblong-linear body called an anther. The anther consists of two 

lobes called microsori. Each lobe or microsorus contains two anther 

sacs or micros porangia in which when mature are to be found micro- 

spores or pollen grains. In the center of ihe flower will be noted 

the gynoecium or female system of organs. This, upon dissection, 

will be found to consist of three fused carpellary leaves termed megasporophylls 

(carpels) forming a somewhat flask-shaped structure 

called a pistil-. The swollen basal portion of the pistil is called the 

ovary; the stalk which arises from it is called the style and the knob- 

like viscid summit of the style is termed the stigma. 

Microscopical examination of sections of the ovary will reveal it 

to be composed of three chambers called locules, within each of which 

are to be noted several inverted ovules. Each of these ovules is 

developed upon a nourishing tissue termed "placenta" which connects 

the ovules to the inner angle of the wall of the locule. The 

ovule is composed of a central prominent megasporangium or nucellus 

which is almost completely invested by two upgrown integuments 

or coverings. 

The opening between the tips of the inner integument 

is called the micropyle (little gate). This is the gateway for the 

entrance of the pollen tube on its way to the nucellus. It is also

LIFE HISTORY OF AN ANGIOSPERM 55 

the exit door for the hypocotyl of the embryo 

and ripened ovule becomes a seed. Within the nucellus, 

after the fertilized 

if the sec- 

tions examined have been properly fixed, will be found a megaspore 

or embryo sac. 

Development of the Female Gametophyte Through the Maturation 

of the Embryo Sac. In its immature condition the embryo 

sac (megaspore) contains a mass of protoplasm surrounding a nucleus. 

This nucleus undergoes three divisions forming as a result eight 

nuclei which ultimately arrange themselves within the protoplasm 

of the embryo sac as follows: three of them occupy a position at the 

apex, the lower nucleus of the group being that of the egg or ovum, the 

other two nuclei being the synergids or assisting nuclei; at the oppo- 

site end of the sac three nuclei known as antipodals take their posi- 

tion; the two remaining nuclei called polar nuclei take up a position 

near the center of the embryo sac. In this condition the contents 

of the embryo sac constitutes the female gametophyte. See Fig. 

28 (1-8). 

MATURATION OF THE POLLEN GRAIN AND FORMATION OF THE 

MALE GAMETOPHYTE 

The pollen grains (microspores), within the anther sacs, all arise 

from a number of tetrads (groups of four) which are formed by the 

division and redivision of pollen mother-cells preceeding them. Each 

pollen grain, after the tetrads have separated into their components, 

consists of an outer firm wall or exosporium, an inner wall or endo- 

sporium, within which will be found the. region called the fovilla, 

which is nothing other than a mass of protoplasm containing a 

nucleus. Before the pollen is shed from the anther its protoplasmic 

contents undergo a series of changes leading up to the development 

of the male gametophyte. The nucleus and protoplasm enveloping 

it divides to form two cells, one a generative-cell containing a generative 

nucleus, the other a tube-cell containing a tube nucleus. The 

generative nucleus then divides to form two sperm nuclei and the 

partition wall between the two cells disappears. In this condition 

the protoplasmic contents of the pollen grain constitute the male 

gametophyte.

56 


POLLINATION AND FERTILIZATION 

The mature pollen grains are discharged from the ripened anther 

through the splitting open of its wall. They are transferred to the 

stigma of the pistil of another Erythronium flower through the 

agency of insects. Here they germinate, each putting forth a tube 

FIG. 28. At the left, diagram of the anatomy of an angiospermous flower 

shortly after pollination; anth., anther; fil., filament; St., stamen; stig., stigma; 

p.g., pollen grains germinating; sty., style; pt., pollen tube; o.w., ovary wall; o. 

ovule, containing embryo-sac; pet., petal; sep;, sepal. 1-8, Stages in the development 

of the female gametophyte (embryo-sac); meg.sp., megaspore-mothercell; 

i.i., inner integument; o.i., outer integument ',f.un., funiculus; chal., chalaza; 

nu., nucellus (megasporangium) ; emb., embryo-sac. All diagrammatic. (Gager.) 

(pollen tube). The pollen tubes, carrying within it two sperm 

nuclei and a tube nucleus embedded in protoplasm, penetrate 

through the style canal until they reach the micropyles of various 

ovules. Each enters and passes through a micropyle, then piercing 

the nucellus, grows toward the embryo sac. The tip of the tube 

fuses with the end of the embryo sac and the two sperm nuclei are 

discharged into the sac. One of these sperm nuclei passes between


the synergids and fuses with the nucleus of the egg to form an 

oospore. By this time the tube nucleus has disintegrated. The 

oospore by repeated divisions develops into as many as four embryos 

or young sporophyte plants. Only one of these, however persists. 

The polar nuclei fuse to form the endosperm nucleus which soon 

undergoes rapid division into a large number of nuclei scattered 

about through the protoplasm of the embryo sac. Later cell walls 

are laid down and endosperm is formed. The endosperm cells soon 

become filled with abundant starch which is later to be utilized by 

the embryo during germination. 

RIPENING OF THE OVULE TO FORM THE SEED AND OF THE OVARY 

TO FORM THE FRUIT 

When the embryo and endosperm are being formed, 

the ovule 

enlarges and its integuments become modified to form a hard horny 

seed coat which encloses the endosperm surrounding the embryo. 

The ovary, containing the ovules, has by this time ripened to form 

a three-valved loculicidal capsule enclosing the seeds. 

GERMINATION OF THE SEED AND DEVELOPMENT OF THE 

MATURE SPOROPHYTE 

The seeds are fully developed by June or July when the capsule 

or fruit splits open to discharge them. They fall to the ground and 

lie dormant until the following spring when they germinate or com- 

mence to grow. Each seed absorbs water from the ground which 

stimulates the ferment amylase, contained in the endosperm cells, to 

break up the insoluble starch into soluble sugar which passes into 

solution and diffuses into the cells of the embryo, where the protoplasm 

changes it into additional protoplasm and so the embryo 

increases in size, therefore, grows. The pressure of the swollen 

endosperm and growing embryo becomes so great that the seed coat 

bursts; the hypocotyl emerges first, dragging the cylindrical cotyledon 

out of the seed coat and epicotyl with it. The hypocotyl elon- 

gates and extends itself into the soil where it develops a root near 

its tip. The tip enlarges through the storage of starch, manufac- 

tured by the green cotyledon and becomes a bulb. The bulb soon

58 


develops within it a plumule, the cotyledon withers, and the young 

plant (seedling) passes the following winter in this condition. 

During the next spring the plumule develops into a foliage leaf and 

the bulb gives rise from its base to several slender elongated runners, 

which, at their tips develop runner bulbs. These runner bulbs, the 

third year, give origin to another set of runners similar to those 

formed during the second year which also develop runner bulbs at 

their tips. A foliage leaf is also formed by each. The following 

spring (spring of fifth year) one of these bulbs develops into a 

mature sporophyte plant, bearing a single flower at the summit of 

its elongated scape. See Fig. 27. 

RESEMBLANCES BETWEEN GYMNOSPERMS AND ANGIOSPERMS 

1. In both are developed those structures in which there is no 

homologue, e.g., flowers. 

2. In both the flowers develop at least two sets of leaves (either 

on one or two plants of the same species) called sporophylla or 

sporophyll leaves, the stamens and carpels. The stamens or stamina! 

leaves are also termed microsporophylls. The carpels or 

carpellate leaves are also known as megasporophylls. 

3. Both groups produce microspores or pollen grains and mega- 

spores or embryo sacs. 

4. In both are developed on the evident generation, the plant or 

sporophyte and the gametophyte, the latter concealed within the 

megaspore of the sporophyte. 

5. Both develop seeds with one or two seed coats. 

6. In both groups there is developed from the fertilized egg an 

embryo which lies within the cavity of the megaspore. 

7. In both there exists a root and a stem pericambium. 

8. Both produce collateral vascular bundles. Very rarely 

do we 

meet with concentric bundles in the stem or leaf of Angiosperms. 

FUNDAMENTAL DIFFERENCES BETWEEN GYMNOSPERMS AND 

ANGIOSPERMS 

i. The flowers of Gymnosperms are often monoecious or dioecious 

but very rarely hermaphrodite, as in Welwitchia, whereas those of 

Angiosperms are usually hermaphrodite, rather rarely monoecious, 

still more rarely dioecious.


2. In the Gymnosperms the sporophylls are usually inserted 

either spirally or in whorls around a distinctly elongated axis, 

whereas in Angiosperms the sprophylls are condensed to short 

whorls or spirals set around a shortened axis, the floral axis or recep- 

tacle, torus or thalamus, or, as in the more modified Angiosperms, 

the floral axis may even become hollow. 

3. In Gymnosperms the microsporophylls or stamens are usually 

sessile, whereas in Angiosperms the microsporophylls are nearly 

always stalked. Rarely do we find sessile anthers among Angiosperms, 

an instance of this being seen in Mistletoe (Viscum) where 

the anthers are set on the staminal leaf. 

4. In Gymnosperms there is a traceable prothallus or gametophyte 

plant that later becomes the so-called "endosperm" of the gymnosperm, 

whereas in Angiosperms no recognizable prothallus has been 

proven to exist. 

5. The stored food tissue in Gymnosperm seeds is prothallial tissue 

loaded with starch, etc., whereas in Angiosperm seeds the stored 

food tissue (endosperm) is a special formation after fertilization. 

6. Gymnosperms bear naked ovules and seeds while Angiosperms 

bear covered ones. 

7. In Gymnosperms there are distinct recognizable archegonia 

formed on or imbedded in the prothallus, whereas in Angiosperms 

there are no distinct archegonia, only an isolated egg or eggs. 

8. In Gymnosperms there are not infrequently found several 

embryos from one fertilized egg. This condition is called poly- 

embryony. Polyembryony is unknown in Angiosperms, only a false 

polyembryony being noticed. 

9. In Gymnosperms the secondary xylem (wood) tissue of roots, 

stems and leaves consists either of punctated or scalariform cells, 

whereas in Angiosperms the secondary wood tissue may be varied 

in structural aspect.

CHAPTER V 


Vegetable Cytology treats of plant cells and their contents. 

THE PLANT CELL AS THE FUNDAMENTAL UNIT 

Schleiden, in 1838, showed the cell to be the unit of plant structure. 

The bodies of all plants are composed of one or more of these fundamental 

units. Each cell consists of a mass of protoplasm which 

may or may not have a cell wall surrounding it. While most plant 

cells contain a nucleus and some contain a number of nuclei, the 

cells of the blue-green algae and most of the bacteria have been found 

to lack definitely organized structures of this kind but rather con- 

tain chromatin within their protoplasm in a more or less diffuse 

or loosely aggregated condition. 

A TYPICAL PLANT CELL 

If we peel off a portion of the thin colorless skin or epidermis from 

the inner concave surface of an onion bulb scale, mount in water and 

examine under the microscope, we find it to be composed of a large 

number of similar cells which are separated from one another by 

means of lines, the bounding cell walls. Under high power each of 

these cells will exhibit the following characteristics: 

An outer wall, highly refractile in nature and composed of cellulose; 

which surrounds the living matter or protoplasm (See Fig. 29). This 

wall is not living itself but is formed by the living matter of the cell. 

Somewhere within the protoplasm will be noted a denser-looking 

body. This is the nucleus. Within the nucleus will be seen one or 

more smaller highly refractile and definitely circumscribed bodies, 

the nucleolus or nucleoli. The protoplasm of the cell outside of the 

nucleus is called the "cytoplasm." It will be seen to be clear and 

60


granular-looking. Within the cytoplasm will be observed a number 

of clear spaces. These are vacuoles and because they are filled with 

cell sap (water with nutrient substances in solution) are called "sap 

vacuoles." 

Protoplasm 

cytoplasm to a bounding film of water between it and the cell wall 

forms the outer plasma membrane or ectoplasm, a clear homogeneous 

outer band of cytoplasm; the reaction of cytoplasm to the water 

is in intimate relation to water. The reaction of the 

cpLnn rf cy 

mi' 

FIG. 29. Portion of inner epidermis of Onion bulb scale showing cells at 

various stages of maturity. Young cell (i); old cell (3); cell intermediate in age 

between I and 3 (2); cell wall (c); outer plasma membrane (pi)', middle lamella 

(ml)\ nucleus (w 1 

); nucleolus (w 2 ); nuclear membrane (w 3 ); cytoplasm (cy); 

vacuole (z;). Note that the young cell (i) shows numerous small vacuoles and 

spheroidal nucleus near center of cytoplasm. In 2 (cell of intermediate age) the 

cell has enlarged, larger vacuoles have formed thru the bursting of films of cytoplasm 

separating smaller ones, and the nucleus has moved toward the cell wall. 

In 3, the films have all burst, the cytoplasm and nucleus have been pushed up 

against the cell wall, the nucleus is flattened out, and a large vacuole appears in 

the center of the cell. 

within the sap vacuoles forms the vacuolar membranes; the reaction 

of the dense protoplasm of the nucleus to the water in the cytoplasm 

around it forms the nuclear membrane. Upon mounting another 

portion of epidermis in iodine solution, removing the excess of stain 

and adding a drop of sulphuric acid and then examining under high 

power, we note that the cell walls of cellulose are stained a deep blue. 

A yellow line is evident in the middle of each cell wall and separates 

each cell from its bounding cells. This line is the middle lamella 

which is composed largely of calcium pectate. 

6l


PROTOPLASM AND ITS PROPERTIES 

Protoplasm, or living matter, is the more or less semi-fluid, viscid, 

foamy, and granular substance in which life resides. It is the 

" physical basis of life." 

The peculiar properties which distinguish protoplasm from non- 

living matter are as follows: 

1 . Structure. Protoplasm invariably exhibits structure. No portion 

of it, however small, has been found to be homogeneous. Each 

The 

advance in microscopical technique reveals new complexities. 

protoplasm of a single cell, far from being a single unit, must rather 

be looked upon as a microcosm. 

2. Metabolism. Perhaps the most significant peculiarity of 

living matter is found in its instability and the chemical changes 

which continually go on within it. It is constantly wasting away, 

and as constantly being built up. These losses and gains are not 

upon the exterior surface, but throughout its mass. Its growth and 

renewal are by intussusception, or the taking in of new particles, 

and storing them between those already present. A bit of pro- 

toplasm may retain its indentity while all the matter of which it is 

composed is changed over and over. It is like a whirlpool or wave in 

a river which remains the same while the water of which it is com- 

posed changes continually. Metabolism has been aptly defined by 

Huxley as the whirlpool character of the organism. 

3. Irritability. All living matter responds to stimulation. When 

matter fails to be irritable or responsive to stimuli, we declare it to 

be dead. The stimuli that excite reactions in living matter are of 

two kinds, viz; intrinsic and extrinsic. 

Intrinsic stimuli are inherited stimuli. They determine that the 

plant shall conform to a particular type, carry on certain activities, 

pass through a definite life cycle, and detach a portion of its own 

substance for the formation of new individuals of its kind. 

Extrinsic stimuli initiate, inhibit, accelerate or modify the effects 

of intrinsic stimuli. They comprise agents of the external world 

such as cold, heat, chemicals, food, water, light, oxygen, electricity, 

gravity etc. 

The irritable reactions manifested by protoplasm and living things 

to the effects of these external agents will now be considered briefly.

VEGETABLE CYTOLOGY 63 

Thermotropism is the response of living substance to the stimulus 

of temperature. All living substance is influenced by variations in 

it while excessive heat causes 

temperature. Freezing disintegrates 

its coagulation. Active vital phenomena are therefore only evident 

within these extremes, the limits differing depending upon the 

endurance of the organism under examination. The lowest tem- 

perature at which the activity of an organism becomes evident is 

known as the minimum, that at which the activities are at their 

best, the optimum, and the highest at which they can be continued, 

the maximum. Some plants are able to endure greater extremes 

of temperature variation than others because of special adaptations. 

Thus, certain bacteria produce spores which resist exposure for an 

hour to the temperature of liquid hydrogen (-225C) or to that of a 

hot air oven at iooC. Many higher plants can endure moderately 

low temperatures by the development of a hairy covering; others 

which are killed by frost produce seeds which can endure rigid cold, 

still others adapt themselves to existence through periods 

of cold 

by passing through a latent stage in the form of bulbs, like the Squill 

or the Lily, or rhizomes, as the Blood Root or the Hellebores. 

Chemotropism is the response of protoplasm to chemical stimulation. 

Any substances that possess the property of producing a 

deleterious effect upon protoplasm are termed poisons. Poisons 

may effect an immediate destructive combination with living substance 

when they are called caustics, or they may have an exciting 

or depressing effect which may eventually prove destructive without 

visible structural change, when they are termed toxins. Caustics 

may liquefy the protoplasm, as the alkalies, or coagulate it, as the 

acids or salts of metals. When well diluted, chemicals may occasion 

no destructive effects, but may call forth positive or negative 

responses, known as positive or negative chemotropism. 

Thus, Pfeffer, working with the motile sperms of ferns, found that 

if a capillary tube, containing a solution of malic acid be introduced 

into water containing them, the sperms moved toward it and entered. 

It is now generally believed that the motile male sexual cells of all 

flowerless plants are attracted to the appropriate female sexual cells 

by means of positive chemotropic influences. Among flowering 

plants, it has been observed that pollen grains brought by various

64 


agencies from anthers to stigmas of certain plants of different species 

will not germinate but when they are carried from one plant to an- 

other of the same species or variety they readily send their pollen 

tubes through stigma and style to the ovule below. In the former 

instance, negative chemotropism is illustrated, while, in the latter, 

positive chemotropism is shown. 

matter to the influence of food. 

4 Sitotropism is the reaction of living 

Hertwig found that if a fine capillary tube be filled with a i per cent, 

solution of asparagin or beef extract and held in contact with a drop 

of water containing certain bacteria, a mass of these soon plugged 

the mouth of the tube. His experiment shows that these organisms 

moved from a poorer to a richer nutrient medium in response to a 

positive sitotropic influence. Oxytropism is the response 

to the 

stimulating influence of oxygen. We see evidence of this everywhere 

in nature. No living thing can continue to exist without this ele- 

ment. A mistaken idea is often prevalent regardingobligate ana- 

erobic bacteria. Like all other bacteria or organisms, these plants 

require oxygen but can only assimilate it in its combined form. 

The tetanus bacillus is a good example. Aerobic bacteria, on the 

other hand, require free (uncombined) oxygen for assimilative purposes. 

Thus the tetanus organism grows in the depth of culture 

media, whereas the tubercle bacterium (an aerobe) grows only 

on the surface. 

Hydrotropism is the response of protoplasm to the stimulus of 

water. This reaction is seen in both positive and negative phases 

in the slime molds. The vegetative stage of these lowly plants is 

characterized by a naked many nucleated mass of protoplasm, con- 

fining itself to the moist crevices of rotten logs etc. until the surface 

of the substratum becomes wet when and only when it will emerge. 

As soon, however, as its fruiting stage begins, the whole protoplasmic 

mass wells up from the substratum, away from moisture. The 

roots of young seedling plants show positive hydrotropism by growing 

toward moisture in the soil. 

Heliotropism is the response of living substance to the stimulus of 

light. The stems of higher plants tend to grow toward the light 

and are, therefore, positively heliotropic, whereas the roots grow 

away from the source of light and so are negatively heliotropic.


Geotropism is the response of protoplasm to the stimulus of gravity. 

Roots of Pteridophytes and seed plants invariably grow downward 

toward the center of gravity and so are positively geotropic. 

The 

FIG. 30. Venus fly-traps, Dioncea muscipula, growing in field near Wilmington, 

N. C. (Photograph by Charles Palmer.') 

fruiting organs of the fungi and the main stems of higher plants 

tend to grow perpendicular to the earth's surface and so are negatively 

geotropic. Branches of stems that assume a relation parallel


to the earth's surface are diageotropic. The Lima Bean, Sarsa- 

parilla, Poison Ivy, and other plants whose stems twine about supports 

exhibit lateral geotropism in their horizontal curvatures. 

Galvanotropism is the reaction of protoplasm to electrical stimuli. 

In this connection it may be said that the degree of response bears 

a definite relation to the intensity of the stimulus. No visible 

external electrotropic reactions have been observed in higher plants, 

PIG. 31. Mimosa Spegazzini. Note the expanded condition of the leaves before 

stimulation. (After Steckbeck.) 

although when their cells are examined microscopically, the reaction 

becomes manifest. Kuhne has shown that when an. electric 

current is passed through the hairs of the Spiderwort, the cytoplasm 

becomes gathered into small globular masses. 

Thigmotropism is the response of living matter to mechanical 

stimulation. Examples of this form of irritability appear to be far 

less common among plants than among animals. Certain species

VEGETABLE CYTOLOGY 6 7 

of Mimosa, Oxalis, Drosera, Desmodium and Dioncea muscipula exhibit 

this phenomenon to a marked degree. A few instances only 

will be considered. When the tendrils of climbing plants come into 

contact with the uneven surface of solid bodies they are induced to 

coil. When the tentacles on a modified leaf of the Sundew (Dro- 

sera) are stimulated mechanically by an insect or artifically they are 

induced to curve over. If a good plant of the Venus Fly-trap (Dioncea) 

PiG-32. Mimosa Spegazzini. After the application of a stimulus. Compare 

with Fig. 31. (After Steckbeck.} 

is selected, it will be seen to possess leaves, the terminal portions 

of which are modified as traps for catching insects (Fig. 30). Hairs 

will be seen projecting from the upper surface of each valve of the 

hinged blade. If one of these hairs is touched with a pencil no re- 

action will be evident but if after a lapse of twenty seconds the hair 

is touched again, the 2 valves close. If the stamens of Berberis be 

touched near the base during their pollen shedding stage they will 

be observed to curve toward the stigma.


The most highly specialized form of thigmotropism observed in 

plants appears to be found in Mimosa Spegazzini, a member of the 

Bean family. According to Steckbeck "when a mechanical stimu- 

lus, such as a forceps pinch, is applied to one of the terminal secondary 

leaflets after a latent period of less than Y second, the leaflet 

stimulated rises and its partner almost at the same time. The 

stimulus is then carried down the midrib, the pairs of secondary 

leaflets closing in order; in 9 seconds all the secondary leaflets, have 

closed, the midribs converge followed in 3 seconds by a drop of the 

entire leaf. The stimulus moves up the other leaflet with the result 

that the secondary leaflets close in order. * 

In 20 seconds after the 

stimulus has been applied all of the secondary leaflets are closed. 

The stimulus is propagated through the stem to other leaves." 1 

(Figs. 31 and 32.) 

4. Reproduction. Protoplasm also shows a very remarkable 

ability to increase and to give off detached portions which retain 

the infinitely complex peculiarities and properties of the original. 

The process, moreover, may be continued indefinitely. 

Other physiological characteristics might be added, but the above 

are mentioned as the most satisfactory criteria by which living may 

be distinguished from non-living matter. 

PROTOPLASMIC CELL CONTENTS 

Protoplasm consists of four well-differentiated portions: 

(a) Cytoplasm, or the foamy, often granular matrix of protoplasm 

outside of the nucleus. 

(b) Nucleus or Nucleoplasm, a denser region of protoplasm con- 

taining chromatin, a substance staining heavily with certain basic 

dyes. 

(c) Nucleolus, a small body of dense protoplasm 

nucleus. 

within the 

(d) Plastids, composed of plastid plasm, small discoid, spheroidal, 

ellipsoidal or ribbon-shaped bodies scattered about in the cytoplasm. 

1 "The comparative histology and irritability of sensitive plants" by D. W. 

Steckbeck in Contributions from The Botanical Laboratory of the U. of Pa., 

vol. IV, No. 2, p. 217, 1919.


Sometimes, as in the cells of lower plants like the Spirogyra, plastids 

are large and are then called chromatophores. 

According to the position of the cells in which plastids occur and 

the work they perform, they differ in color, viz: 

Leucoplastids are colorless plastids found in the underground 

FIG. 33. A, embryonic cells from onion root tip; d, plasmatic membrane; c, cytoplasm; 

a, nuclear membrane enclosing the thread-like nuclear reticulum; b, 

nucleolus; e, plastids (black dots scattered about). B, older cells farther back 

trom the root tip. The cytoplasm is becoming vacuolate; /, vacuole. C, a cell 

from the epidermis of the mid-rib of Tradescantia zebrina, in its natural condition 

on the right, and plasmolyzed by a salt solution on the left; g, space left by 

the recedence of the cytoplasm from the wall; the plasma membrane can now be 

seen as a delicate membrane bounding the shrunken protoplast. All highly magnified. 

(Stevens.) 

portions of a plant and also in seeds, and other regions given up to 

the storage of starch. Their function is to build up reserve starch 

from sugar and other carbohydrates as well as to change the reserve 

starch back into sugar when it is needed for the growth of the plant. 

6 9

70 


They are only evident after properly fixing and staining cells containing 

them. 

Chloroplastids are plastids found in cells exposed to light and con- 

tain the green pigment, chlorophyll. 

Chromoplastids are plastids found in cells independent of their 

relation to light or darkness and contain a yellow, orange or red 

pigment called chromophyll. 

CELL FORMATION AND REPRODUCTION 

The cells of plants have all been derived from preexisting cells. 

In the bacteria and many other low forms of plant life, 

the division 

of the cell always results in reproduction; in higher forms, however, 

it merely increases the size of the individual and so is a phenomenon 

of growth. 

There are two kinds of cells formed by plants, viz. : asexual and 

sexual. Both of these are endowed with the possibilities of repro- 

duction, although the former are frequently limited to the process 

of growth. 

Reproduction is the power possessed by an organism of giving rise 

to new individuals. This may take place through the agency of 

either asexual or sexual cells and is accordingly asexual or sexual in 

character. Whenever a union of cells or their protoplasmic con- 

tents takes place the process is called " sexual reproduction;" if, 

however, there is a mere separation of a cell or cells from an individual 

which later form a new organism, 

" 

asexual or vegetative reproduction." 

the process is termed 

There are four modes of asexual reproduction, viz.: Fission, 

Gemmation, Free Cell Formation and Rejuvenescence. 

Fission. This is the separation of a cell into two equal halves, 

each of which may grow to the size of the original parent cell from 

which it was derived. Fission is seen in the reproduction of bac- 

teria, growth of many algae and the formation of tissues of higher 

plants. 

Gemmation or Budding. This is the method of reproduction 

common among yeasts. The cell forms a protuberance called a 

bud which increases in size until it equals the size of the cell which 

formed it and then becomes detached, although frequently not until 

it has developed other buds and these still others.


Free Cell Formation. This is a type of reproduction in which the 

nucleus and protoplasm become separated into two or more masses 

each of which forms a cell wall about itself. Seen in formation of 

ascospores within ascus of Ascomycetes and spores within spore cases 

of molds. 

Rejuvenescence. In this mode of reproduction the protoplasm 

of the cell becomes rounded out, escapes by rupture of the cell wall, 

forms cilia and moves about as a zoospore. Later it looses its cilia, 

develops a cell wall and passes into a resting condition. Under 

favorable circumstances it grows into a new organism. It is found 

in (Edogonium, Ectocarpus, etc. 

There are two kinds of sexual reproduction, viz. : Conjugation and 

Fertilization. In both of these the sexual cells called gametes or 

their nuclei come together and their protoplasm blends to form a 

new cell. This is the common method seen in higher plants. 

Conjugation. A union of two gametes, alike in character, the 

This method of reproduction is 

product being a zygote or zygospore. 

seen in the molds, Spirogyra, Zygnema and Diatoms. 

Fertilization. A union of two unlike gametes or their nuclei, the 

product being an oospore. One gamete, the male sexual cell, is 

smaller and active, while the other, the female sexual cell, is larger 

and passive. This process is seen among the higher and many of 

the lower plants. 

INDIRECT NUCLEAR DIVISION (MITOSIS OR KARYOKINESIS) 

This is the general method of division seen in the formation of 

tissues of higher plants. 

The process begins in the nucleus and ends with the formation of 

a cell wall dividing the new- formed cells. 

When we examine a cell in its resting stage we find the nucleus 

more or less spherical in shape, surrounded by a nuclear membrane 

and containing a nuclear network, nuclear sap and one or more 

nucleoli. The nuclear network consists of a colorless network of 

limn adhering to which are numerous minute granules called chro- 

matin which take the stain of a basic dye. Surrounding the nucleus 

is the cytoplasm. 

As the cell commences to divide, the nucleus elongates and the


g i 

FIG. 34. Semi-diagrammatic representation of nuclear and cell-division, a, 

resting cell ready to begin division; b, the nuclear reticulum is assuming the form 

ot a thickened thread, and the cytoplasm at opposite poles is becoming thread- 

like to form the spindle fibers; c, the nuclear thread has divided longitudinally 

through the middle, and the spindle fibers have become more definite; d, the nuclear 

membrane and the nucleolus have disappeared, and the nuclear thread has 

become segmented into chromosomes which are assembling at the equator 01 the 

cell. All of the phases of division thus far are called prophases. e, the metaphase, 

where the longitudinal halves of the chromosomes are being drawn apart preparatory 

to their journey toward the opposite poles; /, the anaphase, or movement 

of the chromosomes toward the poles, is about completed, connecting 

fibers extend from pole to pole; g, telophase where the chromosomes have begun 

to spin out in the form of a nuclear reticulum. The connecting fibers have begun 

to thicken in the equatorial plane; h, the connecting fibers have spread out and 

come into contact with the wall of the mother cell in the equatorial plane, and 

the thickening of the fibers throughout this plane has made a complete cell plate 

within which the dividing wall will be produced; i, a nuclear membrane has been 

formed about each daughter nucleus, and the dividing cell-wall is completed. 

The two daughter cells are now ready to grow to the size of the parent cell in a, 

when the daughter nuclei will appear as does the nucleus there. All highly magnified. 

(Stevens.)


iinin threads of the nuclear reticulum shorten, drawing the chromatin 

granules together into a thickened twisted chromatic thread. 

This thread splits transversely and thus becomes divided into a 

number of rods termed chromosomes. Each of these then splits into 

two longitudinal halves that may be termed the daughter -chromo- 

somes. They lie within the nuclear cavity united by delicate threads. 

There now begins a phenomenon concerned with the cytoplasm 

which is primarily a process of spindle formation. The granular 

cytoplasm accumulates at the poles of the elongated nucleus forming 

the cytoplasmic caps. Presently it begins to show a fibrillar struc- 

ture, the threads extending outward around the periphery of the 

nucleus forming an umbrella-like arrangement of fibers from both 

cytoplasmic caps. With the formation of fibers comes a breaking 

down of the nuclear membrane and in consequence the fibers enter 

the nuclear cavity and organize the spindle. Some of the fibers 

become attached to the split chromosomes and push, draw or pull 

them to the equatorial plate, halfway between the poles. Mean- 

while the nucleolus disappears. As the chromosomes line up at the 

equatorial plate their daughter halves are drawn apart in V-shaped 

fashion. The split extends and eventually one daughter-chromosome 

is drawn to one pole and the remaining half to the other. At 

the respective poles the daughter chromosomes form a dense com- 

pact knot. A cell membrane, composed of material contributed 

largely through the shrinking of the spindle fibers, is now formed 

through the middle of the spindle. This soon splits to form a thin 

vacuole lying between the two membranes (the plasma membranes) . 

Presently there appears within the vacuole and between the membranes 

a carbohydrate substance. On either side of this deposit the 

plasma membranes form a cellulose membrane. The flattened 

vacuole extends toward the periphery and ultimately a complete 

cell wall is formed. 

The dense compact knots of chromosomes at the poles of the 

spindle, that are to form the daughter-nuclei, now begin to expand 

and clear mesh-like spaces to appear between the expanding threads. 

As this process advances the chromosome substance becomes dis- 

tributed throughout the nuclear cavity. It is soon possible to dis- 

tinguish the chromatin from the Iinin. Eventually an irregular

74 


network of linin carrying chromatin granules is formed through the 

area of the nucleus. A nuclear membrane also is formed and the 

nucleolus reappears. The spindle fibers disappear. The daughter- 

nuclei increase in size and each daughter-cell formed by this process 

now assumes the resting stage. 

NON-PROTOPLASMIC CELL CONTENTS 

i. Sugars. Sugars comprise a group of crystalline substances 

found in the cell sap of many plants either free or in combination 

with glucosides. They may be divided into two main groups: 

monosaccharoses and disaccharoses. To the former belong simple 

sugars containing two to nine atoms of carbon, which are known 

respectively as bioses, trioses, tetroses, pentoses, hexoses, etc. Of 

these the hexoses (CeHÔe) are the most important and of wide 

distribution. Examples of the hexoses found in drug plants are: 

(a) dextrose (grape sugar), found in the leaves, stems, fruits, sprouting 

grains and nectaries of flowers of nearly all plants; (b) fructose 

(levulose or fruit-sugar), commonly associated with dextrose; (c) 

d-mannose, found in the saccharine exudation of the Manna Ash 

(Fraxinus Ornus)\ and (d) sorbinose, found irf ripe Mountain Ash 

berries. Upon evaporating the sap or treating the parts containing 

these principles with alcohol they can be crystallized out. 

Fliickiger's Micro-chemic Test for the determination of different 

kinds of sugars: Dissolve a small portion of copper tartrate in a 

in this place the section and 

drop of sodium hydrate on a glass slide ; 

put on the cover slip. If fructose is present cuprous oxide crystals 

will at once be formed without warming. If grape sugar is also 

present a gentle warming will produce another crop of reddish-yellow 

crystals. If dextrin be present continued heating will still further 

augment the number of crystals. Cane sugar and mannite, on the 

other hand, will respond negatively to this test. The zymase of 

yeasts is capable of fermenting dextrose, levulose and d-mannose 

carbon dioxide and alcohol. Sorbinose is claimed to be 

forming 

non-fermentable. 

The disaccharoses, having the chemical formula of Ci 2H 22Oii, in- 

clude sucrose, maltose, trehalose, melibiose, touranose and agavose. 

Of these sucrose is the most important. It is found in the stems of


sugar cane, sorghum, corn and Mexican grass; in many fleshy roots 

notably the sugar beet; in the sap of the sugar maple and various 

palms including Cocos nucifera, Phoenix syhestris, Arenga saccharifera; 

in various fruits, as apples, cherries, figs, etc., in the nectaries 

of certain flowers; in honey; and in a number of seeds. It crystal- 

lizes in monoclinic prisms or pyramids. When sections of plant parts 

containing cane sugar are placed for a few seconds in a saturated 

solution of copper sulphate, then quickly rinsed in water, trans- 

ferred to a solution of i part of KOH in i part of water, and heated 

to boiling, the cells containing the sugar take on a sky-blue color. 

Invertase of the yeast reduces cane sugar to dextrose and levulose 

and zymase of the same plant ferments these forming carbon dioxide 

and alcohol. 

Maltose is found in the germinating grains of barley and other 

cereals as a product of the action of the ferment diastase on starch. 

It reduces Fehling's solution, forming cuprous oxide, but one-third 

less with equal weights. 

Trehalose or mycose is found in ergot, Boletus edulis, the Oriental 

Trehala and various other fungi. 

Melibiose is formed with fructose upon hydrolyzing the trisac- 

charose melitose which occurs in the molasses of sugar manufacture 

and in Australian manna. 

Touranose is produced upon hydrolyzing melizitose, a trisaccharose 

which occurs in Persian manna, and 

Agavose is found in the cell sap of the American Century Plant, 

Agave americana. 

2. Starch. Starch is a carbohydrate having the chemical formula 

of (CeHijOs^ which is generally found as the first visible product 

of photosynthesis in most green plants. It is found in the chloro- 

plasts and chroma tophores of green parts 

in the form of minute 

granules. This kind of starch is known as Assimilation Starch. 

Assimilation starch is dissolved during darkness within the chloro- 

plasts by the action of ferments and passes into solution as a glucose 

which is conveyed downward to those parts of the plant requiring 

food. In its descent some of it is stored up in medullary ray cells, 

and in various parts of the xylem, phloem, pith and cortex in the 

form of small grains. Considerable, however, is carried down to


the underground parts, such as rhizomes, tubers, corms, bulbs 

or roots, where the leucoplasts store it in the form of larger-sized 

grains called Reserve Starch. This type of starch is generally 

characteristic for the plant in which it is found. It constitutes 

stored-up food for the plant during that period of the year when the 

vegetative processes are more or less dormant. 

Structure and Composition of Starch. Starch grains vary in 

shape from spheroidal to oval to chonchoidal to polygonal. They 

FIG. 35. Cell of Pellionia Daveauana, showing starch-grains. The black, 

crescent-shaped body on the end of each grain is the leucoplast. Greatly enlarged. 

(Gager.) 

are composed of layers of soluble carbohydrate material and probably 

other substances called " lamella," separated from each other by 

a colloidal substance resembling a mucilage in its behavior toward 

aniline dyes. They contain a more or less distinct highly refractile 

point of origin or growth called the " hilum" which also takes the 

stain of an aniline dye. The layers of carbohydrate material stain 

variously, blue, indigo, purple, etc., with different strengths of 

iodine solutions. Each grain is covered with a stainable elastic 

membrane.


Starch grains may be grouped, according to the condition in 

which they are foiihd in the cells of storage regions 

into three 

kinds, viz.: simple starch grains, compound starch grains and fill 

starch grains. 

Simple starch grains are such as occur singly. Compound starch 

grains occur in groups of two, three, four, five, six or more and are 

designated as two, three, four, five, six, etc., compound, according 

to the number of grains making up the group. Fill starch grains 

are small grains filling up the spaces between the larger grains in 

storage cells. These are common in commercial starches. 

Method of Examining Reserve Starches. Many of the reserve 

starches are used commercially, such as potato, corn, rice, maranta, 

oat, wheat, sago, tapioca, etc., and it frequently becomes necesssary 

for the microscopist to determine their purity or their presence in a 

sample of food or drug. The following characteristics should be 

noted in determining the identity or source of the starch. 

1. The shape of the grain. 

2. Whether simple or compound or both; if compound, the number 

or range in numbers of grains composing it. 

3. The size of the grain in microns. 

4. The position of the hilum, if distinct; whether central or excen- 

tric (outside of the center). 

5. The shape of the hilum and the degree to which it is often 

fissured. 

6. The nature of the lamellae, whether distinct or indistinct; if 

distinct whether concentric (surrounding the hilum) 

or eccentric 

(apparently ending in the margin and not surrounding the hilum), 

or both, as in potato starch. 

7. The color of the grains when stained with dilute iodine solu- 

tions; whether indigo, blue, purple, red or yellowish-red, etc. 

8. The appearance under polarized light. 

9. The temperature at which the paste is formed. 

10. The consistency of the paste.

00 d o 


FIG. 36. A, wheat starch grains; B, rye starch; C, barley starch; D, potato 

starch; E, Maranta starch; P, Sago starch. , Explanation in text.


CHARACTERISTCS or IMPORTANT COMMERCIAL STARCHES 

Potato Starch (Solanum tuberosum) 

Mostly simple, conchoidal or ellip- 

soidal, with occasional spheroidal 

and two- to three-compound grains. 

Size: 5 to 12 5 n 

Hilum: circular, at smaller end of 

grain. 

Lamellae: concentric and eccentric. 

Polarization cross very distinct ; beau- 

tiful play of colors with selenite 

plate. 

Maranta Starch (Maranta 

arundinacea) 

Ellipsoidal to ovoid. 

Simple. 

Size: 10 to 65 /*. 

Hilum: a transverse or crescent- 

shaped cleft in center or near 

broad end of grain. 

Lamellae : usually indistinct. 

Polarization cross very distinct; fine 

play of colors with selenite plate. 

X Corn Starch (Zea Mays} 

Polygonal to rounded. 

10 to 35/z. Most grains over i^/j. 

in diameter. 

Simple. 

Hilum: circular or a two- to five- 

rayed cleft. 

Lamellae: indistinct. 

Polarization cross distinct but no 

marked play of color with selenite 

plate. 

Rice Starch (Oryza saliva) 

Polygonal.. 

2 to io/i in diameter. 

Simple or two- to many-compound. 

Hilum: usually indistinct, occasion- 

ally a central cleft. 

Lamella) : indistinct. 

Polarization cross distinct but no 


Wheat Starch (Triticum satwum} 

Circular grains appearing lenticular 

shaped on edge view; simple. 

Large grains 28 to 45^1 in diameter. 

Hilum: central, appearing as dot, 

but usually indistinct. 

Lamellae : generally indistinct, when 

present concentric. 

Polarization cross indistinct; no 


Rye Starch (Secale cereale) 

Grains having a similar shape to 

those of wheat starch but many 

larger; simple. 

Large grains 20 to 52^' in diameter. 

Hilum: a star-shaped central cleft or 

indistinct in some grains. 

Lamellae : concentric. 

Polarization cross distinct. 

Barley Starch (Hordeum distichon] 

Grains having a similar shape to 

those of wheat starch but fre- 

quently tending to bulge on one 

side and so appear sub-reniform; 

large grains smaller; simple. 

Grains appear elliptical to lemon 

shaped in edge view. Large 

grains usually 18 to 25^, occa- 

sionally up to 30/4 in length. 

Hilum: centric or circular or cleft, 

often indistinct. 

Lamellae : concentric, often indistinct. 


Buckwheat Starch (Fagopyrum 

esculentum) 

Grains simple and compound. 

Simple grains polygonal or rounded 

polygonal.

So 

g

Compound grains more or less rod- 

shaped. 

2 to 15^1 in diameter. 

Hilum: central. 

Lamellae : generally indistinct. 


Cassava Starch (Manihot 

utilissima) 

Grains rounded, truncated on one 

side. 

Simple or two- to three- or four- to 

eight-compound. 

6 to 35/x in diameter. 

Hilum: central, circular or triangu- 

lar with radiating clefts frequently. 

Lamellae : indistinct. 

Polarization cross prominent. 

Bean Starch (Phaseolus vulgaris) 

Ovoid, ellipsoidal or reniform shapedsimple 

grains, occasionally ob- 

scurely 3- or 4-sided. 

25 to 60 fj. in length. Generally from 

30-35M- 

Hilum: central, elongated with bran- 

ching clefts. 

Lamellae: distinct, concentric. In 

some indistinct. 

Polarization crosses shaped thus, X 

VEGETABLE CYTOLOGY 8l 

Pea Starch (Pisum sativum) 

Oval-oblong, ellipticolar sub-reni 

form. 

I S~S I M i*1 length. Generally from 

Hilum: similar to that of bean 

starch but less cleft or simply 

elongated. 

Lamellae: distinct, concentric. 

Polarization crosses similar to bean 

starch. 

Canna Starch (Canna edulis and other 

species of Canna) 

Broadly elliptical, flattened, with 

beak or obtuse angle at one end. 

50 to 135/i in length. 

Hilum: excentric near narrower end. 

Lamellae: concentric and excentric. 

Polarization cross very distinct; fine 


Sago Starch (Metroxylon Sagu] 

Ovoid, muller shaped, or irregularly 

3 or 4 sided with rounded angles. 

Some more or less gelatinised. 

Simple or 2, 3 or 4-compound 

3-6oju long. 

Hilum: eccentric often altered by 

gelatinisation. 

Lamellae : Excentric and concentric. 


4. Dextrin. Dextrin is a carbohydrate made from starch (chiefly 

from corn or potato starch) by the application of heat (yellow 

dextrin) or by treatment with both heat and acids (white dextrin). 

It forms a paste with water, the yellow variety tending to swell up 

and dissolve much more readily than the white. When examined 

microscopically in alcohol mounts, the grains, while conforming in 

general outline to those of the type of starch from which the dextrin 

was prepared, nevertheless show more conspicuous striations and 

clefts. . Corn dextrin shows distinct striations, whereas striations 

6


in corn starch are absent. The grains take on a red coloration with 

iodine solutions. 

5. Amylodextrin. This is a carbohydrate intermediate in proper- 

ties between starch and dextrin. It occurs in the form of small ir- 

regularly shaped granules, in Mace, that take on a reddish brown to 

reddish violet color with iodine solutions. 

6. Inulin. Inulin is a carbohydrate isomeric with starch which 

has the chemical formula of Ci 2H 20Oio. 

It is found dissolved in the 

cell sap of many plants, especially those of the Composite. If pieces 

of a plant part containing this substance be placed directly in alcohol 

for at least a week, then sectioned and mounted in alcohol, sphaerocrystals 

of inulin will be seen applied to the walls of the cells. When 

these sections are treated with a 25 per cent, solution of alpha 

naphthol and 2 or 3 drops of strong H 2SO 4 , the sphaerocrystals will 

dissolve with a violet color. Fehling's solution is not reduced by 

inulin. 

7. Hesperidin. Hesperidin is a glucoside having 

the chemical 

formula of C 2 iH 2 . eOi 2 Like inulin it occurs in solution within the 

cell sap. 

It is found in abundance in the Rutaceae family but occurs 

in many other plants. If sections of alcoholic material containing 

this substance such as Buchu leaves or unripe orange peel, are 

mounted in alcohol and examined, sphaerocrystals will be seen. If 

these are then treated with a drop of alpha naphthol solution and 2 

or 3 drops of strong H 2SO 4 , they dissolve with a yellow color. The 

same coloration is evident when 5 per cent, solution of KOH is 

substituted for the alpha naphthol and H 2SO 4 . 

8. Strophanthin. This is a glucoside occurring in the cell sap of 

the endosperm of Strophanthus Kombe, S. hispidus and other species 

of Strophanthus. If sections of fresh Strophanthus seeds are 

mounted in a drop of water and then transferred to a drop of con- 

centrated H 2SO 4 , the cells containing Strophanthin will assume a 

bright green color. 

9. Salicin. Salicin is a glucoside occurring in the cell sap of the 

bark and leaves of the Willows and Poplars. Sections of these 

mounted in concentrated H 2SO 4 will show a red coloration in the 

cells containing this substance. If water be added a red powder is 

thrown down.


10. Saponin, another glucoside, found in Soap Bark, Senega, 

Saponaria and other drugs also takes a red color with strong H 2SO4. 

11. Coniferin is a glucoside, occuring in the cell sap of the spruce, 

pine, and other plants of the Conifera. If sections containing- it are 

first treated with a solution of phenol and then with sulphuric acid, 

the cells containing it take on a deep blue color. 

12. Digitoxin, a glucoside found in the leaves of Digitalis purpurea, 

is colored green with hydrochloric acid. 

The glucosides are very numerous. Those listed above represent 

but a few examples. They arise in the cell sap of plants containing 

them as products of constructive metabolism (anabolism) and are 

thought by many to have the function of protecting plants against 

the ravages of animals. Some are known to serve as reserve food. 

All glucosides are characterized by the property of being split up 

into glucose and other substances when acted upon by a ferment, 

dilute acids or alkalies. 

13. Alkaloids. Chemically, these are basic carbonaceous amines 

which like glucosides are products of metabolism. Their method 

of formation in plants is uncertain. Some hold that they are kata- 

bolic products, resulting from the breaking down of tissues, while 

others believe them anabolic in character. They undoubtedly serve 

as defensive agents in plants containing them on account of their 

bitter taste and poisonous properties. 

Properties of Alkaloids 

Alkaloids are invariably found in combination with acids forming 

salts which dissolve in water or alcohol. They are composed of 

carbon, hydrogen and nitrogen. Some contain oxygen. They are 

precipitated from saline solutions by the addition of alkalies. They 

are mostly colorless and crystallizable. They can be precipitated 

by one or more of the following alkaloidal reagents: tannic acid, 

gold chloride, phospho-molybdic acid, picric acid and potassiomercuric 

iodide. 

O 2 , 

Examples of Alkaloids 

Strychnine. This alkaloid, with a chemical formula of C 2iH22N 2 - 

occurs in the seeds of Strychnos nux vomica, Strychnos Ignatii 

and other species of Strychnos. When sections of strychnine con-

84 


taining seeds, previously treated with petroleum ether and absolute 

alcohol, 

are mounted in a solution of i Gm. ammonium vanadate 

in 100 mils of sulphuric acid, they take on a violet-red color which 

later changes to brown. 

Veratrine. This alkaloid, with a composition of CarE^NOn, 

is found in various parenchyma cells of Veratrum album. If sections 

of the rhizome or roots are mounted in 2 drops of water and a drop 

of concentrated H 2SO 4 and examined microscopically on a glass 

slide, the cell contents and walls of the cells which contain this substance 

first take a yellow color which soon changes to an orange-red 

and then to a violet. 

Nicotine. This is a volatile alkaloid having the formula of 

CioHi 4N2 which is found in the Nicotiana genus of the Nightshade 

family* Sections of tobacco leaves or stems mounted in dilute 

Lugol's solution will show first a carmine-red color and then a reddish-brown 

precipitate which in time loses its color. 

Caffeine. This alkaloid, with a formula of C 8Hi N 4O2 + H 2O, 

occurs in Thea, Cojfea, Cola, Sterculea, Ilex and Neea. If thin sections 

containing it are placed on a glass slide in 2 or 3 drops of con- 

centrated hydrochloric acid and gently heated and then 2 or 3 drops 

of gold chloride solution are added, the sections then pushed to the 

side and the liquid allowed to evaporate, slender yellowish branch- 

ing needles of caffeine gold chloride will be seen to separate. 

Cocaine. This narcotic alkaloid, having the formula Ci7H 2iNO 4 , 

is found in the leaves of Erythroxylon Coca and E. Truxillense. If 

sections of these leaves are prepared in the same manner as indicated 

for those containing Caffeine, but platinum chloride solution substi- 

tuted for that of gold chloride, large feathers or plumes of cocaine- 

chloro-platinate will be seen separating. 

Aconitine (C 33H 43NOi 2 ) is found in various parts of Aconilum 

Napellus. It is particularly abundant in the tuberous root of this 

plant. If sections of aconite root are treated on a glass slide with 

solution of potassium permanganate, a red precipitate of aconitine 

permanganate will appear in the cells containing this alkaloid. 

Colchicine (C 22H25NO 6). This alkaloid occurs in the corm and 

seeds of Colchicum aulumnale. It is very abundant in the cells 

surrounding the fibro-vascular bundles of the corm. If a section of


either corm or seed be treated with a mixture of i part of H 2SO 4 

and 3 parts of H 2O, the cells containing colchicine will be colored 

yellow. If a crystal of KNO 3 then be added the color will change 

to a brownish-violet. 

10. Gluco-alkaloids. These are compounds intermediate in 

nature between alkaloids and glucosides, having characteristics of 

each. To this group belongs solanine (C28H 47NOii) which is found 

in Solanum nigrum, Solanum Dulcamara, Solanum carolinense and 

other species of the Solanacea. When sections of those plant parts 

which contain this constituent are mounted in a solution of i part 

of ammonium vanadate in 1000 parts of a mixture of 49 parts of 

sulphuric acid with 18 parts of water, the cells containing solanin 

take on a yellow color which changes successively to orange, various 

shades of red, blue-violet, grayish-blue and then disappears. 

14. Asparagine (C 4H 8N2 + H 2O). This is an amino compound of 

crystalline nature which occurs widely in the plant kingdom. It 

has been found in certain of the slime molds and fungi, in the roots 

of Alth&a officinalis and Atropa belladonna, in young shoots of 

Asparagus, in the seeds of Castanea dentata, in the tubers of Solanum 

tuberosum and varieties of Dahlia, and is known to play an important 

part in metabolism. Stevens claims that proteids are reduced for 

the most part to asparagine during seed germination. 1 

If thick sec- 

tions are cut from a plant part containing this substance and 

mounted in alcohol, rhombohedral crystals of asparagin in the form 

of plates will be deposited upon the evaporation of the alcohol. If 

to these a few drops of a saturated solution of asparagine are added 

the crystals already formed will increase in size. To get satisfactory 

results the saturated solution must be of the same temperature as 

the mount. 

15. Calcium Oxalate. This substance, occurs in many plants 

always in the form of crystals. It is apparently formed by the reaction 

of salts of calcium, which have found their way into the cell 

sap from the soil, with oxalic acid which is manufactured by the 

plant. Calcium oxalate crystals dissolve readily in mineral acids 

without effervescence. They are insoluble in acetic acid or water. 

1 Stevens' Plant Anatomy, 3d Edit., p. 189.


These crystals are classified according to form and belong either to 

the monoclinic or tetragonal system (See Fig. 38). 

A 

V 

B 

FIG. 38. Various forms of calcium oxalate crystals. A, styloids from the 

bark of Quillaja saponaria; B, rosette aggregate from rhizome of Rheum officinale; 

C, raphide from the bulb of Urginea maritima; D, crystal fiber as seen in longitudinal 

section in either the xylem or phloem regions of Glycyrrhiza; E, microcrystals 

(crystal sand) isolated from the parenchyma of Belladonna root; P, 

monoclinic prisms; and G, twin-crystals from leaves of Hyoscyamus niger. All 

highly magnified. 

Crystals belonging to the Monoclinic System and Examples of Drugs 

Containing them: 

1. Solitary Hyoscyamus, A cer Spicatum, Viburnum Prunifolium. 

2. Rosette Aggregates Althaea, Gossypii Cortex, Stramonium, 

Granatum, Rheum, Fceniculum, Viburnum. 

3. Columnar (Styloids) Quillaja.


4. Raphides Convallaria, Sarsaparilla, Veratrum, -Scilla, 

Phytolacca. 

5. Micro-crystals (Crystal sand) Bellandonnae Radix, Cinchona, 

Stramonium, Phytolacca, Capsicum. 

6. Crystal Fibers Cascara Sagrada, Prunus Virginiana, Gly- 

cyrrhiza, Aspidosperma. 

7. Membrane Crystals Aurantii Dulcis Cortex, Limonis Cortex, 

Condurango. 

Solitary crystals, usually in the form of rhombohedra, occasionally 

in twin crystals, occur as sharp angular bodies, each one often com- 

pletely filling up the lumen of a cell. 

Rosette aggregates consist of numerous small prisms or pyramids, 

or hemihedral crystals arranged around a central axis, appearing 

like a rosette or star. 

Columnar crystals or styloids are elongated prisms. 

Raphides are groups of acicular or needle-shaped crystals, which 

occur in long thin-walled cells containing mucilage. They are 

more frequently found in Monocotyledons than in any other plant 

group. Micro-crystals (sphenoidal micro-crystals or crystal sand) 

are minute arrow-shaped or deltoid forms completely filling the 

parenchyma cells in which they occur and giving these a grayishblack 

appearance. 

Crystal fibers are longitudinal rows of superimposed parenchyma 

cells each of which contains a single monoclinic prism or rosette 

aggregate. Crystal fibers are found adjacent to sclerenchyma fibers 

such as bast or woody fibers. 

Membrane crystals are monoclinic prisms, each of which is sur- 

rounded by a wall or membrane. In the process 

of formation a 

crystal first is formed in the cell sap and then numerous oil globules 

make their appearance in the protoplasm surrounding it; later some 

of the walls of the cell grow around the crystal and completely 

envelop it. 

1 6. Cystoliths. Cystoliths are clustered bodies formed by the 

thickening of the cell wall at a certain point and subsequent ingrowth 

which latter forms a cellulose skeleton consisting of a stalk 

and body. Silica is subsequently deposited on the stalk while 

calcium carbonate is piled up on the body in layers, forming an irregu-


lar spheroidal or ellipsoidal deposit. These structures are abun- 

dantly found in the plants of the Nettle Family and constitute a 

leading peculiarity of the same (see Fig. 87). 

Hair cystoliths differ from the average type in that they are 

devoid of a stalk. Such are seen in the non-glandular 

hairs of 

Cannabis sativa. 

The calcium carbonate incrustation of a cystolith dissolves with 

effervescence on the addition of a mineral or organic acid. 

17. Silica. Silica (SiO 2 ) occurs in a number of plants either as 

an incrustation in the cell wall as in Diatomsv the Equisetinea and 

Graminea or more rarely in the form "silica bodies" such as are 

found in certain Palms, Orchids and Tristicha. It is insoluble in 

all the acids except hydroflouric. It may be obtained in pure form 

by placing tissue containing it in a drop 

or two of concentrated 

sulphuric acid and after a time treating with successively stronger 

solutions of chromic acid (starting with 25 per cent.) and then wash- 

ing with water and alcohol. 

1 8. Tannins. Tannins are amorphous substances occurring in 

plants having an astringent taste, and turning dark blue or green 

with iron salts. They occur in greatest quantity in the bark of 

exogens, and in gall formations. They are soluble in water, alcohol 

glycerine, and a mixture of alcohol and ether. They are almost 

insoluble in absolute ether and chloroform. They give insoluble 

precipitates with organic bases such as alkaloids and with most of 

the salts of the heavy metals. 

According to their behavior with solution of iron chloride or 

other soluble iron salts two kinds of tannic acid are recognized: (a) 

a form of tannic acid giving a blue color, as that which is found in 

Rhus, Castanea, Granatum, Galla, etc.; (b) another tannic acid 

producing a green coloration, as that found in Krameria, Kino, 

Mangrove bark, Quercus, Catechu, etc. 

If sections are placed in a 7 per cent, solution of copper acetate 

for a week or more, then placed on a slide in 0.5 per cent, aqueous 

solution of ferric chloride, and after a while washed with water and 

mounted in glycerin, an insoluble brownish precipitate will be produced 

in those cells containing tannin.

VEGETABLE CYTOLOGY 8 9 

19. Proteins. Proteins are complex nitrogenous substances 

forming the most important of the reserve foods of plants. They are 

found in all the living and many of the dead cells of plants, although 

most abundant in seeds. Protoplasm, itself, is composed largely 

of these substances. They all contain carbon, hydrogen, oxygen, 

nitrogen and sulphur, and many contain in addition phosphorus. 

They are formed by the addition of nitrogen, sulphur and fre- 

quently phosphorus to elements of grape sugar. The nitrogen, 

sulphur and phosphorous elements are obtained from nitrates, 

sulphates and phosphates which are dissolved in the water taken 

in through the roots. The names of proteins recorded may be 

found by the hundreds. These are grouped into chemical classes, 

the most important of which from the standpoint of their occurrence 

in plants are the globulins, albumens, glutelins, nudeins, and gliadins. 

Of these the globulins are found most extensively. Globulins are 

insoluble in water but soluble in sodium chloride solutions. They 

do not coagulate upon the application of heat. 

Albumens are soluble in water and coagulate with heat. 

Glutelins are insoluble in water, sodium chloride solution and 

strong alcohol. 

Gliadins are nearly or wholly insoluble in water but soluble in 

70 to 90 per cent, alcohol. 

Nucleins are insoluble in water but soluble in akaline solutions. 

The following tests are of value in determining the presence of 

proteins. 

Lugol's solution stains proteins yellow or brown. 

Concentrated nitric acid stains proteins yellow. This color 

becomes deeper upon the addition of ammonia water. 

Million's reagent stains proteins a brick-red. 

Concentrated solution of nickel sulphate colors proteins yellow 

or blue. 

If sections are placed for an hour or two in a solution of i Gm. of 

sodium phospho-molybdate in 90 Gm. of distilled water and 5 

Gm. of nitric acid, the proteid substances appear as yellowish 

granules. 

The globulins (phytoglobulins) frequently occur in bodies called 

"aleurone grains."

go 


ALEURONE GRAINS 

Aleurone grains are small bodies found in seeds particularly those 

containing oil, and like starch grains often are characteristic of the 

genus or species. Each aleurone grain consists of a ground substance 

(composed of amorphous proteid matter soluble in water, 

dilute alkali or acid), in which are usually embedded one or more 

phyto-globulins (insoluble in cold water, but soluble in less than i per 

cent, solution of an alkali, in dilute HC1 and acetic acid), one or more 

transparent globular globoids composed of Ca and Mg phosphate 

PIG. 39. To show aleurone grains. A , cells from cotyledon of seed of garden 

bean; n, aleurone grains; m, starch; B, cells from endosperm of castor bean; a, 

a, aleurone grain; I, ground substance; k, phytoglobulin; I, globoid. (A, After 

Sachs; B, after Frank.) 

(insoluble in water and dilute potash solution but soluble in i per 

cent, acetic acid solution), and frequently a crystal of calcium 

oxalate, the whole being enclosed by a protoplasmic membrane (so- 

luble in water). (Fig. 39$.) 

The proteins insoluble in the cell-sap water are made soluble for 

translocation by means of proteolytic enzymes which change them 

into proteoses and peptones. 

20. Mucilages and gums are those substances occurring in plants 

which are soluble in water or swell in it, and which are precipitated 

by alcohol. 

Mucilage is formed in plants in several ways, viz.; either as a 

product of the protoplasm, as a disorganization product of some of 

the carbohydrates, as a secondary thickening or addition to the cell 

wall, or as a metamorphosis of it. In the first two cases the mucilage 

is called cell-content mucilage; in the last two, membrane mucilage.

VEGETABLE CYTOLOGY 9 I 

Mucilage is stored as reserve food in the tubers of Salep and 

many other Orchids and also in the seeds of some species .of the 

Leguminosae. 

Cell-content mucilage has been found in the leaves of Alos, 

the rhizomes of Triticum, the bulb scales of Squill and Onion and 

in certain cells of many other Monocotyledons, especially those 

containing rap hides. 

Membrane mucilage has been observed in Barosma, Ulmus, 

Althaa, Linum, Astragalus, and Acacia species, in the Blue-green 

Algae, and many of the Brown and Red Algae. 

When mucilage is collected in the form of an exudate from shrubs 

and trees it constitutes what is termed a gum. Many of these gums 

are used in pharmacy, medicine and the arts. The three most im- 

portant from a pharmaceutical standpoint are: Acacia, yielded by 

Acacia Senegal and other species of Acacia; Tragacanth, yielded by 

Astragalus gummifer and other Asiatic species of Astragalus; and 

Cherry Gum, obtained from Prunus Cerasus and its varieties. 

Mucilage may be demonstrated in plant tissues containing it by 

placing sections of these in a deep blue solution of methylene-blue 

in equal parts of alcohol, glycerin and water on a glass slide, allowing 

them to remain in the solution for several minutes, then draining 

off the stain and mounting in glycerin. Those cells containing muci- 

lage will exhibit bluish contents. 

21. Fixed Oils and Fats. These are fatty acid-esters of glycerin 

which are found in the vacuoles of cells or formed with the cell 

walls from which they may be liberated as globules upon treating 

sections with chloral hydrate or sulphuric acid or heating them. 

They are quite soluble in ether, chloroform, benzol, acetone and 

volatile oils but insoluble in water, and, with the exception of castor 

from the 

oil, insoluble in alcohol. They are readily distinguished 

volatile oils in that they leave a greasy stain upon paper which does 

not disappear. Fixed oils and fats take a brownish to black color 

with osmic acid, a red color with alkannin or Sudan III and a blue 

color with cyanin. In Vaucheria, the Diatoms and a few of the 

other Thallophytes, fixed oil is formed in the chromatophores in- 

stead of starch as the first visible product of photosynthesis. In 

higher plants it is generally found in storage regions, 

such as the

0,2 


parenchyma of seeds, fruits and the medullary ray cells and chymaparen- 

of barks, roots and rhizomes. 

22. Volatile Oils. These are volatile odoriferous principles found 

in various parts of numerous plants which arise either as a direct 

product of the protoplasm or through a decomposition of a layer of 

the cell wall which Tschirch designates a " resinogenous layer." 

They are readily distilled from plants, together with watery vapor, are 

slightly soluble in water, but very soluble in fixed oils, ether, chloroform, 

glacial acetic acid, naphtha, alcohol, benzin and benzol. 

They leave a spot on paper which, however, soon disappears. They 

respond to osmic acid, alkannin, Sudan III, and cyanin stains 

similar to the fixed oils and fats. 

Volatile oils may be grouped into four classes: 

A. Pineries or Terpenes, containing carbon and hydrogen and 

having the formula of doHi 6 . Examples: Oil of Turpentine and 

various other volatile oils occurring in coniferous plants. 

B. Oxygenated oils, containing carbon, hydrogen and oxygen. 

Oil of cassia and other cinnamons. 

Examples: 

C. Nitrogenated oils, containing carbon, hydrogen and oxygen 

with nitrogen (from HCN). Example: Oil of Bitter Almonds. 

D. Sulphurated oils, containing carbon, hydrogen and sulphur. 

Example: 

Volatile oil of mustard. 

23. Resins, Oleoresins, Gum Resins, and Balsams. These substances 

represent products of metabolism in many plants which are 

formed either normally as Turpentine, Asafcetida, Mastiche, etc., 

or as a result of pathological processes through injury to the plant 

tissues as Styrax, Benzoin, Balsam of Tolu and Peru, etc. They 

occur usually in special cavities such as secretion cells, glands, or 

secretion reservoirs. 

Resins are insoluble in water but mostly soluble in alcohol. They 

combine with alkalies to form soap. Many of them are oxidized 

oils of plants. Examples: Guaiacum, Resina. 

Oleoresins are mixtures of oil and resin. Examples: Terebin- 

thina, Terebinthina Canadensis. 

Gum resins are natural compounds of resin, gum and oil. Ex- 

amples: Asafcetida, Myrrha, Cambogia.


Balsams are mixtures of resins with cinnamic or benzoic acid or 

both and generally a volatile oil. Examples: Balsamum Tolu- 

tanum, Styrax, Balsamum Peruvianum. 

If sections of a resin containing plant part are placed in a saturated 

aqueous solution of copper acetate for a week or two and mounted 

in dilute glycerin, the resin will be stained an emerald green. 

24. Pigments. These are substances which give color to various 

plant parts in which they are found. They occur either in special 

protoplasmic structures, as chloroplasts, chromoplasts or chromatophores, 

or dissolved in tjae cell sap. Of the pigments named 

the following will be considered: Chlorophyll, Xanthophyll, 

Chromophyll, Etiolin, Anthocyanin, Phycocyanin, Phycophaein, 

and Phyccerythrin. 

Chlorophyll is the yellowish-green pigment found in the chloro- 

plastids or chromatophores of leaves or other green parts of plants. 

Its composition is not definitely known although it yields products 

similar to the haemoglobin of the blood when decomposed. Iron 

is known to be essential to its formation. If an equal portion of 

xylene be added to a fresh alcoholic solution of chlorophyll and the 

mixture shaken, the chlorophyll in solution will break up into a 

yellowish and greenish portion. The greenish portion dissolves in 

the xylene which rises forming the upper stratum, while the yellowish 

portion dissolves in the alcohol forming the lower stratum. To this 

isolated greenish portion of chlorophyll has been given the name of 

" " 

chlorophyllin while the yellowish portion has been designated 

"xanthophyll." 

Chlorophyllin when examined spectroscopically produces absorption 

bands in the red, orange, yellow and green of the spectrum, the 

broadest and most distinct band being in the red. 

Chromophyll also called " xanthophyll" and "carotin" is the yellow 

or orange pigment found in chromoplastids. By some the term 

carotin is limited to the orange pigment found in the carrot. Sul- 

phuric acid forms a blue color with chromophyll. 

Etiolin is a pale yellow pigment which appears when green plants 

are kept for some time in darkness. It is probably identical with 

xanthophyll.

94 


Anthocyanins are applied to the blue, purple and red pigments 

which occur in the cell sap. The character of the color is claimed to 

be due to the alkalinity or acidity of the cell sap. 

Phycocyanin is the blue pigment found in the blue-green algae, 

associated with chlorophyll. It is soluble in water. 

Phycophaein is the brown pigment found in the brown algae. 

Phycoerythrin is the red pigment found in many of the red algae. 

The last two are always associated with chlorophyll but frequently 

conceal it. 

25. Latex. This is an emulsion of varying composition and color 

found in special passages, as latex cells and laticiferous vessels of 

many plants. It may contain starch, sugar, proteid, oil, enzymes, 

tannins, alkaloids, gum, resins, caoutchouc and mineral salts. The 

color may be absent as in Oleander; whitish as in Asclepias, Papaver, 

Hevea, and Apocynum; yellowish to orange as in Celandine, or red as 

in Sanguinaria. 

Chlor-zinc-iodine solution imparts to latex a wine red color. 

The latex of the following plants is of value to pharmacy and 

the arts: 

Papaver somniferum and its variety album which yields Opium. 

That from the unripe capsules is alone used for this drug. 

Palaquium Gutta which yields Gutta Percha. 

Hevea species, Ficus elastica, Landolphia species, Castittoa elastica, 

Hancornia speciosa, Forsteronia species, Funtumia elastica and F. 

africana, Manihot species, Clitandra species and various species of 

Euphorbia furnish most of the Rubber of commerce. 

Lactuca mrosa and other species of Lactuca yield the drug Lactu- 

carium. 

26. Enzymes. An enzyme or ferment (according to Hepburn) 

is a soluble organic compound of biologic origin functioning as a 

thermolabile catalyst in solution. Ostwald has defined a catalyst 

as an agent which alters the rate of a reaction without itself entering 

into the final product, 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. The biologic catalysts (enzymes)


differ from the inorganic catalysts in that they are sensitive to heat 

and light. According to Haas and Hill they are destroyed at iooC. 

and most of them cannot be heated safely above 6oC. Enzymes 

are soluble in water, glycerin or dilute saline solutions. They are 

stimulated to activity by substances known as "activators" and their 

activity is checked by other substances called " paralyzers ." Frequently 

the paralyzers consist of products of enzyme action. Cold 

inhibits and warmth accelerates enzyme action. Moisture must 

always be present for enzymic activity. 

CLASSIFICATION OF ENZYMES 

A. According to Di/usibility through Cell Wall. 

Endocellular : Those that cannot diffuse out of the cell. Example : 

Zymase 

of Yeast. 

Extracellular: Those that can diffuse out of the cell. Example: 

Invertase of Yeast. 

B. According to Kind of Substances Acted upon and Transformed. 

i. Carbohydrate enzymes: 

Diatase found in the germinating seeds of barley and 

other grains and in Aspergillus oryza, etc., converts starch to 

maltose and dextrin. 

Invertase, secreted by yeasts, and found in younger parts 

of higher plants, transforms cane sugar, producing dextrose and 

levulose. 

Maltase, found in malt and Saccharomyces octosporus, transforms 

maltose to dextrose. 

Trehalase, found in Polyporus, hydrolyzes trehalose to dextrose. 

Cytase, found in Nux Vomica seeds, in barley, dates, etc., decom- 

poses hemicellulose and cellulose to galactose and mannose. 

Lactase, found in Kephir grains, hydrolyzes lactose to dextrose 

and galactose. 

Inulase, found in Compositaceous plants, transforms inulin to 

levulose. 

Zymase, found in yeast, hydrolyzes glucose (dextrose and levulose) 

to alcohol and carbon dioxide.

96 

2. Fat and Oil Ferment : 


It is 

Lipase splits up fats and oils into fatty acids and glycerin. 

found m various mildews, molds and numerous oily seeds and other 

fatty-oil storage regions of higher plants. 

3. Proteinaceous Ferments : 

Pepsin converts proteids into proteoses and peptones. 

Trypsin, found in yeast, Boletus edulis, Amanita species, etc., 

resolves proteins to peptones and amino-acids. 

Bromelin, found in the fruit of the Pineapple and Papayin (Pa- 

pain), found in the latex of the fruit of the Papaw, act similarly to 

trypsin. 

Nepenthin, found in the pitchers of Nepenthes species, acts simi- 

larly to pepsin. 

4. Glucoside Ferments : 

Emulsin (synaptase), found in the seeds of the Bitter Almond, 

Cherry Laurel leaves, in the barks of the Wild Black Cherry and 

Choke Cherry and in other Rosaceous plant parts, in Manihot 

utilissima, 'Polygala species, etc., hydrolyzes the glucoside present 

(either amygdalin or 1-mandelonitrile glucoside) to hydrocyanic 

acid, benzaldehyde and glucose. 

Myrosin (myronase), found in the seeds of Brassica nigra and 

other members of the Crucifera, converts the glucoside, Sinigrin, 

into ally-iso-sulphocyanide and glucose. 

Rhamnase, found in Rhamnus Frangula and probably other species 

of Rhamnus, hydrolyzes the glucoside frangulin to rhamnose and 

emodin. 

Gaultherase, found in Gaultheria procumbens and other Ericaceous 

plants, resolves the glucoside, gaultherin, to methyl-salicylate 

and glucose. 

CELL WALLS 

The cell walls of plants make up the plant skeleton. They are all 

formed by the living contents of the cells (protoplasts) during cell- 

divisions. In most plants the cell wall when first formed consists of 

cellulose, (CeHioC^n, a carbohydrate, or closely 

allied substances. 

It may remain of such composition or become modified to meet cer-


tain functions required of it. Thus, in the case of outer covering 

cells as epidermis and cork, whose function is that of protecting the 

underlying plant units, 

the walls become infiltrated with cutinand 

suberin, waxy-like substances, which make them impermeable to 

water and gases, as well as protect them against easy crushing. 

Again, in the case of stone cells and sclerenchyma fibers whose 

function is that of giving strength and support to the regions wherein 

found, the walls become infiltrated with Hgnin which increases their 

strength, hardness, and in the case of sclerenchyma fibers, their 

elasticity also. Moreover, in the case of the cells comprising the 

testa or outer seed coat of the pumpkin, squash, mustard and flax, 

etc., whose function is that of imbibing quantities of water, the walls 

undergo a mucilaginous modification. 

Growth in Area and Thickness. The cell wall when first formed 

is limited in both extent and thickness. As the protoplast within 

enlarges new particles are placed within the wall by the process called 

intussusception. This increases its area. New particles, also, are 

deposited on its surface which gradually increases its thickness. 

The latter process is known as growth by apposition.


VARIOUS KINDS OF CELL WALLS AND BEHAVIOR or EACH TO MICRO- 

CHEMIC REAGENTS 

Nature of wall Where found Reagent and behavior toward same 

Cellulose. 

Lignocellulose 

(Lignified wall). 

Reserve cellulose 

Mucilaginous 

modification of 

cellulose. 

Suberized walls. . 

Cutinized walls.. 

Callus of sieve 

plates. 

Silicified walls . . 

Parenchyme cells, trichomes 

such as 

cotton, etc. 

Wo o d y parts of 

plants, 

such as stem 

cells, bast fibers, 

wood fibers, etc. 

Found in certain seeds 

such as nux vomica, 

ignatia, ivory nut, date 

coffee, etc. 

In various parts of 

plants. 

In cork, wounded 

areas of plants, endo- 

dermis. 

Forming outer walls 

of many epidermal 

cells. 

Plates of sieve tubes. 

Epidermis of Equi- 

setacese, Gramineae, 

etc.; Diatoms. 

Cuoxam dissolves it. Chlorzinc- 

iodine solution imparts a blue or vio- 

let color. Iodine solution followed 

by sulphuric acid colors it blue. 

Phloroglucin with HC1 imparts a red 

color except to bast fibers of flax. 

Corallin-soda solution imparts pink 

color. Aniline sulphate with H2SO4 

colors it a golden-yellow. 

iodine imparts a yellow color. 

As for cellulose. 

Chlorzinc- 

Alcoholic or glycerin solution of meth- 

ylene-blue imparts a blue color. 

Alcoholic extract of chlorophyll, in the 

dark, imparts a green color. Alcannin 

and Sudan III impart a red colora- 

tion. Converted into yellowish droplets 

and granular masses upon heating 

with a strong solution of KOH. 

Sulpuhric acid is resisted. 

As for suberized walls. 

Corallin-soda solution imparts pink 

color. 

Soluble in hydrofluoric acid.

CHAPTER VI 

PLANT TISSUES 

A tissue is an aggregation of cells of common source, structure 

and function in intimate union. 

THE TISSUES OF SPERMATOPHYTES AND PTERIDOPHYTES 

The tissues of seed plants and pteridophytes are all derived from 

a fertilized egg (oospore) which has undergone repeated divisions. 

At first either an apical cell arises or a mass of cells is formed which 

are essentially alike, but gradually we find that a division of labor has 

become operative setting aside many different groups of cells, each 

group of which has its particular role to perform in the economy of 

the whole. Each group of cells similar in source, structure and 

function is called a tissue. The tissues found in higher plants range 

from those whose component cells are more or less rounded, in a 

rapid state of division, and whose thin cellulose cell walls enclose 

a mass of protoplasm, devoid of vacuoles, or with exceeding small 

ones to those whose cells through various physical and chemical 

factors become compressed, elongated, and highly modified in respect 

to their contents and walls. 

As was shown by Hanstein, 1 the embryo of Angiosperms, while 

still constituted of only a few cells in the process of division, becomes 

differentiated into three layers of cells which differ in their arrangement 

and direction of division; these were called by him, Derma- 

togen, Periblem and Plerome. In roots a fourth layer of cells is 

sometimes evident at the apex. This was termed by Janczewski 2 the 

Calyptrogen layer. These primary layers or groups of cells are 

called primary meristems or generative tissues. They are composed 

1 

Hanstein, "Die Scheitelzellgruppe im Vegetationspunkt der Phanerogamen," 

Bonn, 1868. 

2 Am. Sci. Nat. 5 serie, torn. xx. 

99


of more or less rounded cells having delicate cell walls of cellulose 

which enclose protoplasm and nucleus and wherever found in living 

embryos are in a rapid state of division. 

The generative tissues are found in the growing apices of plant 

organs, such as root, stem and leaf apex. By the division and redivi- 

sions of their cells they give rise to the mature or adult tissues of 

plants. 

1. Dermatogen originates epidermal 

tissue and derivative struc- 

tures such as stomata, non- glandular and glandular hairs, glands, 

and cork cambium. 

2. Periblem originates cortex tissue, chlorophylloid cells (chlor- 

enchyma) colloid cells (collenchyma), strengthening cells (scleren- 

chyma), crystal cells (raphiderchyma) latex cells (lacterchyma), 

endodermis and cork cambium. 

3. Plerome originates fibre-vascular bundles, fundamental tissue, 

pericambium and cambium. 

According to structure the following tissues are found in various 

forms of higher plants: 

1. Meristem 7. Cork 

2. Parenchyma 8. Laticiferous tissue 

3. Collenchyma 9. 

Cribiform or sieve tissue 

4. Sclerenchyma 10. Tracheary tissue 

5. Epidermis n. Medullary rays 

6. Endodermis 

MERIS1EM 

Meristem, frequently called embryonic tissue, 

tissue composed of cells -in the state of rapid division. It is found 

is undifferentiated 

in the growing apices of roots, stems and leaves and is in these 

regions called primary meristem, since it is the first meristem to 

appear. Such meristem gives rise to the permanent or mature 

tissues of plants and retains the power of independent growth and 

capacity for division as long as the plant part 

survives which con- 

tains it. Meristem is also found in other regions of plant organs 

and is there 

such as the cambium, cork cambium and pericambium 

called secondary meristem. Secondary meristem loses with its de- 

velopment the power of division and independent growth.

PLANT TISSUES IOI 

PARENCHYMA 

Parenchyma or Fundamental Tissue is the soft tissue of plants, 

consisting of cells about equal in length, breadth and thickness 

(isodiametric) with thin cellulose cell walls enclosing protoplasm and 

a nucleus and frequently substances of a non-protoplasmic nature. 

There are four generally recognized types of parenchyma, viz.: 

Ordinary Parenchyma (Soft Ground Tissue, Fundamental Tis- 

sue). Next to the meristem this is the least modified of all plant 

tissues. It is generally composed of thin-walled cells, commonly 

polyhedral or spheroidal in form and often of approxinately the 

same length, breadth, and thickness (isodiametric), the cell walls are 

composed of cellulose which is usually unmodified. Occasionally 

the outline of the cells is star- shaped, as in the Wood Rush or Pick- 

erel Weed or the cells may be several times as long as wide, as in 

Pelargonium, etc. Moreover, markings may occur on the walls. 

These may be of the nature of pores, as in the parenchyma cells of 

the pith of the Elder or Sassafras, annular or reticulate thickenings, 

as in the Mistletoe, or spiral thickenings, as in certain Orchids. 

Protoplasm and a nucleus are always present, but in old cells are 

only seen as a thin layer pushed up against the cell wall. Ordinary 

Parenchyma may be seen composing the soft tissues of roots, stems, 

and barks. 

Assimilation Parenchyma (Chlorophyll or Chromophyll Parenchyma, 

Chlorenchyma). This form of parenchyma tissue is found 

in foliage leaves, floral leaves, in the outer region of young green 

stems and fruits. Its cells are thin walled and vary in shape from 

more or less isodiametric to irregular and elongated forms. The 

cells always contain chloroplasts or plastids, in whose pores may be 

found some other coloring substance. 

Conducting Parenchyma. This type of .parenchyma functions 

in the rapid translocation of food materials to distant regions in .the 

plant. It includes the wood parenchyma cells of the xylem which 

convey a portion of the crude sap (water with mineral salts in 

solution) and the phloem parenchyma (soft bast) which transports 

the elaborated sap (carbohydrate and proteid material in solution). 

Conducting parenchyma cells differ from those of ordinary paren-

102 


FIG. 40. Transverse section of part of leaf-stalk of a begonia, e. Epidermis; 

c, cuticle; B, collenchyma, with walls thickened* at the angles v, chl, chloroplasts. 

(Sayre after Vines.) 

FIG. 41. Stone cells from different sources. I. From coffee; 2, 3, and 4, 

from stem of clove; 5 and 6, from tea leaf; 7, 8, and 9, from powdered star-anise 

seed. (Stevens, after Moeller).

PLANT TISSUES 103 

chyma in being usually more elongated and in conducting soluble 

food materials with greater celerity. 

Reserve Parenchyma. This resembles ordinary parenchyma in 

many particulars of structure but differs from it mainly by its cells 

being filled with starch, protein crystals, or oil globules. It is 

usually found in seeds, fleshy roots, or underground stems 

tubers, corms, and bulbs. 

such as 

Collenchyma. This form of tissue is characterized by 

its cells 

being prismatic, more elongated than ordinary parenchyma, and 

thickened in their angles with a colloidal substance. The cells, like 

those of parenchyma tissue contain piotoplasm and a nucleus, and 

frequently chloroplasts (Fig. 40). Collenchyma is generally found 

underneath the epidermis, and gives strength to that tissue. It is 

frequently observed forming the "ribs" of stems and fruits of the 

Umbellifera and "ribs" of stems of the Labiata. In many leaves 

it has been found as the supporting and strengthening tissue between 

the stronger veins and the epidermis. 

Sclerenchyma or stony tissue comprises a variety of supporting 

elements having thickened cell walls composed of lignocellulose. 

When first formed these cells resemble those of ordinary parenchyma 

in having walls of pure cellulose, but later lignin becomes deposited 

on the inner surface of the walls in one or more layers. (Occasionally 

as in the rhizomes of Ginger no lignin is deposited on the walls 

of the sclerenchyma fibers). When sclerenchyma is composed of 

cells which are more or less isodiametric or moderately elongated, 

with thickened lignified walls and conspicuous pores, its elements 

are called Stone Cells. Stone cells are distributed in fruits, seeds 

and barks of many plants, rarely in woods. They have been found 

forming the gritty particles in the "flesh" of certain fruits as the 

Pear, the endocarp or stone region of drupaceous fruits as the Olive, 

Peach, Cubeb, Pepper, etc., the hard portions of seed coats as in 

Physostigma, Walnuts, etc. Each stone cell presents for examination 

a cell wall of cellulose with one or several layers of lignin on its 

inner surface which surround a central lumen. The latter is in 

communication with radial pore canals leading outward to the 

middle lamella. Longitudinal pore canals are also evident. 

When sclerenchyma is composed of cells which are greatly elon-

IO4 


FIG. 42. Stone cells from various sources. I, From olive pit; 2, from cocoanut 

endocarp; 3, from flesh of pear; 4, from aconite root; 5, from capsicum; 6. 

from hazelnut; 7, from allspice. (Drawing by Hoffstein.)


gated and more or less obtusely or taper ended, its component elements 

are termed Scierenchyma fibers. These fibers are frequently 

spindle-shaped, contain air and exhibit oblique slits in their walls. 

They are either polygonal, rectangular or somewhat rounded in 

FIG. 43. Scierenchyma fibers from different sources. I, From powdered 

cinnamon bark; 2, End of bast fiber of flax stem showing transverse markings 

(6); 3, middle portion of flax fiber showing characteristic cross markings at 6; 4, 

bast fiber from cinchona bark; 5, branched bast fiber from choke cherry bark; 

6, above, end, and below, median portion of bast fiber of jute. All highly mag- 

nified. 

transverse section. They occur in various parts of roots, stems, 

leaves, fruits and seeds as supporting elements. When Scierenchyma 

fibers occur in the xylem region of fibro-vascular bundles they are 

termed Wood Fibers; when they appear in the ph"o2m region, 

Bast Fibers.


EPIDERMIS 

Epidermis is the outer covering tissue of a plant and is protective 

in function. Its cells may be brick-shaped, polygonal, equilateral or 

wavy in outline. Their outer walls are frequently cutinized (infiltrated 

with a waxy-like substance called cutin) . Among the epidermal 

cells of leaves and young green stems may be found numerous 

pores or stomata (sing, stoma) surrounded by pairs 

of crescent- 

Pic. 44. Upper epidermis of Comptonia asplenifolia leaf (surface view) showing 

epidermal cells and two non-glandular trichomes. 

shaped cells, called guard cells. The stomata are in direct communication 

with air chambers beneath them which in turn are in 

communication with intercellular spaces of the tissue beneath. The 

function of the stomata is to give off watery vapor and take in or 

give off carbon dioxide, water and oxygen. In addition to stomata 

some leaves possess groups of water stomata which differ from trans- 

piration stomata in that they always remain open, are circular in out- 

line, give off water in droplets directly, and lie over a quantity of 

small-celled glandular material which is in connection with one or 

more fibro-vascular bundles. Examples: Leaves of Crassula, Saxi- 

Jraga and Ficus.

PLANT TISSUES 

3 

FIG. 45. Trichomes from different sources, i, Unicellular non-glandular 

trichomes as seen growing out of epidermal cells of Senna; 2, uniseriate nonglandular 

trichomes of Digitalis; 3, unicellular stellate trichomes from Deutzia 

scabra; 4, unicellular twisted trichomes from lower epidermis of Eriodictyon; 5, 

clavate non-glandular trichomes from scraping of epidermis of the fruits of 

Rkus glabra; 6, 2-branched trichomes of Hyoscyamus muticus, a substitute for 

Heixbane; 7, branched multicellular trichome of Marrubium; 8, glandular trichomes 

from strobile of Humulus (Lupulin); 9,' glandular trichomes from leaves 

of Digitalis purpurea; 10, aggregate, non-glandular trichomes of Kamala; n, 

lateral view (to left) and vertical view (to right) of glandular trichomes of 

Kamala; 12, vertical view (above) and profile view (below) of 8-celled glandular 

hair from Mentha piper ita. All highly magnified. 

107

io8 PHARMACEUTICAL BOTANY 

The epidermis of leaves, stems, fruits, and seeds of many plants 

frequently give rise to outgrowths in the form of papillae, hairs 

and scales. Epidermal papilla are short protuberances of epidermal 

cells. They may be seen to advantage on the upper epidermis of the 

FIG. 46. FIG. 47. 

PIG. 46.' i, Epidermis of oak leaf ; 2, epidermis of Iris leaf, both viewed from 

the surface; 3, group of cells from petal of Viola tricolor; 4, two epidermal cells 

in cross-section showing thickened outer wall differentiated into three layers, 

namely, an outer cuticle, cutinized layer (shaded), and an inner cellulose layer; 

5 and 6, epidermal outgrowths in the form of scales and hairs, (i, 2, 6 after 

Stevens, 3 after Strasburger, 4 after Sachs, and 5 after de Bary.) 

PIG. 47. Different forms of epidermal outgrowths, i, Hooked hair ffom 

Phaseolus multiflorus; 2, climbing hair from stem of Humulus Lupulus; 3, cod- 

Hke wax coating from the stem of Saccharum officinarum; 4," climbing hair of 

Loasa hispida; 5, stinging hair of Urtica urens. (Fig. 3 after de Bary; the remainder 

from Haberlandt.) 

ligulate corolla of various species of Chrysanthemum, on the lower 

epidermis of the foliage leaves of species of Erythroxylon and upon the 

upper epidermis of the petals of the Pansy (Viola tricolor}. Epidermal 

hairs or trichomes are more elongated outgrowths of one 

or more epidermal cells. They may be unicellular (Cotton) or 

5

PLANT TISSUES IOQ 

multicellular, non-glandular (simple) or glandular. The non- 

glandular hairs may be of various shapes, viz. : clavate (club-shaped) 

as on Rhus glabra fruits', stellate for star-shaped) as on Deutzia 

leaves; candelabra-shaped, as on Mullein leaves; filiform as on 

Hyoscyamus, Belladonna and Digitalis leaves; hooked, as on stems 

of Phaseolus multiflorus or Hops; barbed, as on the stems of Loasa 

species; or tufted, as found on the leaves of Marrubium vulgar e. 

They may be simple as in Cotton, etc., or branched as in Hyoscyamus 

muticus. 

The glandular hairs comprise those whose terminal cell or cells are 

modified into a more or less globular gland for gummy, resinous or 

oily deposits. They are generally composed of a stalk and a head 

region although rarely the stalk may be absent. The stalk may be 

unicellular, bicellular or uniseriate (consisting of a series of superimposed 

cells). The head varies from a one- to many-celled struc- 

ture. The drug Lupulin consists of the glandular hairs separated 

from the strobiles of Humulus lupulus. 

Scales are flat outgrowths of the epidermis composed of one or 

several layers of cells. They occur attached to the stipes of Aspid- 

ium, Osmunda and other ferns, where they are called "chaff scales." 

They are also found on a number of higher plants. 

Plant hairs are adapted to many different purposes. They may 

absorb nourishment in the form of moisture and mineral matter 

in solution, e.g., root hairs. Those which serve as a protection to the 

plant may be barbed and silicified, rendering them unfit for animal 

food, or,-as in the nettle, charged with an irritating fluid, penetrating 

the skin when touched, injecting the poison into the wound. A 

dense covering of hairs also prevents the ravages of insects and the 

clogging of the stomata by an accumulation of dust. They 

important office in the dispersion of seeds and fruits, as with their 

aid such seeds as those of the milkweed and Apocynum are readily 

scattered by the wind. 

The reproductive organs of many Cryptogams are modified 

hairs, as the sporangia of Ferns. 

ENDODERMIS 

fill an 

Endodermis is the "starch sheath" layer of cells, constituting 

the innermost layer of the cortex. In Angiospermous stems it


usually resembles the other parenchyma layers of cortex as to structural 

characteristics, save that it frequently contains more starch. 

In fern stems, roots of Monocotyledons and of Dicotyledons of 

primary growth, however, its cells are clearly distinguished from the 

other cells of the primary cortex by their elongated form and suberized 

(occasionally lignified) radial walls. In the roots of Mexican 

Sarsaparilla the inner as well as the radial walls are suberized; in 

those of the Hcnduras variety, inner, radial and outer walls all show 

suberization. Endodermal tissue is devoid of intercellular-air-spaces. 

Its cells contain protoplasm and nucleus. Its functions seem to be 

to give protection to the stele (tissues within it) and to reduce per- 

meability between primary cortex and stele. 

CORK 

Cork or suberous tissue is composed of cells of tabular shape, 

whose walls possess suberized layers. Its cells are mostly filled 

with air containing a yellow or brownish substance. It is derived 

from the phellogen or cork cambium which cuts off cork cells out- 

wardly. Cork tissue is_ devoid of intercellular-air-spaces. It 

forms a protective covering to the roots of secondary growth, stems 

(after the first season) of Dicotyledons and Gymnosperms, and 

wounds of stems and branches. Living cork cells contain protoplasm 

and cell sap while dead cork cells are filled with air. 

The walls of cork cells resist the action of concentrated sulphuric 

acid. They are colored green, when in contact with alcoholic 

extract of chlorophyll for several days in the dark. 

LATICIFEROUS TISSUE 

This form of tissue comprises either latex cells, laticiferous vessels, 

or secretory cells differing from each other in origin and method of 

development. Latex cells are elongated tubes which take their' 

origin from meristematic cells of the embryo. Elongating with the 

growth of the plant, they branch in various directions and traverse 

at maturity all of its organs. Such cells are abundant in the 

following families: Apcoynacece, Asclepiadacea, Urticacea and 

Euphorbiacea.

PLANT TISSUES III 

Laticiferous vessels are long simple or branching tubes, which owe 

their origin to chains of superimposed cells whose transverse walls 

PIG. 48. Laticiferous vessels from"the"cortex of 

rootofScorozonorahispanica. 

A, As seen under low power, and B, a smaller portion under high power. (Stevens, 

after Sachs.) 

have early become absorbed, the lumina of the cells then becoming 

filled with latex; They are found in various parts of roots, stems, 

and leaves. When branched the branches Connect with those of


other tubes forming anastomosing 

network. These vessels occur 

in the following families: Composites, Papaveracea, Campanulacea, 

Convolvulacea, Euphorbiacea, Aracea, Oleacea, Geraniacea, and 

Musacea. 

Secretory cells with a latex-like content are probably of secondary 

oiigin in plants. They resemble in many respects latex cells and are 

seen in various species of the Celastracea, Urticacea, Tiliacece, and 

Oleacea families. 

All laticiferous elements contain a colorless, milky-white, or 

otherwise colored emulsion of gum-resins, fat,, wax, coautchouc and 

in some cases, alkaloids, tannins, salts, ferments, etc. This emulsion 

is called "latex." 

SIEVE (LEPTOME OR CRIBIFORM) TISSUE 

This tissue found in the phloem (rarely in the xylem) region of 

nbro-vascular bundles consists of superimposed, elongated, tubular 

cells whose longitudinal walls are thin and composed of cellulose and 

whose transveise walls, called "sieve plates," are perforated, permitting 

of the passage of proteids from one cell to another. Occasionally 

sieve plates are formed on the longitudinal walls. Sieve 

tubes are usually accompanied by companion cells excepting in 

Pteridophytes and Gymnosperms. Both companion cells and sieve 

tubes arise by the division of the same mother-cell. The companion 

cells may be distinguished from the sieve tubes by their abundant 

protoplasmic contents, and also by the fact that they retain their 

nuclei after complete maturation. Besides sieve tubes, companion 

cells, and bast fibers, parenchyma cells are often found in the 

phloem. 

TRACHEARY TISSUE 

The tracheary tissue of plants comprises two kinds of elements, the 

trachea (ducts or vessels) and tracheids. Both of these conduct 

crude sap (water with mineral salts in solution). The trachea are 

very long tubes of a cylindrical or prismatic shape which are formed 

by the disintegration of the transverse walls between certain groups 

of superimposed cells, during the growth of the plant. The tubes 

frequently retain some of, their transverse walls. The longitudinal

PLANT TISSUES 

walls of these tubes are of varying thickness, usually, however, 

thinner than those of woody fibers. The thickness is due to an infil- 

d b 

B 

FIG. 49. 

c d b 

6 C 

FIG. 50. 

o v o 

y$fi 

OOoC 

00 

FIG. 49. Stages in the development of sieve tubes, companion cells, and 

phloem parenchyma. A, a. and b, Two rows of plerome cells; in c and d, a has 

divided longitudinally and c is to become companion cells; d, a sieve tube, and 

b, phloem parenchyma. B, c, Companion cells, and d, a beginning sieve tube 

from c and d, respectively in>4. The cross-walls in d are pitted; b, phloem parenchyma 

grown larger than in A. C, The same as B with the pits in the crosswalls 

of the sieve tubes become perforations, and the nuclei gone from the cells 

composing the tube. (From Stevens.) 

FIG. 50. Vascular elements. A, annular tracheal tube; B, spiral trachea 

tube; C, reticulated tracheal tube; D, pitted tracheal tube; E, cross-section 

through plate of seive tube, and adjoining companion cell; F, length-wise section 

of sieve tube; G, portions of two companion cells. (A, B, C, D, Robbins; E, F, 

and G, after Strasburger.) 

tration of lignin upon the original cellulose wall, 

characteristic thickenings on their inner surfaces. 

8 

The walls show


FIG. 51. Stages in the development of the elements of the xylem. A, progressive 

steps in the development of a tracheal tube, i, Row of plerome or 

cambial cells that are to take part in the formation of a tube; 2, the same at a 

later stage enlarged in all dimensions; 3, the cells in 2 have grown larger, their 

cross-walls have been dissolved out, and the wall has become thickened and 

pitted; 4, the walls in 3 have become more thickened, the pits have an overhanging 

border, the walls have become lignified as indicated by the stippling, and 

finally the protoplasts have disappeared, and the tube is mature and dead. B, 

Stages in the formation of tracheids from plerome or cambial cells. The steps

PLANT TISSUES 

Tracheae are classified according to their markings as follows 

Annular, with ring-like thickenings. 

Spiral, with spiral thickenings. 

Reticulate, with reticulate thickenings. 

PIG. 52. Closed collateral bundle of stem of Zea mays. VG, Bundle sheath; 

L, intercellular space; A, ring from an annular tracheal tube; SP, spiral tracheal 

tube; M, pitted vessels; V, sieve tubes; S, companion cells; CP, crushed primary 

sieve tubes; F, thin-walled parenchyma of the ground or fundamental tissue. 

(From Sayre after Strasburger.) 

Porous or pitted with spherical or oblique slit pores. 

Annulo- spiral, with both ring and spiral thickenings. 

Scalariform, with ladder-like thickenings. 

are the same as in A, excepting that the cross-walls remain and become pitted. 

C, steps in the development of wood fibers from cambial cells, i, Cambial cells; 

2, the same growth larger in all dimensions with cells shoving past each other 

as they elongate; 3, a later stage with cells longer and more pointed and walls 

becoming thickened and pitted; 4, complete wood fibers with walls more thickened 

than in the previous stage and lignified, as shown by the stippling. The 

protoplasts in this last stage have disappeared and the fibers are dead. D, steps 

in the formation of wood parenchyma from cambial or procambial cells, i, 

Group of cambial or pierome cells; 2, the same enlarged in all dimensions; 3, the 

same with walls thickened and pitted; 4 and 5 show the same stages as 2 and 3, 

but here the cells have enlarged radially or tangentially more than they have 

vertically. The walls of these cells are apt to become lignified, but the cells are 

longer lived than the wood fibers. (From Stevens.)

Il6 RHARMACEUTICAL BOTANY 

Tracheids are undeveloped ducts having bordered pores and fre- 

quently scalariform thickenings. Like tracheae their walls give the 

characteristic lignin reaction with phloroglucin and HC1. The 

bordered pores of coniferous tracheids (Fig. 77) exhibit a wall 

surrounding the pore which forms a dome shaped protrusion into 

the cell. Like tracheae, also, tracheids convey water with mineral 

salts in solution. Tracheids and medullary rays make up most of 

the wood of Conifers. 

PIG. 53. 

Transverse section of a concentric bundle from the rhizome of Iris 

(a monocotyledon). Xylem surrounding the phloem. /, Tracheae; f 1 , protoxylem; 

s, sieve. tubes; g, companion cells of the internal phloem portion. (From 

Sayre after Vines.) 

MEDULLARY RAYS 

These are bands of parenchyma cells which extend radially from 

the cortex to the pith (primary medullary rays) or from a part of the 

xylem to & part of the phloem (secondary medullary rays). In 

tangential-longitudinal sections they usually appear spindle shaped 

'while in radial-longitudinal sections they are seen crossing the other 

elements. Their primary function is to supply the cambium and 

wood with elaborated sap formed in the leaves and conveyed away 

by the sieve tubes, and phloem parenchyma and to supply the cam-


bium and phloem with crude sap which passes up ma inly through the 

tracheae and tracheids from the absorptive regions of the roots. 

They furthermore serve as storage places for starch, alkaloids, resins, 

and other substances. 

Fibro-vascular Bundles are groups of fibers, vessels and cells cours- 

ing through the various organs of a plant and serving for conduction 

PIG. 54. Diagrams illustrating the arrangement of the regions in different 

types of nbrovascular bundles. In each diagram x represents xylem; P, phloem 

and C, cambium. A, Radial bundle; B, concentric bundle of fern stem type; 

C, concentric bundle of monocotyl type; D, closed collateral bundle; E, open 

collateral bundle; F, bi-collateral bundle. 

and support. According to the relative structural arrangement of 

their xylem and phloem masses they may be classed as follows: 

I. Closed collateral, consisting of a mass of xylern lying alongside 

of a mass of phloem, the xylem facing toward the center, the phloem 

facing toward the exterior. Stems of most Monocotyledons and 

Horsetails. 

II. Open collateral, consisting of a mass of xylem facing toward 

the pith and a mass of phloem facing toward the exterior and sepa- 

rated from each other by a cambium. Stems and leaves of Dicoty- 

ledons and roots of Dicotyls and Gymnosperms of secondary growth. 

III. Bicollateral, characterized by a xylem mass being between 

an inner and an outer phloem mass. There are two layers of cambium 

cells, one between the xylem and inner phloem mass, the other

Il8 PHARMACEUTICAL BOTANY 

between the xylem and outer phloem mass. Seen chiefly in stems 

and leaves of the Cucurbitacece and Solanacece. 

IV. Concentric, characterized by a central xylem mass surrounded 

by a phloem mass or vice versa. No cambium present. 

(a) Concentric, with xylem central in bundle. Seen in stems and 

leaves of nearly all ferns and- the Lycopodiaceos. 

FIG. 55. Cross-section through a portion of a root of A corns calamus. A. 

Cortical parenchyma; B, endodermis; C, pericycle; E, phloem; F, xylem. At 

F, F, are large tracheal tubes, which were formed last, the narrow tubes near 

the periphery of the xylem being formed first. At the center of the root, within 

the circle of the radial vascular bundle, occur thin-walled parenchymatous pith 

cells. (From Sayre after Frank.} 

(b) Concentric, with phloem central in bundle. Seen in stems and 

leaves of some Monocotyledons. Examples'. Calamus and Conval- 

laria rhizomes. 

V. Radial, characterized by a number of xylem and phloem masses 

alternating with one another. Seen in the roots of all Spermato- 

phytes and Pteridophytes.

PLANT TISSUES IIQ 

Xylem is that part of a fibro-vascular bundle that contains wood 

cells and fibers. It may also contain tracheae, tracheids, seldom 

sieve tubes. 

Phloem is that part of a fibro-vascular bundle that contains sieve 

tubes, phloem cells, and often bast fibers. 

SECRETION SACS (SECRETION CELLS) 

These were formerly parenchyma 

cells which sooner or later lost 

their protoplasm and nucleus and became receptacles for oil, resin, 

oleoresin, mucilage or some other secretory substance. They are 

generally found in parenchyma regions of stems, roots, leaves, 

flower or fruit parts and frequently possess suberized walls. Good 

illustrations of these structures may be seen in Ginger and Calamus. 

INTERCELLULAR AIR SPACES 

Intercellular air spaces are cavities filled with air found between 

cells or groups of cells throughout the bodies of higher plants. 

Their 

function is to permit of the rapid movement of atmospheric gases 

through the entire plant body. They are formed either by the 

breaking down of the middle lamella of the cell walls, where several 

cells come together, and a later separation of the cells at these 

places (Schizogenous intercellular- air-spaces), or by a breaking down 

and disappearance of cell walls common to groups of cells (lysigenous 

intercellular-air-spaces). In terrestrial plants which live in middle 

regions (mesophytes) and in desert plants (xerophytes) the intercellular-air-spaces 

are averagely small and more or less angular. In 

plants of swamp or marsh habit they are medium-sized, while in 

those which live entirely in the water (hydrophytes) they are of large 

size and more or less rounded. 

SECRETION RESERVOIRS 

These structures are either found as globular or irregular spaces, 

as in Orange and Lemon Peel and Eucalyptus leaves, containing oil 

or oil and resin when they are called internal glands, or, as tube-like 

such as are found in Pine 

spaces filled with hydrocarbon principles 

leaves and stems, when they sometimes receive the name of secretion


canals. Occasionally they are named according 

to the nature of 

their contents resin or oil canal or reservoir, etc. They are generally 

lined with a layer of cells, usually more or less flattened, which 

are characterized by possessing large nuclei. To this layer has been 

assigned the name " epithelium."- 

PIG. 56. Resin duct (secretion reservoir) in leaf of Pinus silvestris, in cross 

section at A, and in longitudinal section at B; h, cavity surrounded by the secret- 

ing cells; /, /, sclerenchyma fibers surrounding and protecting the duct. (Stevens, 

after Haberlandt. 

Classification of Tissues According to Function. According to 

their particular function, tissues may be classified as follows: 

I. CONDUCTING TISSUES 

II. PROTECTIVE TISSUES 

III. MECHANICAL TISSUES 

.f 

Parenchyma (fundamental tissue) 

Medullary rays 

Xylem cells (wood parenchyma) 

Tracheae (ducts) 

Phloem cells 

Sieve tubes 

Companion cells 

f Epidermis (outer cell walls cutinized) 

\ Cork (suberized tissue) 

Bast fibers 

f 

I 

Wood fibers 

Sclerenchyma fibers 

Stone cells 

Collenchyma

CHAPTER VII 

PLANT ORGANS AND ORGANISMS 

An organ is a part of an organism made up of several tissues and 

capable of performing some special work. 

An organism is a living entity composed of different organs or 

parts with functions which are separate, but mutually dependent, 

and essential to the life of the individual. 

The organs of flowering plants are either Vegetative or Reproductive. 

The vegetative organs of higher plants, are roots, stems, and 

leaves. They are concerned in the absorption and elaboration of 

food materials either for tissue-building or storage. 

The reproductive organs of higher plants include those structures 

whose function it is to continue the species, viz.: the flower, fruit and 

seed. 

The ripened seed is the product of reproductive processes, and the 

starting point in the life of all Spermatophytes. The living part of 

the seed is the embryo, which, when developed, consists of four parts, 

the caulicle, or rudimentary stem, the lower end of which is the be- 

ginning of the root, or radicle. At the upper extremity of the stem 

are one, two, or several thickened bodies, closely resembling leaves, 

known as cotyledons, and between these a small bud or plumule. 

The function of the cotyledon is to build up nourishment for the 

rudimentary plantlet until it develops true leaves of its own. 

THE ROOT 

The root is that part of the plant that grows into or toward the 

.soil, that never develops leaves, rather rarley produces buds, and 

whose growing apex is covered by a cap. 

The functions of a root are absorption, storage and support. Its 

principal function is the absorption of nutriment and to this end it 

generally has branches of rootlets covered with root-hairs which 

largely increase the absorbing surface. 

121 

These root-hairs are of


minute and simple structure, being merely elongations of the 

epidermis of the root back of the root cap into slender tubes with 

thin walls. 

PIG. 57. Cross-section of rootlet in the region of the root-hairs. (From Stevens.) 

The tip of each rootlet is protected by a sheath- or scale-like cover- 

ing known as the root cap, which not only protects the delicate grow- 

ing point, but serves as a mechanical aid in pushing its way through 

FIG. 58. Root-hairs, with soil-particles adhering. (Gager, after Sachs.) 

the soil. The generative tissues in the region of the root cap are: 

plerome, producing fibro-vascular tissue; periblem, producing cortex; 

dermatogen, producing epidermis; and calyptrogen, producing the 

root cap.

PLANT ORGANS AND ORGANISMS 123 

DIFFERENCES BETWEEN ROOT AND STEM 

The Root The Stem 

1. Descending axis of plant. i. Ascending axis of plant. 

2. Growing point sub-apical. 2. Growing point apical. 

3. Contains no chlorophyll. 3. Chlorophyll sometimes present. 

4. Branches arranged irregularly. 4. Branches with mathematical regu- 

larity. 

5. Does not bear leaves or leaf rudi- 5. Bears leaves and modifications, 

ments. 

6. Structure comparatively simple. 6. Structure better defined. 

Classification of Roots as to Form. i. Primary or first root, a 

direct downward growth from the seed, which, if greatly in excess of 

the lateral roots, is called the main or tap root. : Examples Taraxa- 

cum, Radish. 

2. Secondary roots are produced by the later growths of the stem, 

such as are covered with soil and supplied with moisture. Both 

primary and secondary roots may be either fibrous or fleshy. 

The grasses are good examples of plants having fibrous roots. 

Fleshy roots may be multiple, as those of the Dahlia, or may assume 

simple forms, as follows: 

Fusiform, or spindle-shaped, like that of the radish or parsnip. 

Napiform, or turnip-shaped, somewhat globular and becoming 

abruptly slender then terminating in a conical tap root, as the roots 

of the turnip. 

Conical, having the largest diameter at the base then tapering, as 

in the Maple. 

3. Anomalous roots are of irregular or unusual habits, subserving 

other purposes than the normal. 

4. Adventitious roots are such as occur in abnormal places on the 

plant. Examples: Roots developing on Bryophyllum and Begonia 

leaves when placed in moist sand. 

5. Epiphytic roots, the roots of epiphytes, common to tropical 

forests, for example, never reach the soil at all, but cling to the bark 

of trees and absorb nutriment from the air. Example: 

Vanilla. 

Roots of 

6. The roots of parasitic plants are known as Haustoria. These 

penetrate the bark of plants upon which they find lodgement, known

124 


asjiosts, and absorb nutritious juices from them. The Mistletoe, 

Dodder and Geradia are typical parasites. 

Duration of Root. Plants are classified according to the duration 

of the root, as follows: 

1. Annual plants are herbs with roots containing no nourishment 

for future use. They complete their growth, producing flower, 

fruit and seed in a single season, then die. 

2. Biennial plants develop but one set of aerial organs the first 

year, e.g., the leaves, and, as in the beet and turnip, etc., a large 

amount of reserve food material is stored in the root for the support 

of the plant the following season when it flowers, 

fruits and dies. . 

3. Perennial plants live indefinitely, as trees. 

Root Histology. Monocotyledons. The histology of mono- 

cotyledonous roots varies, depending upon relations to their surroundings, 

which may be aquatic, semi-aquatic, mesophytic, or 

xerophytic. In this connection we will discuss only the type of 

greatest pharmacognic importance, i.e., -the mesophytic type as 

seen in its most typical form in the transverse section of Honduras 

Sarsaparilla root. 

Examining such a section from -periphery toward the center, one 

notes the following: 

i. Epidermis of a single layer of cells many of which give rise to 

root-hairs. 

2. Hypodermis of two or three layers 

extremely thickened. 

of cells whose walls are 

3. Cortex, consisting of a broad zone of parenchyma cells many 

of which contain starch grains. 

4. Endodermis of one layer of endodermal cells whose walls are 

extremely thickened through the infiltration of suberin and lignin. 

5. Pericambium of one or two layers of meristematic cells whose 

walls are extremely thin. 

6. A radial fibro-vascular bundle of many alternating xylem and 

phloem patches and hence poly arch. The phloem 

tissue consists 

of phloem cells and sieve tubes. The xylem is composed of xylem 

cells, tracheae and wood fibers. 

7. Medulla or pith composed of parenchyma cells containing starch 

and often showing xylem patches cut off and enclosed within it.


Dicotyledons. The typical dicotyl root is a tetrarch one, four 

xylem alternating with four phloem patches. These roots have an 

unlimited power of growth. 

p m t en 

PIG. 59. Part of a transverse section of Honduras sarsaparilla root showing 

epidermis (e), root hair (ha), hypodermis (h), cortex (c), rendodemis (en), pericambium 

(p), trachea of one of the numerous xylem patches (/), and pith (m). 

The phloem patches are the small oval cellular areas wedged in between the outer 

portions of adjacent xylem masses. (Photomicrograph.) 

A. Of Primary Growth. 

A transverse section of a dicotyl root in its young growth shows 

the following structure from periphery toward center: 

i. Epidermis with cutinized outer walls, the cells often elongating 

to form root-hairs. 

ta


2. Hypodermis. 

3. Primary cortex with usually small intercellular spaces. 

4. Endodermis, or innermost layer of cells of the cortex* with 

lenticularly thickened radial walls. 

f , 5. Pericambium of one to two layers 

of actively growing cells which may 

produce side rootlets. 

6. Radial fibro-vascular bundle of 

four, rarely two or three or five or six 

phloem patches alternating with as 

many xylem arms. Not uncommon to 

find bast or phloem fiber along outer 

face of each phltfem patch. Xylem 

has spiral tracheae, internal to these a 

few pitted vessels, then, as root ages, 

more pitted vessels, also xylem cells 

and wood fibers make their appearance. 

7. Pith, a small zone of parenchyma 

cells. 

B. Of Secondary Growth (Most official 

roots). 

At about six weeks one notes cells 

dividing by tangential walls in the inner 

curve of phloem patches. This is in- 

trafascicular cambium. A single layer 

of flattened cells starts to cut off on 

elusive; g, primary xylem bun- 

; 

d Vc 

its inner side a quantity of secondary 

xylem and pushes out the patches of 

bast fibers, adds a little secondary 

r e 

-L S 

portion of root: lettered as in A; 

* C0rk " 

phloem on the outer side. Secondary 

xylem finally fills up the patches between 

the arms. The patches of bast fibers get flattened out. The 

pericambium has a tendency to start division into an inner and 

outer layer. The outer layer becomes a cork cambium (phellogen) 

surrounding the bundle inside of the endodermis. It cuts off cork 

tissue on its outer face, hence all liquid material is prevented from 

filtering through and cortex including endodermis, as well as the


epidermis, shrivel and dry up and separate off at the age of two 

to three months. The cork cambium (phellogen) may lay down 

secondary cortex internal to itself and external to the phlcem. 

Patches of cells of the inner layer of pericambium divide rapidly 

and are called interfascicular cambium. These join the intrafasci- 

cular cambium to form a continuous cambium ring which then 

cuts off additional secondary xylem on its inner face and secondary 

phloem on its outer face pushing inward the first-formed or protoxylem 

and outward the first-formed or protophloem. The medullary 

rays become deepened. 

Thus, in a transverse section made through a portion of a Dicotyl 

root showing secondary growth, the following regions are noted pass- 

ing from periphery to center: 

43 

1. Cork 

2. Cork cambium (phellogen) 

3. Secondary cortex 

4. Protophloem 

5. Secondary phloem 

6. Cambium 

7. Secondary xylem 

8. Protoxylem 

Strands of cells extending radially from the cortex to the center 

of the section separating each open fibro-vascular bundle from its 

neighbors. These are called medullary rays. 

Histology and Development of a Dicotyl Root (California Privet) 

A. Make a permanent mount of a T. S. of the root of the California 

Privet (Ligustrum Californicum) cut just above the root cap, and 

note the following structures, passing from periphery toward the 

center (see Fig. 61): 

1. Epidermis, composed of a layer of epidermal cells whose 

outer walls have been infiltrated with a substance called Cutin. 

2. Hypodermis, a layer of somewhat thick walled cells just be- 

neath the epidermis. 

3. Cortex, composed of cortical parenchyme 

angular intercellular air spaces. 

cells with small


4. Endodermis, or innermost layer of cells of the cortex, whose 

radial walls are lenticularly thickened. 

5. Pericambium, of a layer of actively growing meristematic 

cells, which has the power of producing lateral rootlets. 

6. Radial nbro-vascular bundle of five xylem arms alternating 

with as many phloem patches. Note the narrow spiral tracheae in 

the xylem patches. 

PIG. 61. Photomicrograph of a transverse section of a California Privet root 

of primary growth showing epidermis (e) ; hypodermis (k) cortex ; (c) endodermis 

; 

(en); pericambium (p); a xylem arm of the radial bundle (/) and pith (m). 

The section you have just studied illustrated in general the appear- 

ance of any Dicotyl root of primary growth. 

B. Mount permanently another T. S. cut through the same root a 

short distance above the first. 

Note that this is somewhat larger in diameter. Observe the root 

hairs starting from the epidermis; a broad cortex; a large clear and 

open looking endodermis; then pericambium; next, a central patch 

of xylem showing a faint pentarch relation. Pushed out are five


phloem tracts. Each of these constitutes a mass of protophloem 

(first formed phloem). On the inner face of each phloem mass may 

be seen intrafascicular cambium. At the outer end of each xylem 

tract there has been Cut off a patch of fine cambial cells (interfascicular 

cambium) which becomes joined to the intrafascicular cambium 

FIG. 62. Photomicrograph of a transverse section of a California Privet root 

made about 1)2 inches above the root tip and showing transition structure. The 

epidermis (e), primary cortex (pc) and endodermis are in the process of stuffing 

off, since cork (ck) has been laid down by the cork cambium (ph) directly beneath 

the endodermis. The cork cambium has also formed several layers of secondary 

cortex (sc) on its inner face. The protophloem represented largely by hard 

bast (hb) has been pushed out, while a small amount of secondary phloem represented 

by soft bast (sb) has been deposited beneath it by the cambium (c) which 

now is nearly circular in aspect. The protoxylem (px) has been pushed into the 

center by the encroaching secondary xylem (x) which has been laid down by 

the cambium on its inner face. Highly magnified. 

to develop secondary phloem on the outer face and secondary xylem 

on the inner face. 

C. Mount permanently a third T. S. out through the same root a 

short distance above the second. Note that this is still larger in 

diameter than the second. The pericambium has already divided

130 


into an inner and an outer layer. The outer layer has become the 

cork cambium, cutting off cork on its outer face beneath the endo- 

dermis. Cork being an impermeable barrier to water has prevented 

the nourishing sap from percolating through to the endodermis, cor- 

tex and epidermis. These regions have consequently begun to sluff 

off. Note that the cambium has begun to spread out into the form 

FIG. 63. Transverse section of California Privet root made about an inch and 

a half above the section shown in Fig. 6 1 and showing secondary structure. 

Note that epidermis, primary cortex and endodermis have completely disappeared. 

Cork (ck)', phellogen (ph); secondary cortex (sc); protophloem (p'); secondary 

phloem (p 2 

); cambium (c); secondary xylem (x z ) and protoxylem (*') 

(Photomicrograph) . 

of a ring. More secondary xylem has been formed on its inner face 

and additional secondary phloem has appeared on its outer face. 

(Fig. 62.) 

D. Make a permanent mount of a fourth Ti S. cut through the 

same root some distance above the third. Note that the epidermis, 

primary cortex and endodermis have completely peeled off. Cork 

is found as the external bounding layer and underneath it, cork 

ck 

ph 

sc


cambium. This cork cambium has developed secondary cortex on 

its inner face. The cambium has assumed a circular aspect. Just 

beneath the secondary cortex will be found flattened patches of 

protophloem, and beneath these secondary phloem 

masses have 

PIG. 64. Photomicrograph of a transverse section of an old 

California Privet root, showing completed secondary development. 

portion of 

Note the 

prominent medullary rays (mr); cork (ck)\ phellogen (ph~); secondary cortex 

(between ph and 2 

p')\ protophloem (p')\ secondary phloem ( ); cambium (c); 

secondary xylem (# 2 ); tracheae (/); wood fibers (wf); and protoxylem (#') 

been formed through the activity of the cambium. The cambium 

has developed new or secondary xylem on its inner face which has 

pushed the first formed or protoxylem toward the center of the root. 

(Fig. 63.)

132 


Abnormal Structure of Dicotyl Roots. In certain Dictoyl roots 

as Amaranthus, Jalap, Pareira, and Phytolacca, after the normal 

bundle system has been formed, there then develop successive cam- 

biums outside of these bundles, producing concentric series of open 

collateral bundles. 

Histology of a Dicotyl Tuberous Root (Aconitum). A transverse 

section made through the tuberous root of Aconitum Napellus near 

its middle shows a cork region of one or more layers of blackish or 

brownish cells; a broad cortex of two regions, viz.: an outer narrower 

and an inner broader zone. The narrower zone consists of from 

eight to fifteen layers of cortical parenchyma cells, interspersed among 

which are numerous irregular-shaped stone cells. Separating this 

zone from the broader one is an endodermis of a single layer of tan- 

genitally elongated endodermal cells. The broader zone consists of 

about twenty layers of parenchyme cells. Next, a five- to sevenangled 

cambium, within the angles of which and frequently scattered 

along the entire cambial line, occur collateral fibro-vascular bundles. 

In the center is found a broad five- to seven-rayed pith composed of 

parenchyma cells. The parenchyma cells of the cortical regions and 

pith contain single or two- to five-compound starch grains. 

ROOT TUBERCLES 

The roots of plants of the Leguminosa, Myricacece as well as some 

species of Aristolochiacece and of the genera Alnus and Ceanothus are 

characterized by the appearance upon them of nodule-like swellings 

called root tubercles. In the case of the Leguminosce the causative 

factor is a species of bacteria named Pseudomonas radicicola. This 

is a motile rod-shaped organism which appears widely distributed 

in soils. It is apparently attracted to the root-hairs of leguminous 

plants by a chemo tactic influence probably due to the secretions 

poured out by these structures. A number of these organisms penetrate 

the walls of the root-hairs by enzymic action. Upon entering 

the hairs they form bacterial tubes which branch and rebranch and 

extend into the middle cortex cells carrying the bacteria with them. 

Within the cortex cells the organisms multiply rapidly producing 

nest-like aggregations. Their presence here causes the formation of 

nodules or tubercles. Under oil-immersion magnification these


bacteria are found to exhibit variously shaped involution forms called 

bacterioids. They remain within the cells of the medio-cortex region 

gradually swelling up into zooglosa masses, until finally their bodies 

break down into soluble nitrogenous substances which are partly 

absorbed and assimilated and partly stored as reserve nitrogenous 

food for the green leguminous plant. 

FIG. 65. Root system of a legume showing tubercles. (Marshall.) 

In the modern rotation of crops, plant growers plough under the 

leguminous crops or their nodule-producing roots which decay and 

enrich the soil with ample nitrogenous material to supply the next 

season's crop of nitrogen-consuming plants. 

The writer has found tubercles on Myrica cerifera, Myrica Car- 

oliniensis and Myrica Macfarlanei seedling primary roots of 5 to 6 

months' growth, and from thence onward on the secondary roots 

inserted on the hypocotyl axis, on nearly all the adventitious roots of 

subterranean branches and on the subterranean branches of Myrica

134 


FIG. 66. Ps. radicicola. i, From Melilotus alba; 2 and 3, frorgi Medicago saliva; 

4, from Vicia villosa. (Marshall, after Harrison and Barlow from Lipman.) 

FIG. 67. Tubercular clusters on underground stem and roots of Myrica Mac- 

farlanei observed by the author at North Wildwood, N. J., Jan. 31, 1915.


cerifera, M. Caroliniensis, M. Gale, M. Macfarlanei, and Comptonia 

asplenifolia. The inciting organism has been isolated by him in 

pure culture according to Koch's postulates and named Actinomyces 

Myricarum Youngken. 

The tubercles occur either singly, as is frequently the case on 

subterranean branches, in small groups the size of a pea, or in larger 

coralloid loose or compact clusters which frequently attain the size 

of a black walnut. Each tubercle is a short cylindrical blunt-ended 

root-like structure which branches di- or trichotomously after attaining 

a certain length. The branches frequently rebranch at their tips 

which grow out into long thread-like structures from 1-3 cm. in 

length that may also branch and become entwined about the roots of 

other plants. The color of the youngest tubercles is a pinkish-gray 

brown. As the tubercles become older their color changes to brown, 

dark-brown and even black. (For a detailed description of the 

Myrica and Comptonia tubercles and their inciting organism, con- 

sult, "The Comparative Morphology, Taxonomy and Distribution 

of the Myricaceae of the Eastern United States" by Youngken, in 

Contributions from the Botanical Laboratory of the University of 

Pennsylvania, vol. iv, no. 2, 1919.) 

> 

THE BUD 

Buds are short young shoots with or without rudimentary leaves 

(bud scales) compactly arranged upon them. 

The plumule represents the first bud on the initial stem or caulicle. 

Scaly buds are such as have their outer leaf rudiments transformed 

into scales; there are often coated with a waxy or resinous substance 

without and a downy lining within, to protect them from sudden 

changes in climate. Buds of this character are common among 

shrubs and trees of temperate regions. 

Naked buds are those which are devoid of protective scales. 

They are common to herbaceous plants. 

Classification of Buds According to Development. i. A leaf bud 

is a young shortened shoot bearing a number of small leaves. It is 

capable of elongating into a branch which bears leaves. 

2. A flower bud is a rudimentary shoot bearing one or more 

concealed and unexpanded young flowers.

136 


3. A mixed bud is a young shoot bearing concealed unexpanded 

leaves and flowers. 

Classification of Buds According to Position on the Stem. i. A 

terminal bud is one which is located on the end of a stem (shoot). 

It is capable of elongating into a shoot which bears leaves or both 

leaves and flowers. 

2. An axillary or lateral -bud is one which arises in the leaf axil. 

It is capable of giving rise to a side branch or to a flower. Occa- 

sionally axillary buds do not develop and are then called dormant 

buds. 

3. An adventitious bud is one which occurs on some position of the 

stem other than at its apex or in the axil of a leaf. Such buds may 

be seen developing along the veins of a Begonia leaf or along the 

margin of a Bryophyllum leaf after these have been planted in 

moist soil for several days. 

4. An accessory bud is an extra bud which forms in or near the 

leaf axil. 

Classification of Buds According to Their Arrangement on the Stem. 

1. When a single bud is found at each joint or node of a stem, the 

buds are said to be alternate. 

2. When two buds are found at a node they are opposite. 

3. When several buds occur at a node they are whorled. 

THE STEM 

The stem is that part of the plant axis which bears leaves or modi- 

fications of leaves and its branches are usually arranged with mathe- 

matical regularity. 

Stems usually grow toward the light and so are heliotropic. 

The functions of a stem are to bear leaves or branches, connect 

roots with leaves, and conduct sap. 

When the stem rises above ground and is apparent, the plant is 

said to be caulescent. 

When no stem is visible, but only flower or leaf stalks, the plant is 

said to be acaulescent. 

Stems vary in size from scarcely ^ 5 inch in length, as in certain 

mosses, to a remarkable height of 400 feet or more. The giant 

Sequoia of California attains the height of 420 feet. Some of the


Eucalyptus trees of Australia and Tasmania are reported to attain 

the height of 500 feet. 

Nodes and Internodes. The nodes are the joints of stems. They 

represent the parts of the stem from which leaves or branches arise. 

Internodes are the parts of stems between nodes. 

Direction of Stem Growth. Generally the growth 

erect. Very frequently it may be: 

of the stem is 

Ascending, or rising obliquely upward. Example: Saw Palmetto. 

Reclining, or at first erect but afterward bending over and trailing 

upon the ground. Example: Raspberry. 

Procumbent, lying wholly upon the ground. : Example Pipsissewa. 

Decumbent, when the stem trails and the apex curves upward. 

Examples: Vines of the Cucurbitacea. 

Repent, creeping upon the ground and rooting at the nodes, as the 

Strawberry. 

Stem Elongation. At the tip of the stem there is found a group of 

very actively dividing cells (meristem) which is the growing point of 

the stem. All the tissues of the stem are derived from the cells of 

the growing point whose activity gives rise in time to three generative 

regions which are from without, inward: 

1. Dermatogen, forming epidermis; 

2. Periblem, forming the cortex; and 

3. Plerome, forming the fibre-vascular elements and pith. 

Duration of Stems. 

Annual, 

Examples: Corn. 

the stem of an herb whose life terminates with the season. 

Biennial, where the stem dies at the end of the second year. 

Example: Burdock. 

Perennial, when the stem lives for many years. Example: Oak. 

Stem Modifications. (i) twining, by elongation and marked 

circumnutation of young internodes as in Convolvulus, Dodder, 

etc. (2) Tendriliform by thread-like modification and sensitivity 

to contact of a side branch as in Passion flower, Squash, etc. (3) 

Spiny, by checking and hardening of a branch that may then become 

defensive ecologically as in hawthorn, honey locust, etc. (4) 

Aerial tuberous, in which one or more internodes, enlarge above 

ground and store reserve food as in pseudobulbs of orchids, Vitis

138 


gongylodes, etc. (5) Subterranean tuberous in which a subterranean 

stem or branch enlarges as a food-storing center: (a) annual 

type, tuber as in potato, etc., corm as in crocus, etc.; (b) perennial 

type, bulbs as in lily (scaly) and onion or hyacinth (tunicated). 

(6) Phylloid or leaf-like in which flattening branch expansion occurs, 

when leaves become reduced in size as in Asparagus, Ruscus } etc. 

(7) Cactoid, in which reduced condensed branches or stems become 

swollen for water (and food) storage as in Cacti, Euphorbia sp., etc. 

Above-ground Stems. A twining stem winds around a support, 

as the stem of a beam or Morning Glory. 

A culm is a jointed stem of the Grasses and Sedges. 

A climbing or scandent stem grows upward by attaching itself 

to some support by means of aerial rootlets, tendrils or petioles. 

Examples: Ivy, Grape, etc. 

The scape is a stem rising from the ground and bearing flowers 

but no leaves, as the dandelion, violet, or blood root. 

A tendril is a modification of some special organ, as of a leaf 

stipule or branch, capable of coiling spirally and used by a plant in 

climbing. Present in the Grape, Pea, etc. 

A spine or thorn is the indurated termination of a stem tapering 

to a point, as the thorns of the Honey Locust. 

Prickles are outgrowths of the epidermis and cortex and are 

seen in the roses. 

A stolon Is a prostrate branch, the end of which, on coming in contact 

with the soil, takes root, so giving rise to a new plant. Exam- 

ples: Currant and Raspberry. 

An herbaceous stem is one which is soft in texture and readily 

broken. Example: Convallaria majalis. 

An undershrub or sujfruitoose stem is a stem of small size and 

woody only at the base. Examples: Bitter-sweet, Thyme, etc. 

A shrubby or fruitcose stem is a woody stem larger than the pre-* 

ceding and freely branching near the ground. Example: Lilac, etc. 

A trunk is the 'woody main stem of a tree. , 

HERB AND TREE 

A tree is a perennial woody plant of considerable size, attaining a 

height of 15 or more feet, and having as the above-ground parts a 

trunk and a crown of leafy branches.


There are two plans of branching in trees. When the trunk, or 

main stem, extends vertically upward to the tip, as it does in the 

junipers, spruces and other conical trees, the type of branching is 

called excurrent] when it divides into several more or less equal 

divisions as in the elm and other spreading trees, it is said to be 

deliquescent. The deliquescent plan is the more common one among 

our deciduous trees. 

An herb is a plant whose stem does not become woody and perma- 

nent, but dies, at least down to the ground, after flowering. 

Underground Stems. A rhizome is a creeping underground stem, 

more or less scaly, sending off roots from its lower surface and stems 

from its upper. The rhizome grows horizontally, vertically or ob- 

liquely, bearing a terminal bud at its tip. Its upper surface is 

marked with the scars of the bases of aerial stems of previous years. 

Examples: Triticum, Rhubarb, etc. 

The tuber is a short and excessively thickened underground stem, 

borne usually at the end of a slender, creeping branch, and having 

numerous eyes or buds. Example: Tubers of the Potato. 

The corm is an underground stem excessively thickened and solid 

of buds from the center of the 

and characterized by the production 

upper surface and rootlets from the lower surface. Examples: 

Colchicum, Jack-in-the-Pulpit, etc. 

A bulb is a very short and scaly stem, producing roots from the 

lower face and leaves and flower from the upper. 

Tunicated bulbs are completely covered by broad scales which 

form concentric coatings. Examples: Onion, Squill, Daffodil. 

Scaly bulbs have narrow imbricated scales, the outer ones not en- 

closing the inner. Example: Lily. 

Tubers and corms are annual. Bulbs and Rhizomes are perennial. 

Exogenous and Endogenous Stems. Exogenous stems are typical 

of Gymnosperms and Dicotyledons and can increase materially in 

thickness due to presence of a cambium. Such stems show differen- 

tiation into an outer or cortical region and an inner or central cylinder 

region. 

Endogenous stems are typical of most Monocotyledons and cannot 

increase materially in thickness due to absence of cambium. The 

limited increase in diameter that does take place is due to the en-

140 


largement of the cells of the primary tissues. Such stems show no 

differentiation into cortical and central regions. 

Histology of Annual Dicotyl Stem. (In both annual and perennial 

dicotyledonous stems endodermis and pericambium are rarely seen 

since each has become so similar to cortex through passage of food, 

etc.) 

FIG. 68. Photomicrograph of cross-section of stem of Aristolochia sipho, 

where cambial activity is just beginning, a, Epidermis; b, collenchyma; c, thinwalled 

parenchyma of the cortex, the innermost cell layer of which is the starch 

sheath or endodermis; d, sclerenchyma ring of the pericycle; e, thin-walled parenchyma 

of the pericycle; /, primary medullary ray; g, phloem; h, xylem; i, interfascicular 

cambium; j, medulla or pith. X 20. (From Stevens.) 

1. Epidermis, cutinized, with hairs. 

2. Cortex composed of three zones: an outer or exocortex, whose 

cells are thin walled and contain chloroplasts; a middle ormedio- 

cortex, consisting of cells of indurated walls giving extreme pliability 

and strength, an inner or endocortex, a very broad zone of thin- and 

thick-walled parenchyma cells. 

3. The innermost layer of cells of the cortex called endodermis. 

(Not generally distinguishable.) 

4. Pericambium. (Not generally distinguishable.) 

J

fr 


I 

1 i 

UjJUi 111 -i 

: 

- 

FIG. 69. A diagram to show the character of the tissues and their disposition 

in a young stem of the typical dicotyledon type. (From Stevens.)

142 


5. Fibro-vascular bundles of open collateral type arranged in a 

circle with primary medullary rays between the bundles. 

6. Pith. 

FIG. 70. Diagram similar to the preceding but representing a later^stage and 

showing the tissues formed by the cambium. (From Stevens.)


Growth of Perennial DicotylStem and its Histology. A perennial 

dicotyl stem in the first year does not differ in structure from an 

annual. By the close of the year a cork cambium (phellogen) has 

originated beside the epidermis. In origin of cork cambium one 

of two methods: (a) either the epidermis may divide into an outer 

layer of cells that remains epidermis and an inner layer of cells that 

becomes cork cambium, or, (b) the outermost layer of cortex cells 

underneath the epidermis becomes active after being passive for 

one year, and lays down walls, the inner layer becoming cork cam- 

bium, the outer becoming a layer of cork. The cork cuts off water 

and food supplies from epidermis outside and so epidermis separates 

and falls off as a stringy layer. The cork cambium produces cork 

on its outer face and secondary cortex on its inner. 

Between the bundles certain cells of the primary medullary rays 

become very active and form interfascicular cambium which joins 

the cambium of the first-formed bundles (intrafascicular cambium) 

to form a complete cambium ring. By the rapid multiplication of 

these cambial cells new (secondary) xylem is cut off internally and 

new (secondary) phloem externally, pushing inward the first-formed, 

or protoxylem, and outward the first-formed, or protophloem, thus 

increasing the diameter of the stem. The primary medullary rays 

are deepened. Cambium may also give rise to secondary medullary 

rays. 

Sometimes, as in Grape Vines, Honeysuckles, and Asclepias, instead 

of cork cambium arising from outer cortex cells it may arise 

at any point in cortex. It is the origin of cork cambium at varying 

off. That 

depths that causes extensive sheets of tissue to separate 

is what gives the stringy appearance to the stems of climbers. 

At close of first year in Perennial Dicotyl Stem we note: 

1. Epidermis development of dermatogen or periblem in process of 

peeling off, later on entirely absent. 

2. Cork tissue or periderm. 

3. Cork cambium or phellogen. 

4. Sometimes zone of thin-walled cells containing chloroplasts cut off by 

cork cambium on inner face and known as phelloderm. 

5. Cortex in perennial stem cells of cortex may undergo modification into 

mucilage cells, into tannin receptacles, crystal cells, spiral cells, etc.

144 


6. Fibro-vascular bundles of open collateral type which are now arranged 

into a compact circle, and between which are found primary and often 

FIG. 71. 

secondary medullary rays. 

From without inward the following tissues make f. up v. bundles. 

Protophloem J Hard Bast long tenacious bast fibers. 

Secondary Phloem \ Soft Bast phloem cells and sieve tubes. 

Cambium active layer giving rise to secondary phloem on outer and 

secondary xylem or inner face, and adding to depth of med. rays. 

Secondary xylem wood fibers, pitted vessels, tracheids. 

Protoxylem spiral tracheae. 

7. Pith. 

Portion of cross-section of four-year-old stem of Aristolochia sipho, as 

shown by the rings of growth in the wood. The letters are the same as in Fig. 68 

but new tissues have been added by the activity of the cambium; and a cork cambium 

has arisen from the outermost collenchyma cells and given rise to cork. Tho 

new tissues are; I, cork cambium; k, cork; g, secondary phloem from the cambium, 

and just outside this is older crushed phloem; n, secondary xylem produced by 

the cambium; m, secondary medullary ray made by the cambium (notice that this 

does not extend to the pith). Half of the pith is shown. Notice how it has been 

crushed almost out of existence. Compare Figs. 68 and 71, tissue for tissue, to 

find out what changes the primary tissues undergo with age, and to what extent 

new tissues are added. Photomicrograph X 20. (From Stevens.) 

n


EXCEPTIONAL TYPES OF DICOTYL STEMS 

In a number of Dicotyledons and Gymnosperms, the secondary 

growth in thickness of the stem and frequently of the root differs from 

that which is found in the vast majority of species and so is called 

exceptional or anomalous. 

InPhytolacca, etc., there first arises a ring of primary bundles with 

broad loose medullary rays. Then the stem cambium ceases its 

PIG. 72. White birch (Betula populifolia). Portion of a branch showing the 

prominent lenticels. (Gager.) 

activity, and, outside the bast of the bundles already formed in the 

a new cambium starts 

pericambium or tissue developing from it, 

to lay down another ring of bundles in rather irregular fashion. 

Then after developing a wavy ring of bundles and connecting tissue 

that cambium closes up. Still another cambium ring arises without 

this, arid in a single season quite a number of these are found successively 

arranged in concentric fashion. 

In Gelsemium, species of Solanacece, Combretacece, Cucurbitacece, 

etc., there arises a cambium on the inner face of the xylem which

146 


forms internal phloem (or intraxylary phloem), thus giving rise to 

bicollateral bundles. 

In Strychnos Nux Vomica internal phloem exactly 

as in Gelse- 

mium, etc., appears but in addition interxylary phloem is developed. 

In the wood region of this plant axis the cambium starts at a certain 

age to lay down patches of phloem which become wedged in between 

xylem tissue as interxylary phloem. 

Lenticels and Their Formation. The epidermis in a great ma- 

jority of cases produces stomata, apertures, surrounded by a pair of 

guard cells, which function as passages for gases and watery vapor 

from and to the active cells of the cortex beneath. 

FIG. 73. Cross-section through a lenticel of Sambucus nigra. E, Epidermis; 

PH, phellogen; L, loosely disposed cells of the lenticel; PL, cambium of the 

lenticel; PS, phelloderm; C, cortical parenchyma containing chlorophyll. (From 

Sayre after Strasburger.) 

There very early originate in the region beneath the stomata 

loosely arranged cells from cork cambium which swell up during 

rain and rupture, forming convex fissures in the cork layer, called 

lenticels. 

The function of lenticels is similar to that of stomata, namely, 

to permit of aeration of delicate cells of the cortex beneath. 

Annual Thickening. In all woody exogenous 

stems such as 

trees and shrubs the persistent cambium gives rise to secondary 

xylem thickening every spring, summer and autumn. Soon a great 

cylinder of xylem arises which constitutes the wood of the trunk and 

branches. In the spring, growth is more active, and large ducts 

with little woody fiber are produced while in summer and autumn


growth is lessened and small ducts and much mechanical woody 

fiber are formed. Thus the open, loosely arranged product of the 

spring growth abuts on the densely arranged product of the last 

summer and autumn growth and the sharp contrast marks the 

periods of growth. To the spring, summer and autumn regions of 

growth of each year is given the term of "annual ring" By count- 

ing the number of these rings it is possible to estimate the age of 

the tree or branch. 

FIG. 74. Part of a transverse section of a twig of the linden, four years old. 

m, Pith; ms, medullary sheath; x, secondary wood; Ph, phloem; 2, 3, 4, annual 

rings; c, cambium; pa, dilated outer ends of medullary rays; b, bast; pr, primary 

cortex; k, cork. (From Sayre after Vines.) 

Bark. Bark or bork is a term applied to all that portion of a 

woody exogenous plant axis outside of the cambium line. 

In pharmacognic work, bark is divided into three zones, these 

from without inward being: 

1. Outer Bark or Cork. 

2. Middle Bark or Cortical Parenchyma. 

3. Inner Bark or Phloem. 

Periderm. Periderm is a name applied to all the tissue produced 

externally by the cork cambium (Phellogen). This term appears 

01 ten in pharmacognic and materia medica texts. 

Phelloderm. Phelloderm or secondary cortex is all that tissue 

produced by the cork cambium on its inner face. Its cells frequently 

contain chloroplasts.

148 


FIG. 75. Part of a cross-section through branch of Cytisus laburnum. (The 

branch was cut from the tree at the end of October.) From A to the last 

annual ring of wood; from A to B the spring growth with large tracheal tubes 

(T, T, D; between B and C and D and D are wood-fibers; between C and D and 

D and E, wood parenchyma; from E to F, cambium; F to G, phloem portion; G to 

H, cortical parenchyma; M, medullary ray. 

-A 

Below A the last wood-fibers and 

wood parenchyma formed the previous year. (From Sayre after Haberlandt.)


Histology of a Typical Bark, Cascara Sagrada. In transverse 

section passing from outer to inner surface, the following structural 

characteristics are evident: 

1. Cork, or outer bark, composed of several layers of rectangular 

cork cells. The most external layers are dead and appear black 

because they are filled with air. The inner layers of this region 

are living and contain brownish contents. 

2. Cork cambium (phellogen), a layer of delicate cells with pro- 

toplasmic corrtents in the process of division. 

3. Cortex, or middle bark, consisting of two regions, viz.: an outer 

zone of two or three rows of brownish collenchyma cells, and an 

inner broader zone of tangentially elongated cortical parenchyma 

cells. Imbedded within this zone will be noted numerous groups 

oi stone cells. 

4. Phloem, or inner bark, a very bjroad zone composed of irregular- 

shaped, elongated phloem masses separated from each other by 

medullary rays which converge in the outer phloem region. Each 

phloem mass consists of numerous sieve tubes and phloem cells, some 

of which latter contain spheriodal starch grains while others contain 

monoclinic prisms or rosette aggregates of calcium oxalate. Em- 

bedded within the phloem masses in tier-like fashion will be noted 

groups of bast fibers, each group of which is surrounded by a row 

of crystal fibers, individual cells of which can only be made out in 

this kind of a section. Each of these contains a monoclinic prism of 

calcium oxalate. The medullary rays possess brownish contents 

which take a red color with an alkaline solution. 

In radial longitudinal section a lengthwise view of the tissues will 

be seen. The medullary rays appear 15 to 25 cells in height and 

crossing at right angles to the other elements. The crystal fibers 

here will be seen to be composed of vertical rows of superimposed 

thin-walled cells each of which contains a monoclinic prism of 

calcium oxalate. The bast fibers appear elongated and taper ended 

and are associated with crystal fibers. 

In a tangential longitudinal section which has been cut through 

the phloem, the exact range in width of the medullary rays may be 

ascertained. In this bark the medullary rays are spindle-shaped 

in tangential view and one to four cells in width.


Wood. From a pharmacognic standpoint as well as that of the 

lumber trade, wood is all that portion of woody exogenous plant 

axis inside of the cambium line. In Dicotyl and Gymnosperm 

stems it therefore includes the xylem regions of the bundles, the 

FIG. 76. Photomicrograph of transverse section of Cascara Sagrada bark; 

k, cork; g, cork cambium; c, cortex; st, group of stone cells; bf, group of bast fibers; 

mr, medullary ray. 

xylem portions of the medullary rays and the pith, while in the roots 

of secondary growth of these plants it comprises the xylem portions 

of the bundles and the xylem medullary rays. 

As the cambium year after year adds new layers of wood to that 

already present on its inner face, the coveying of sap and storing


of starch, etc. is gradually relegated to the outer wood layers, since 

the inner layers, step by step, lose their protoplasmic contents and 

PIG. 77. Diagrammatic representation of a block of pine wood highly magnified, 

a, Early growth; b, late growth; c, intercellular space; d, bordered pit in 

tangential wall of late growth; m, /-and e, bordered pit in radial wall of early 

growth from different points ot view; h, row ot medullary cells for carrying food; 

g, row of medullary ray cells for carrying water; k, thin place in radial wall of ray 

cells that carry food (From Stevens.) 

power of conducting sap and become filled with extractive, resinous 

and coloring matters. The outer whitish layers of wood which con-

152 


tain living cells, functioning in the vegetative processes of the plant, 

constitute ,the alburnum or sap-wood. The drug Quassia is a good 

example of this kind of wood. The inner dead colored layers constitute 

the duramen or heart-wood. Important examples of this 

kind of wood used in pharmacy are Lignum Guaiaci, Haematoxylum, 

and Santalum Album. 

Microscopic Characteristics of Angiospermous and Gymnospermous 

Woods. The wood of Angiosperms is characterized by the pres- 

ence of tracheae (vessels) with various markings on their walls, 

FIG. 78. Photomicrograph 

of cross-section of very young 

cornstalk, where certain plerome 

strands have just gone 

over into vascular bundles. 

For comparison with Fig. 79. 

(Stevens.) 

particularly by small pits in the walls 

ot some of the tracheae, together with 

wood fibers, wood parenchyma and 

medullary rays. 

The wood of Gymnosperms is made 

up for the larger part ot tracheids with 

bordered pits which latter are charac- 

terized in radial longitudinal section by 

the presence of two rings, one within 

the other. A single row of these is 

seen on the tracheid wall. Medullary 

rays, frequently diagnostic for different 

species and woody parenchyme cells, 

are also found. 

Histology of Typical Herbaceous 

Monocotyl Stems (Endogenous). Passing from exterior toward 

center the following structures are seen: 

1. Epidermis whose cells are cutinized in their outer walls. 

2. Hypodermis, generally collenchymatic. 

3. Cortex. 

4. Endodermis or innermost layer of cortex. 

5. A large central zone of parenchyma matrix in which are found 

scattered nbro-vascular bundles of the closed collateral or rarely 

concentric type (amphivasal) . In this latter type, which is typical 

of old monocotyl stems, the xylem grows completely around phloem 

so that phloem is found in the center and xylem without and sur- 

rounding it.


Histology of a Typical Woody Monocotyl Stem. The stem of the 

Greenbrier, a woody monocotyl, will here be considered. In transverse 

section passing from periphery toward the center the following 

structural details will be noted: 

i. Epidermis, of a single layer of epidermal cells whose outer walls 

are strongly cutinized. Cutin is a wax-like substance which forms 

a protective coat to the epidermis, preventing the evaporation of 

water, the ingress of destructive parasites, and injury from insects. 

FIG. 79. Cross-section of cornstalk stern; a, epidermis; b, cortex and 

c, ground tissue. {After Stevens.) 

2. A cortex, composed of about ten or twelve layers of thick-walled 

parenchyma cells, the outer two or three layers of which are termed 

hypodermis. 

3. An endodermis, wavy in character and composed of endodermal 

cells whose brownish walls are strongly suberized. 

4. A sclerenchymatous cylinder sheath composed of somewhat 

separated masses of sclerenchymatous fibers and undeveloped fibrovascular 

bundles of the closed collateral type. 

5. A central matrix of strongly thickened parenchyma cells in 

which are scattered, irregularly, numerous closed collateral bundles. 

cells. Examine 

Small starch grains will be found in the parenchyma 

a representative bundle, and note the two very large tracheae and

154 


several smaller ones in the xylem portion of the bundle which faces 

toward the center of the section, In the outer or phloem portion 

of the bundle will be seen an area of soft, small-celled sieve tubes and 

phloem parenchyme. The entire bundle is enclosed by_a several 

FIG. 80. Photomicrograph of a representative portion of Greenbrier stem 

showing epidermis (e.p.), cortex (c), endodermis (e.n.d.), cylinder sheath (c.s.) 

sclerenchyma fibers of closed collateral bundle (&), fundamental parenchyma 

(/..), trachea (t). X 22. 

layered ring of sclerenchyma fibers, which on the inner face are 

called wood fibers, on the outer, bast fibers. The wood fibers con- 

stitute the supporting elements of the xylem, while the bast fibers 

are the supporting elements of the phloem.


THE LEAF 

The leaf is a usually flattened, rarely semi-centric, or centric-lateral 

expanse developed by the stem or by branches and in whose axil one 

or more branches arise. 

Leaves seldom develop buds over their surface or along their 

margin and in connection therewith roots. The capacity for bud 

development is restricted to three families, viz.: Crassulacece, Begoniacea 

and Gesneracea. 

Leaf Functions . The most essential function of plants is the conversion 

of inorganic into organic matter; this takes place ordinarily 

in the green parts, containing chlorophyll, and in these when exposed 

to sunlight. Foliage is an adaptation for increasing the extent of 

green surface. 

The functions of a leaf are photosynthesis, assimilation, respiration 

and transpiration. 

Photosynthesis is the process possessed by all green leaves or other 

green parts of plants of building up sugar, starch or other complex 

organic substances by means of chlorophyll and sunlight. This 

process takes place in nature, only during sunlight. CO 2 is taken 

in and O given off. 

Assimilation is the process of converting food material into proto- 

plasm. 

Respiration or breathing is the gaseous interchange whereby all 

living organisms take in oxygen and give off carbon dioxide. 

Transpiration is the giving off of watery vapor. 

Types of Leaves Developed in Angiosperms. These may be 

tabulated as follows: 

1 . Cotyledons (the primitive or seed leaves) . 

2 . Scale leaves. 

3. Foliage leaves. 

4. Bract leaves: (a) primary at base of inflorescence: (b) bracteo- 

lar leaves at a base of individual flowers. 

5. Sepals. 

6. Petals. 

7. Microsporophylls (stamens). 

8. Megasporophylls (carpels).

156 


Cotyledons. Cotyledons are the first leaves to appear upon the 

ascending axis and are single in Monocotyledons, double in Dicotyledons. 

Occasionally, as in certain Maples, there may be three 

cotyledons shown. This is due to a splitting of one of the cotyledons. 

There exist no true cases of polycotyledony (development of many 

cotyledons) among Angiosperms, as in Gymnosperms. In Mono- 

cotyledons the single cotyledon is a terminal structure and truly 

axial in relation to the hypocotyl and radicle. From a primitively 

Monocotyl-like ancestry Dicotyledons develop a second cotyledon 

on the Epicotyledonary node. Later, by a suppression of the .second 

node the second cotyledon is brought to the level of the first. 

Scale Leaves. Scale leaves are reduced foliage leaves. They are 

found on certain rhizomes, above ground stems, such as Dodder, 

etc., on bulbs, and forming the protective scales of scaly buds. 

Foliage Leaves. These are the common green leaves so familiar 

to all. 

Bract leaves are modified leaves appearing on inflorescence axes. 

Sepals, petals, microsporophylls and megasporophylls are floral 

leaves and will be treated at length under the subject of the flower. 

Origin and Development of Leaves. Leaves arise around the 

growing apex region of a stem or branch as lateral outgrowths, each 

consisting at first of a mass of cells called the primordial leaf. 

Through continued cell-division and differentiation of these cells in 

time the mature leaf is developed. The primordial leaf is formed 

by a portion of the dermatogen of the growing stem apex, which 

becomes epidermis, a portion of the periblem, producing mesophyll 

which grows into this, and a part of the plerome, which becomes 

vascular tissue within the mesophyll. 

In the sub-divisions of cells around the growing stem-apex, the 

primordial leaves (primordia) do not arise exactly at the same time. 

There is a tendency toward spiral arrangement. 

Phyllotaxy. Phyllotaxy is the study of leaf arrangement upon 

the stem or branch, and this may be either alternate, opposite, 

law in the 

^whorled, or verticillate, or fascicled. It is a general 

arrangement of leaves and of all other plant appendages that they 

are spirally disposed, or on a line which winds around the axis like 

the thread of a screw. The spiral line is formed by the union of

PLANT ORGANS AND ORGANISMS 1 57 

two motions, the circular and the longitudinal, and its most common 

modification is the circle. 

In the alternate arrangement there is but one leaf produced at 

each node. Examples: Aconite, Magnolias. 

Opposite, when a pair of leaves is developed at each node, on 

opposite sides of the stem. Examples: Mints, Lilac. 

Decussate, when the leaves are arranged in pairs successively along 

the stem, at right angles to each other. Example: Thoroughwort. 

Whorled or Verticillate, when three or more form a circle about 

the stem. Examples: Canada Lily and Culver's root. 

Fascicled or Tufted, when a cluster of leaves is borne from a single 

node, as in the Larch and Pine. 

The spiral arrangement is said to be two-ranked, when the third 

leaf is over the first, as in all Grasses; three-ranked, when the fourth 

is over the first. Example: Sedges. The five-ranked arrangement 

is the most common, and in this the sixth leaf is directly over the 

first, two turns being made around the stem to reach it. Example : 

Cherry, Apple, Peach, Oak and Willow, 

etc. As the distance be- 

tween any two leaves is two-fifths of the circumference of the stem, 

the five-ranked arrangement is expressed by the fraction %. In 

the eight-ranked arrangement the ninth leaf stands over the first, 

and three turns are required 

to reach it, hence the fraction 

% expresses it. Of the series 

of fractions thus obtained, the 

numerator represents the num- 

ber of turns to complete a 

. 

cycle, or to reach the leaf ^^ which is directly over the first; pio 8l _Three principal types of 

the denominator, the number vernation. (Robbins.) 

of perpendicular rows on the 

f coupllcate 

stem, or the number of leaves, counting along the spiral, from any 

one to the one directly above it. 

Vernation. Prefoliation or Vernation relates to the way in which 

leaves are disposed in the bud. A study of the individual leaf 

enables u to distinguish the following forms. When the apex is bent 

inward toward the base, as in the leaf of the Tulip Tree,-it is said to be

158 


inflexed or reclinate vernation; if doubled on the midrib so that 

the two halves are brought together as in the Oak or Peach, it is 

conduplicate; when rolled inward from one margin to the other, as 

in the Wild Cherry, it is convolute; when rolled from apex to base, 

as in Ferns, it is circinate; when folded or plaited, like a fan as in 

Ricinus, Maples, Aralias, etc., it is plicate; 

if rolled inward from each 

margin toward the midrib on the upper side, as the leaves of the 

Apple or Violet, involute; when rolled outward from each margin as 

FIG. 82. Stereogram of leaf structure. Part of a veinlet is shown on the right. 

Intercellular spaces are shaded. (From Stevens.) 

Dock or Willow leaves, revolute. The inner surface is always that 

which will form the upper surface when expanded. 

TJie Complete Leaf. The leaf when complete consists of three 

parts, lamina, petiole, and stipules. The lamina or blade is the expansion 

of the stem into a more or less delicate framework, made up 

of the branching vessels of the petiole. 

The petiole is the leaf stalk. The stipules are leaf-like appendages 

appearing at the base of the petiole.


The leaf of the Tulip Poplar or Liriodendron affords a good exam- 

ple of a Complete Leaf. 

Sometimes the lamina or blade is attached directly to the stem 

by its base and is then said to be sessile. If the petiole is present, 

petiolate. 

When leaf stipules are absent, the leaf is said to be exstipulate, 

when present, stipulate. 

The petiole is seldom cylindrical in form, but usually channelled 

on the %>per side, flattened, or compressed. The stipules are always 

in pairs and closely resemble the leaf in structure. 

The blade of the leaf consists of the framework, made 3p of branch- 

ing vessels of the petiole, which are woody tubes pervading the soft 

tissue called mesophyll, or leaf parenchyma, and serve not only as 

supports but as veins to conduct nutritive fluids. 

in simple leaves such as many of the Mosses. 

Veins are absent 

Leaf Venation. Furcate or Forked Venation is characteristic of 

many Ferns. 

Parallel Venation is typical of the Monocotyledons, as Palms, 

Lilies, Grasses, etc. 

Reticulate or Netted Veins characterize the Dicotyledons, as the 

Poplar or Oak. The primary veins in these are generally pinnate while 

the secondary ones and their branches are arranged in netted fashion. 

Pinni-veined or Feathered-veined leaves consist of a mid-vein with 

lateral veinlets extending from mid-vein to margin at frequent 

intervals and in a regular manner. Example: Calla. 

Palmately Veined leaves consist of a number of veins of nearly the 

same size, radiating from petiole to margin. Example: Maple leaf. 

Veins are said to be anastomosing when they subdivide and join 

each other, as the veins near the margin of Eucalyptus leaves. 

Leaf Insertion. The point of attachment of the leaf to the stem is 

called the insertion. A leaf is: 

Radical, when inserted upon an underground stem. 

Cauline, when upon an aerial stem. 

Ramal, when attached directly to a branch. 

When the base of a" sessile leaf is extended completely around the 

stem it is 

perfoliate, the stem appearing to pass through the blade. 

Example: Uvularia perfoliata or Mealy Bellwort.

i6o 


FIG. 83. Leaf outlines: Linear (i); lanceolate (2); oblong (3); elliptical (4): 

ovate-lanceolate (5); oblanceolate (6); spatulate (7); obovate-lanceolate (8): 

orbicular (9); reniform (10); cuneate (n).


When a sessile leaf surrounds the stem more or less at the base, 

it is called clasping or amplexicaul. Example: Poppy (Papaver 

somniferum) . 

When the bases of two opposite leaves are so united as to form 

one piece, they are called connate-perfoliate, as Eupatorium perfoliatum 

or Boneset. 

Leaves are called equitant when they are all radical and succes- 

sively folded on each other toward their bases, as in Iris sp. 

The Forms of Leaves. Simple leaves are those having a single 

blade, either sessile or petiolate. 

Compound 

leaves are divided into two or more distinct subdivi- 

sions called leaflets, which may be either sessile or petiolate. 

Simple leaves and the separate blades of compound leaves are described 

as to general outline, apex, base, marginal indentations, sur- 

face and texture. 

(a) General Outline (form viewed as a whole without regard to 

indentations of margin). Dependent upon kind of venation. 

When the lower veins are longer and larger than the others, the 

leaf is Ovate, or Egg-shaped. Parallel-veined leaves are usually 

linear, long and narrow of nearly equal breadth throughout 

(Linaria), or lanceolate, like the linear with the exception that the 

broadest 

Buchu. 

part is a little below the center. Example: Long 

Elliptical, somewhat longer than wide, with rounded ends and 

sides. Example: Leaf of Pear. 

Oblong, 

Matico. 

when longer than broad, margins parallel. Example: 

Inequilateral, margin longer on one side than the other, as the 

Hamamelis, Elm and Linden. 

Orbicular, circular in shape. Example: Nasturtium. 

Peltate, or shield-shaped, having the petiole inserted at the center 

of the lower surface of the lamina. Example: Podophyllum. 

Filiform, or thread-like, very long and narrow, as Asparagus leaves. 

Ovate, broadly elliptical. Example: Boldo. Obovate, reversely 

ovate. Examples: Short Buchu and Menyanthes. 

Oblanceolate, reversely lanceolate. Example: Chimaphila. 

Cuneate, shaped like a wedge with the point backward.


FIG. 84. Leaf bases (12-17); leaf apices (18-26); compound leaves (27-31). 

Cordate (12); auriculate (13); connate-perfoliate (14); sagittate (15); hastate 

(16) ; peltate (17). Acuminate (18) ; acute (19) ; obtuse (20) ; truncate (21) ; retuse 

(22) ; emarginate (23) ; cuspidate (24) ; mucronate (25) ; aristate (26). Imparipinnate 

(27); paripinnate (28); bi-pinnate (29); decompound (30); palmately 

5-foliate (31).

Spatuldte, like a spatula, 

apex. Example: Uva Ursi. 


with narrow base and broad rounded 

Ensiform, when shaped like a sword. Example: Calamus. 

Acerose or acicular, tipped with a needle-like point, as Juniper. 

Falcate, sythe or sickel shaped as Eucalyptus. 

Deltoid, when the shape of the Greek letter A, as Chenopodium. 

(b) Apex of Leaf. Acute, when the margins form an acute angle 

at the tip of the leaf. Examples: Eriodictyon, Digitalis. 

Acuminate, when the point is longer and more tapering than the 

acute. Examples: Pellitory, Coffee. 

Obtuse, blunt or round. Example: Long Buchu. 

Truncate, abruptly obtuse, as if cut square off. Example: Meli- 

lotus leaflets. 

Mucronate, terminating in a short, soft point. Example: Senna 

leaflets. 

Cuspidate, like the last, except that the point is long and rigid. 

Aristate, with the apex terminating in a bristle. 

Emarginate, notched. Example: Pilocarpus. 

Retuse, with a broad, shallow sinus at the apex. Example: Petal 

of Rosa gallica. 

Obcordate, inversely heart-shaped. Example: Oxalis. 

(c) Base or Leaf. Cordate, heart-shaped. Examples: Lime and 

Coltsfoot. 

Reniform, kidney-shaped. Examples: Ground Ivy, Asarum. 

Hastate, or halbert-shaped, when the lobes point outward from 

the petiole. Example: Aristolochia Serpentaria. 

Auriculate, having ear-like appendages at the base. Example: 

Philodendron. * 

Sagittate, arrow-shaped. Example: Bindweed. 

Cuneate, wedge shaped. Examples: 

(d) Margin of Leaf. Entire, when the margin 

Short Buchu and Uva Ursi. 

is an even line. 

Example: Belladonna. 

' 

Serrate, with sharp teeth which incline forward like the teeth of a 

hand-saw. Examples: Peppermint, Yerba Santa, Buchu. 

Dentate, or toothed, with outwardly projecting teeth. Chestnut. 

Crenate, or Scalloped, similar to the preceding forms, but with the 

teeth much rounded. Examples: Digitalis, Salvia.

164 


FIG. Ss.^Leaf margins: Pinnately-lobed (32); pinnately-cleft (33); pinnatelyparted 

(34); pinnately-divided (35); palmately tri-lobed (36); palmately tri-cleft 

(37); palmately 3-parted (38); palmately 3-divided (39); crenate (40); serrate 

(41); dentate (42); repand or undulate (43); sinuate-dentate (44).


Repand, or Undulate, margin a wavy line. Example: Hama- 

melis. 

Sinuate, when the margin is more distinctly sinuous than the last. 

(Stramonium.) 

Incised, cut by sharp, irregular incisions. Example: Hawthorn. 

Runtin ate, the peculiar form of pinna tely incised leaf observed in 

the Dandelion and some other Composite 

in which the teeth are 

recurved. 

A Lobed leaf is one in which the indentations extend toward the 

mid-rib, or the apex of the petiole, the segments or sinuses, or 

both, being rounded. Example: Sassafras. 

Cleft is the same as lobed, except that the sinuses are deeper, and 

commonly acute. Example: Dandelion. 

A Parted leaf is .one in which the incisions extend nearly to the 

mid-rib or the petiole. Example: Geranium maculatum. 

In the Divided leaf the incisions extend to the mid-rib, or the 

petiole, but the segments are not stalked. Example: Watercress. 

If the venation is pinnate, the preceding forms may be described 

as pinnately incised, lobed, parted, or divided. If the venation is 

radiate, then the terms radiately or palmately lobed, incised, etc., 

are employed. 

The transition from Simple to Compound Leaves is a very gradual 

one, so that in many instances it is difficult to determine whether a 

given form is to be regarded as simple or compound. The number 

and arrangement of the parts of a compound leaf correspond with 

the mode of venation, and the same descriptive terms are applied 

to outline, margin, etc., as in simple leaves. 

Leaves are either pinnately or palmately compounded. The term 

pinnate is frequently given to the former while that of palmate is 

often assigned to the latter. They are said to be abruptly pinnate 

or paripinnate when the leaf is terminated by a pair of leaflets; odd 

pinnate or imparipinnate when it terminates with a single leaflet. 

When the leaflets are alternately large and small, the leaf is inter- 

ruptedly pinnate, as the Potato leaf. When the terminal leaflet is 

the largest, and the remaining ones diminish in size toward the base 

the form is known as lyrate, illustrated in the leaf of the Turnip. 

leaves have the leaflets attached to the 

Palmately compound


apex of the petiole. When these are two in number the leaf is 

bifoliate, or binate; if three in number, trifoliate, or ternate, 

as in 

Menyanthes; when four in number, quadrifoliate, etc. If each of 

the leaflets of a palmately compound leaf divides into three, the leaf 

is called biternate; if this form again divides, a triternate leaf results. 

Beyond this point the leaf is known as decompound. In the case 

of pinnately-compound leaves, when division progresses so as to 

separate what would be a leaflet into two or more, the leaf becomes 

bipinnate, as the compound leaves of Acacia Senegal or on the new 

wood of Gleditschia; if these become again divided, as in many 

Acacia species, the leaf is termed tripinnate. Examples of decom- 

pound leaves seen in Cimicifuga and Parsley. 

Leaf Texture. Leaves are described as : 

Membranous, when thin and pliable, as Coca. 

Succulent, when thick and fleshy, as Aloes, and Live Forever. 

Coriaceous, when thick and leathery, as Eucalyptus, Uva Ursi 

and Magnolia. 

Leaf Color. Petaloid, when of some brilliant color different from 

the usual green, as the Coleus and Begonia, and other plants which 

are prized for the beauty of their foliage rather than their blossoms. 

Leaf Surface. Any plant 

surface is : 

Glabrous, when perfectly smooth and free from hairs or protuber- 

ances. Example: Tulip. 

Glaucous, when covered with bloom, as the Cabbage leaf. 

Pellucid-punctate, when dotted with oil glands, as the leaves of 

the Orange family. 

Scabrous leaves have a rough surface with minute, hard points. 

Pubescent, covered with short, soft hairs. Example: Strawberry. 

Villose, covered with long and shaggy hairs. Example: Forgetme-not. 

Sericious, silky. Example: Silverleaf. 

Hispid, when covered with short, stiff hairs. Example: Borage. 

Tomentose, densely pubescent and felt-like, as the Mullein leaf. 

Spinose, beset with spines, as in the Thistle. 

Rugose, when wrinkled. Example: Sage. 

Verrucose, covered with protuberances or warts, as the calyx of 

Chenopodium.


Duration of Leaves. Leaves vary as to their period of duration. 

They are: Persistent, or evergreen, if they remain green 

for a year or more. 

Deciduous, if unfolding in spring and falling in autumn. 

Caducous, or fugacious, if falling early in the season. 

on the tree 

Parts of Typical Leaf. The parts of a typical leaf are petiole or 

leaf stalk, lamina or blade, and stipules. 

Gross Structure and Histology of the Petiole. The petiole in 

Monocotyledons is usually a broadened, sheathing basal structure 

which connects the lamina to the stem. Into this a set of closed 

collateral vascular bundles of the stem extend, these showing xylem 

uppermost and phloem beneath; but in the Palmacece, Aracece, 

Dioscoreacece and Musacea the petiole in part or throughout may be 

much thickened, strengthened and developed as a semi-cylindric or 

cylindric structure frequently showing, as in Palmacea, generally, 

two sets of bundles. In all of these the petiole shows distinct scat- 

tered closed collateral bundles embedded in parenchyma and sur- 

rounded by epidermis. In the Monocotyl genus Maranta a special 

swelling is found at the apex of the petiole which is termed a pulvinus. 

In Dicotyledons the petiole attains its most perfect development 

and here usually shows differentiation into a pulvinus or leaf cushion 

and stalk portion. The pulvinus is sensitive to environal stimuli 

and in some groups as Oxalidacece and Leguminoscz a gradual increase 

in sensitivity up to a perfect response can be traced. Moreover, in 

these, if we start with the simpler less sensitive pulvini and pass by 

stages to the most complex, we note that a special substance known 

as the aggregation body develops in the pulvinar cortex cells and 

that this substance undergoes rapid molecular change on stimulation 

of the leaf. The stalk portion of the petiole in Dicotyledons is 

usually plano-convex or nearly to quite circular in outline; rarely in 

certain families does it simulate Monocotyledons in becoming 

abruptly or gradually thinned or flattened or widened out so as to 

sheath round the stem. The most striking example of this is seen in 

the Umbellifera where the flattened sheathing leaf stalk is known as 

the peridadium. Such a structure is not peculiar to the Umbelli- 

fera for in many Ranunculacece, etc., a similar sheathing development 

is observed. The stalk may bear the laminar tissue on its extremity.


This is most commonly the rule, but when the plant is exposed to 

xerophytic conditions, as the Acacias of Australia, the stalk, instead 

of being cylindric or sub-cylindric, becomes flattened from side to 

side, until there is produced a bifacial vertically placed petiole, with 

a large green surface that wholly takes the place of the lamina. 

The petiolar structure in primitive types of Dicotyls resembles 

that^seen in Monocotyls except 

that the bundles are more con- 

densed side by side. In these the petiole is somewhat dorsiventral, 

shows an external epidermis, a flattened cortex with a set of parallel 

vascular bundles, each with xylem uppermost and phloem below. 

From this we pass to another group in which the bundles form 

three-fourths of a circle and in which the upper bundles show incurv- 

ing orientation, to still another in which, as in Nepenthes, all of the 

bundles form nearly a cylinder. Finally in Ficus, Geranium, Podo- 

phyllum and other plants showing conpletely formed cylindric 

petioles, the bundles form a continuous ring enclosing pith and surrounded 

by cortex and epidermis, as in Dicotyl stems. 

Stipules. Stipules are lateral leafy or membranous outgrowths 

from the base of the petiole at its junction with the stem. They 

may be divided into two groups, viz.: lateral and axillary. The 

lateral group includes four types, namely, free lateral, lateral adnate, 

lateral connate and lateral interpetiolar. 

Free lateral stipules are seen in Leguminoscz, Rosacece, Beeches, 

etc. They are free on either side of the petiole and supplied by 

vascular tissue from the petiolar bundle mass. In appearance and 

duration they may be either green, foliaceous and persistent or mem- 

branous to leathery, scale-like and caducous. Caducous scaly stip- 

ules only function as bud scales through the winter and fall in spring 

as the buds expand. 

Lateral adnate stipules are such as fuse with and are carried up 

with the petiole as wing-like appendages. This type is seen in the 

genus Rosa, in Clovers, etc. 

Lateral connate stipules are such as join and run up with the 

petiole to form a structure which is called a ligule. This structure 

is common to the Graminese or Grass family. 

Lateral interpetiolar stipules are common to many species of the 

Rubiacea. In the genus Cinchona the leaves are opposite and orig-


inally had free lateral stipules which latter gradually fused with the 

stem, slid across it and adjacent stipules, then fused together to 

form a median structure on either side of the stem. 

The axillary group represent stipules which stand in the axil of 

the leaf with the stem. Such may be free axillary structures, arising 

as distinct processes, or connate, when the two stipules unite at their 

margins and sheath the stem, as in many species of the Polygonacecz 

such as Buckwheat, Rhubarb, Yellow Dock, Knot Weeds, etc. The 

sheath formed is called an ochrea. 

Modified Stipules. In some plants such as the Locust and several 

other trees and shrubs of the Legume family, the stipules become 

modified for defensive purposes as spines or prickles. 

In the Sarsa- 

parilla-yielding plants and other species of the genus Smilax they 

undergo modification into tendrils which are useful in climbing. 

The Lamina. This as was previously indicated represents an ex- 

pansion of the tissues of the petiole, but in sessile leaves is attached to the stem and so a direct stem outgrowth. 

directly 

Mode of Development of the Lamina of Leaves. The lamina of 

leaves develops in one of six ways. 

1. Normal' or Dorsoventral. 

2. Convergent. 

3. Centric. 

4. Bifacial. 

5. Reversed. 

6. Ob-dorsi-ventral. 

The first foui will be considered. 

A. Dorsoventral (the commonest). 

(a) Dorsoventral Umbrophytic. Flattened from ab6ve downward. 

Plants with such leaf blades tend to grow in the shade. 

(6) Dorosoventral Mesophytic. Similar to the former, but plants 

usually grow directly in the open and exposed to sunlight and winds. 

(c) Dorsovertral Xerophytic. Similar to former, but plants not 

only grow exposed, but exposed to hot desert conditions or to cold 

vigorous conditions. 

(d) Dorsoventral Hydrophytic. All transitions between typical 

mesophytic forms to those of marshy places, to swamps and borders 

of streams and finally with leaves wholly emersed, the last a com- 

pletely hydrophytic type.

i yo 


Gross Structure and Histology of Different Types of Dorsoventral 

Leaf Blades. i. Umbrophytic. Characterized by leave^ mostly 

undivided and having the largest and most continuous leaf expanse. 

Usually the deepest green leaves we have, to enable the leaves to 

FIG. 86. Transverse section through portion of dorsoventral leaf blade of 

horehound (Marrubium vulgare). Upper epidermis devoid of stomata (up.ep.); 

lower epidermis which possesses stomata (I. ep)\ palisade parenchyma (pal.); 

spongy parenchyma (sp. p); xylem (x) and phloem (ph) regions of fibrovascular 

tissue of stronger vein; long-pointed non-glandular trichome (t); branched trichomes 

(ft 1 , ft z , ft 3 ); several types of glandular trichomes (gt, gt 2 , gt s , gl 4 ). 

absorb scattered and reduced rays that pass in through high trees 

and shrub overhead. Their texture is usually thin and soft. In 

microscopic structure they are covered with a cutinized epidermis 

which has all the stomata on the lower surface. The mesophyll is 

fairly spongy, the spongy parenchyma having decided intercellular


spaces. The lower epidermis is more or less hairy. Examples: 

Dog's Tooth Violet, Asteis. 

2. Mesophytic. Leaves tend to subdivision, either to slight or 

moderate lobing, seldom to complete subdivision in pinnate or 

tripinnate fashion. Example: Dandelion. In microscopic struc- 

ture, they consist of an upper and lower epidermis, the upper epider- 

FIG. 87. Photomicrograph of cross-section through a portion of the leaf of a 

xerophyte, Ficus elastica, showing upper epidermis (u.e.), water storage tissue 

(iv.s.), cystolith suspended on stalk within a cystolith sac (cys), palisade parenchyma 

(p.p.), spongy parenchyma (s.p.), vein (v), lower epidermis (I.e.), and 

stoma (s). (Highly magnified.) 

mis being the thicker of the two. The stomata are wholly or are 

mainly on the lower epidermis. Hairs are seldom seen. The palisade 

mesophyll is toward the upper surface, the spongy mesophyll 

toward the lower. The intercellular-air-spaces in the spongy paren- 

chyma 

are small. 

and lower 

3. Xerophytic. Leaves characterized by a thick upper 

cuticle and by having their numerous, small stomata restricted to

172 


the lower surface or present more or less equally on both surfaces, 

where they are sunken in depressions. They may be either firm, 

leathery, tough, fibrous, or may become swollen .up in their meso- 

phyll chiefly in their spongy parenchyme cells and store considerable 

mucilage. Examples: Yucca, Ficus, Aloe, Agave. Succulent 

forms like Aloe generally possess a thin but tenacious cuticle. 

4. Hydrophytic. All gradations are seen. In pond plants, such 

as the Water Lily, the leaves have long split petioles which bring 

the blade up to the surface of the water. The stomata are entirely 

on the upper surface. In Ranunculus, the lower leaves are cut up 

into filiform segments. These are devoid of stomata. Their mesophyll 

is soft, open, and spongy. The epidermis is quite thin. The 

upper leaves are floating, trilobed, and have stomata only on their 

upper surface. In Utricularia, some of the filiform submerged leaves 

are modified into bladders which trap insect larvae and smaller 

Crustacese. 

B. Convergent. In Phormium tenax, the base of the blade is 

sheathing, it then converges and opens 

out above. In the various 

species of Iris the petiole is sheathing, the upper part being fused 

(mostly seen in monocotyls). 

C. Centric. Succulent. Nearly always 

associated with Xero- 

phytes. 

Xerophytic. Centric laminae are produced gradually by an encroachment 

of the under on the upper surface, and the swelling of the 

whole. In a completely centric leaf of the succulent kind, like that 

of Sedum, the difference between the upper and lower surface is lost. 

Stomata are found scattered over the entire epidermis. The bundles 

are arranged in a circle, the mid-rib being in the center. A great 

deal of mucilage is found stored in the central cells. In a typical 

Xerophytic Centric leaf, like that of the Pine or Sansemera . 

cylin- 

drica, the epidermis shows a tnick cuticle; the stomata are sunken in 

cavities of the epidermis; the epidermis and leaf tissue are strength- 

ened by scleroid bands in the centric mesophyll. 

D. Bifacial. Leaves with laminae which stand edge on in relation 

to the sun's rays. The best illustrations are seen among dicotyledons, 

such as Eucalyptus, Callistemnon, and other genera of Myr- 

tacece. Both surfaces are similar, having stomata about equal in


number. The mesophyll is differentiated into a central spongy paren- 

chyme containing bundles, and a zone of palisade cells on either 

side facing the epidermises. 

Structure and Development of Stomata. Stomata are slit-like 

openings in the epidermis of leaves or young green stems surrounded 

ep. v. p.p. 

ep. p.p. 

PIG. 88. Photomicrograph of a transverse section of a bifacial leaf of Eucalyptus 

globulus showing epidermis (ep.), palisade parenchyma (p.p.), toward both 

surfaces, spongy parenchyma (s.p.), vein (v), and oil reservoir (o.r.) lined with 

secretory epithelium. (Highly magnified.) 

by a pair of cells, called guard cells, whose sides opposite one another 

are concave. They form a communication between the intercellularair-space 

(respiratory cavity) beneath them and the exterior. The 

slit-like opening taken with the guard cells, constitutes what is known 

as the stomatal apparatus.

174 


The epidermal cells which abut on the stomatal apparatus are 

called neighboring cells or subsidiary cells. These in many cases, as 

in species of Helleborus, Sambuscus, Hyacinthus, Paonia, Ferns, etc., 

are very similar to the other epidermal cells, but in a large number of 

plants they differ in size, arrangement and shape 

from the other 

cells of the epidermis which do not abut upon the stomatal apparatus. 

In Senna they are two in number one larger than the other and 

arranged parallel to the guard cells of the stoma; in Coca a similar 

arrangement occurs but the cells are more even in size, nevertheless 

they lack the characteristic papillae found on the other epidermal 

cells; in Pilocarpm they are usually four in number but quite narrow 

and more or less crescent-si aped; in Uva Ursi their number is usu- 

ally seven to eight and their arrangement radial around the stomata 

apparatus. 

On all dorsoventral leaves, the stomata arise more abundantly on 

the lower epidermis, less abundantly on the upper. Exceptions to 

this rule are due to the peculiar readaptation of the leaf to its sur- 

roundings. Thus, in the reversed types of leaves (twisted in a half 

circle) the stomata, formerly on the lower surface, have migrated to 

the upper surface which now has become the physiological lower 

surface. 

In Umbrophytic (shade) plants the stomata are either wholly on 

the lower surface or partly so with a number on the upper surface. 

Where the plants are Mesophytic and exposed to dense sunlight and 

leaves remain dorsoventral, the stomata are on the lower surface; 

these stomata are large, if the surroundings are damp. If such 

plants live in dry soil and dry air, the stomata are of small size and 

and dense 

numerous; if they dwell in dry soil in hot surroundings 

light they are very small and frequently sunk. If the plants are 

Xerophytic and the leaves dorsoventral, the stomata are quite abun- 

dant, small, with narrow slit, and depressed below the level of the 

epidermis. 

There are five types of stomatal development, viz.: 

First Type. Each primitive epidermal cell (or the majority, or 

only certain ones of the epidermis) at the close of the dermatogen 

stage, gradually lengthens and then cuts off a smaller from a larger 

cell. The smaller one is equilateral, has a very large nucleus, and is 

termed the Stoma Mother-cell; the larger, quadrangular, and called


the Epidermal Daughter-cell. The latter, upon maturing, becomes a 

normal epidermal cell. A partition is laid down lengthwise through 

the Stoma Mother-cell dividing it into two stomatal daughter-cells. 

The wall laid down lengthwise splits and thus forms the orifice of the 

stoma; 

the cells on either side of the orifice are called Guard Cells. 

FIG. 89. Types of stomatal apparatuses and neighboring cells from different 

sources. In A, a portion of the lower epidermis of Easter Lily leaf. The stomatal 

apparatus is surrounded by neighboring cells that are similar to other 

epidermal cells adjacent to them; in B, lower epidermis of Senna leaflet, note 

the two neighboring cells parallel to the guard cells, one being larger than the 

other; C, lower epidermis of Coca leaf showing two neighboring cells, parallel 

to the guard cells but nearly equal in size as well as papillated regular epidermal 

cells; D, lower epidermis of Pilocarpus showing rounded stomatal apparatus 

and four crescent-shaped neighboring cells; E, lower epidermis of Uva Ursi, 

showing eight neighboring cells arranged radiately around stomatal apparatus; 

F, lower epidermis of Stramonium. 

These, while at first flat and inoperative, soon become bulged and 

crescent-shaped. This mode of development is seen in Squill, 

Hyacinth, Daffodil, Sambucus, Silene, etc. 

Second Type. After the cutting off of the stomal mother-cell 

there are cut off on either side portions of neighboring epidermal cells 

which form subsidiary cells to the stoma. This condition is seen in 

Graminece, Cyperacece, Juncacece, in various species of Aloe, Musa and 

Proteacea.

.17 


Third Type. Instead of two parallel subsidiary cells, four are cue 

off, as in Heliconia, in species of Tradescantia, Araucaria, or four co 

five, as in Ficus elastica, or four to five or more, as in the Coniferce 

and Cycads. 

Fourth Type. Instead of only four subsidiary cells, each of these 

again subdivides by parallel walls, more rarely by radial walls, into 

eight radiating subsidiary cells, as in Maranta bicolor, Pothos argyraa, 

some of Proteacece, etc. 

Fifth Type. The "stomal mother-cell" divides once or several 

times before becoming the true mother-cell of the stoma. As a 

result of the divisions there are also formed one or more subsidiary 

cells. This mode of development is seen in the Labiates, Papilio- 

nacece, Cruciferce, Solanacece, Crassulacecz, Cactacece, and Begoniacece, 

also in a number of ferns. 

Histologic Differences between Leaves of Dicotyledons and 

Monocotyledons. The following may be cited as broad comparative 

histologic differences between Dicotyl and Monocotyl leaves: 

Dicotyl Leaves 

Epidermal cells usually iso-diametric 

or sinuous. 

The stomata are on the whole more 

numerous but smaller. 

Non-glandular and glandular hairs 

frequent, or upper but more frequent 

on lower surface, or both. 

Leaf glands which excrete varied 

products are rather abundant. 

Water stomata over the upper sur- 

face, more rarely over the lower 

surface, are frequent, especially 

along margins of leaves. 

Palisade and spongy parenchyma 

. in dicotyledons are more distinct 

and palisade parenchyma is 

denser. 

The vascular bundles, in their in- 

trinsic elements, are more indur- 

ated but the accessory fibrous 

sheath is feebly developed. 

A greater variety of accessory products 

of assimilation are de- 

veloped. 

. Epidermal 

. Stomata 

Monocotyl Leaves 

cells usually elongate 

and equilateral. 

larger. 

3. Hairs rare in MonocotyJs. 

4. Leaf glands rare and only seen as a 

rule on the sepals. 

5. Water stomata absent or very rare. 

Present in some Araccce. 

6. Palisade and spongy parenchyma 

are less distinct and dense. 

The vascular bundles, in their intrinsic 

elements, are less indur- 

ated. The fibrous sheath is 

strongly developed. 

A comparatively small variety of 

accessory products of assimilation 

are developed.


INFLORESCENCE 

Inflorescence or Anthotaxy. A typical flower consists of four 

whorls of leaves modified for the purpose of reproduction, and com- 

pactly placed on a stem. The terms Inflorescence and Anthotaxy 

are applied to the arrangement of the flowers and their position on 

the stem, both of which are governed by the same law which determines 

the arrangement of leaves. For this reason flower buds are 

always either terminal or axillary. 

In either case the bud may de- 

velop a solitary flower or a compound inflorescence consisting of 

several flowers. 

p 

p 

p 

p 

^ 

B 

FIG. 90. Types of indeterminate inflorescence. A, A raceme; B, a spike; C, a 

catkin; D, a corymb; E, an umbel. The flowers are represented by circles; the 

age of the flower is indicated by the size. (From Hamaker.) 

Determinate, cymose, descending, or centrifugal 

7 

inflorescence is that 

form in which the flower bud is terminal, and thus determines or 

completes the growth of the stem. Example: 

Ricinus communis. 

is that form in 

Indeterminate, ascending, or centripetal inflorescence 

which the flower buds are axillary, while the terminal bud continues 

to develop and increase the growth of the stem indefinitely. 

Exam- 

ple: the Geranium. 

Mixed inflorescence is a combination of the other two forms. 

Example: Horse Chestnut.

i 7 8 PHARMACEUTICAL BOTANY 

The flower stalk is known as the peduncle, and its prolongation the 

rachis, or axis of the inflorescence. 

The flower stalk of a single flower of an inflorescence is caHed a 

pedicel. When borne without such support the flower is sessile. 

A peduncle rising from the 

PIG. 91. Photomicrograph of longitudinal 

section through a staminate 

catkin of Comptonia asplenifolia X 10, 

showing catkin axis (ax) , anther-lobe (a) , 

and bract (6). 

ground is called a scape, 

previously mentioned under 

the subject of stems. 

The modified leaves found 

on peduncles are termed bracts. 

These vary much the same as 

leaf forms, are described in a 

similar manner, and may be 

either green or colored. When 

collected in a whorl at the 

base of the peduncle they 

form an involucre, the parts of 

which are sometimes imbri- 

cated or overlapping, like 

shingles. This is generally 

green, but sometimes petaloid, 

as in the Dogwood. The 

modified leaves found on pedi- 

cels are called bracteolar leaves. 

The Spathe is a large bract 

enveloping the inflorescence 

and often colored, as in the 

Calla, or membranous, as in 

the Daffodil. 

Indeterminate Inflores- 

cences. In the indeterminate 

or axillary anthotaxy, either 

flowers are produced from base to apex, those blossoming first 

which are lowest down on the rachis or from margin to center. The 

principal forms of this type are: 

Solitary Indeterminate. The simplest form of inflorescence in 

which a single flower springs from the axil of a leaf. A number of


these are generally developed on the same stem. Example: 

Periwinkle. 

Raceme, or simple flower-cluster in which the flowers on pedicels 

of nearly equal length are arranged along an axis. Examples: 

Convallaria, Cimicifuga, and Currant. 

Corymb, a short, broad cluster, differing from the raceme mainly in 

its shorter axis and longer lower pedicels, which give the cluster a 

flat appearance by bringing the individual florets to nearly the same 

level. Example: Cherry. 

Umbel, which resembles the raceme, but has a very short axis, and 

the nearly equal pedicels radiate from it like the rays of an umbrella. 

Many examples of this mode of inflorescence are seen in the family 

Umbellifera, as indicated by the name, including Anis~, Fennel and 

other drug-yielding official plants. 

A Spike is a cluster of flowers, sessile or nearly so, borne on an 

elongated axis. The Mullein and common Plantain afford good 

illustrations. 

The Catkin or Ament resembles the Spike, but differs in that it 

has scaly instead of herbaceous bracts, as the staminate flowers of the 

Oak, Hazel, Willow, Comptonia, etc. 

The Head or Capitulum is like a spike, except that it has the rachis 

ak in the 

shortened, so as to form a compact cluster of sessile flowers , 

Dandelion, Marigold, Clover, and Burdock. 

The Strobile is a compact flower cluster with large scales concealing 

the flowers, as the inflorescence of the Hop. 

The Spadix is a thick, fleshy rachis with flowers closely sessile or 

embedded on it, usually with a spathe or sheathing bract. Example : 

Calla, Acorus Calamus, Arum triphyllum. 

The compound raceme, particularly if irregularly compounded, is 

called a panicle. Ex. Hagenia abyssinica. 

Determinate Inflorescences. Determinate Anthotaxy is one in. 

which the first flower that opens is the terminal one on the axis, the 

other appearing in succession from apex to base or from center to 

margin. The principal varieties are: 

The Solitary Determinate, in which there is a single flower borne 

on the scape, as in the Anemone, or Windflower, and Hydrastis. 

The Cyme, a flower cluster resembling a corymb, except that the

i8o PHARMACEUTICAL BOTANY 

buds develop from center to circumference. Example: 

Elder. If 

the cyme be rounded, as in the Snowball, it is a globose cyme. 

A Scorpioid Cyme imitates a raceme, having the flowers pedicelled 

and arranged along alternate sides of a lengthened 'axis. 

A Glomerule is a cymose inflorescence of any sort which is condensed 

into a head, as the so-called head of Cornus florida. 

A Verticillaster is a compact, cymose flower cluster which resembles 

a whorl, but really consists of two glomerules situated in the axils 

of opposite leaves. Clusters of this kind are seen in Catnip, Hore- 

hound, Peppermint and other plants of the Labiates. 

o 

\ 

\ 

FIG. 92. Cymose inflorescences. F, A terminal flower; G, a simple cyme; H, 

a compound cyme. (From Hamaker.) 

The raceme, corymb, umbel, etc., are frequently compounded. 

The compound raceme, or raceme with branched pedicels, is called a 

panicle. Examples: Yucca and paniculate inflorescence of the Oat. 

A Thyrsus is a compact panicle, of a pyramidal or oblong shape. 

Examples: Lilac, Grape and Rhus glabra. 

A Mixed Anthotaxy is one, in which the determinate and indeterminate 

plans are combined, and illustrations of this are of frequent 

occurrence. 

The order of flower development is termed ascending when, as in 

the raceme, the blossoms open first at the lower point on the axis and 

continue to the apex. Examples: White Lily, and many other

PLANT ORGANS AND ORGANISMS l8l 

plants of the same family. In the cyme the development is centrifugal, 

the central florets opening first, while in the corymb it is 

centripetal, or from margin 

to center. 

PREFLORAT1ON 

Prefloration. By prefloration is meant the arrangement 

of the 

tioral envelopes in the bud. It is to the flower bud what vernation is 

to the leaf bud, the same descriptive terms being largely employed, 

as convolute, involute, revolute, plicate, imbricate, etc. 

In addition to those already defined, the following are important. 

Vahate Prefloration, in which the margins meet but do not overlap. 

Of this variety the induplicate has its two margins rolled 

inward as in Clematis. In the reduplicate they are turned outward, 

as the sepals of Althaea. 

Vexillary, the variety shown in the corolla of the Pea, where the 

two lower petals are overlapped by two lateral ones, and the four in 

turn overlapped by the larger upper ones. 

Contorted, where one margin is invariably exterior and the other 

interior, giving the bud a twisted appearance, as in the Oleander and 

Phlox. 

THE FLOWER 

The flower is a shoot which has undergone a series of changes so as 

to serve as a means for the propagation of the individual. 

A Typical or Complete Flower possesses four whorls of floral leaves 

arranged upon a more or less shortened stem axis called a receptacle, 

torus or thalamus. These whorls passing from periphery toward the 

center are: calyx, composed of parts called sepals; corolla, composed 

of parts termed petals; andrcecium, composed of parts called stamens 

or microsporopylls; and gyn&cium, composed of one or more parts 

termed carpels or megasporophylls. 

The stamens and carpels constitute the essential organs, and a 

flower is said to be Perfect when these are present and functional. 

A Hermaphrodite flower is one which possesses 

both stamens and 

carpels which may or may not be functionally active. In some cases 

the stamens may alone be functional while in others the carpels 

only may function.


A Regular Flower possesses parts of each whorl of the same shape 

and size, as the flower of Veratrum. 

It is Symmetrical when the parts of each whorl are of the same 

number, or multiples of the same number. 

An Imperfect Flower shows one set of essential organs wanting. 

When either petals or sepals, or both, are present in more than the 

usual number, the flower is said to be "double" as the cultivated 

PIG. 93. Diagrams of floral structures. A, Shows the relations of the floral 

parts in a hypogynous flower; B, the same in a perigynous flower; C, the same 

in an epigynous flower; D, a stamen; E, a simple pistil in longitudinal section; 

F, the same in cross-section; G, transitional forms between true petals (left) and 

true stamens (right); H, slight union of two carpels to form a compound pistil; 

/ and J, union of carpels more complete; K and L, cross-sections of compound 

pistils, of three carpels. In B: a, stamen; b, petal; c, sepal; d, pistil; e, receptacle; 

/, pedicel. In D: a, anther cell; b, connective; c, filament. In E: a, stigma; 

b, style; c, ovules; d, ovary. (From Hamaker.) 

Rose and Carnation. The doubling of flowers is brought about 

cultivation and is due either to the transformation of sta- 

through 

mens (as in cases cited), and occasionally of carpels into petals, to a 

division of the petals, or to the formation of a new series of petals. 

If the pistils are present and stamens wanting, the flower is called 

Distillate, or female; if it possesses stamens but no pistil, it is described 

as staminate, or male; if both are absent, neutral, as marginal flow- 

ers of Viburnum. Some plants, as the Begonias and Castor oil, bear 

both staminate and pistillate flowers, and are called Monoecious.

PLANT ORGANS AND" ORGANISMS 183 

When the staminate and pistillate flowers are borne on different 

plants of the same species, they are termed Dicecious, as the Sassafras 

and Willow. When staminate, pistillate and hermaphrodite flowers 

are all borne on one plant, as on the Maple trees, they are polygamous. 

of the flowers of 

Connation and Adnation. In the development 

primitive species of flowering plants, the parts of each whorl are disjoined 

or separate from each other. In many higher types, however, 

the parts of the same whorl frequently become partly or completely 

united laterally. This condition is termed connation, coalescence, 

cohesion or syngenesis. Illustrations of this may be seen in Belladonna, 

Stramonium and Uva Ursi flowers, where the petals have 

joined laterally to form gamopetalous corollas. When the one or 

more parts of different whorls are united, as of stamens with petals 

(Rhammus] or stamens with carpels (Apocynum) the union is called 

adnation or adhesion. 

The Receptacle. The Receptacle, Torus or Thalamus is a more or 

less shortened axis (branch) which bears the floral leaves. It is 

usually flat or convex, but may be conical and fleshy as in the Strawberry, 

concave as in the Rose and Fig or show a disc-like modification 

as in the Orange. The internodes of the receptacle in many 

species lengthen and separate various whorls. When the lengthen- 

ing of the internode occurs between calyx and corolla, as in Lychnis, 

the structure resulting is called an anthophore] if between corolla and 

andrcecium as in Passiflora, a gonophore\ if betwen androcecium and 

gyncecium as in Geum, a gynophore. If the flowers of the Umbel- 

lifers the receptacle elongates between the carpels producing the 

structure called a carpophore. 

The Perigone. The perigone or perianth is the floral envelope 

consisting of calyx and corolla (when present). 

When both whorls, i.e., calyx and corolla, are present the flower is 

said to be dichlamydeous; if only calyx is present, monochalmydeous . 

The Calyx. The Calyx is the outer whorl of modified leaves. Its 

parts are called Sepals, and may be distinct (Chorisepalous, from a 

Greek word meaning disjoined) or more or less united (Gamosepalous) 

. They are usually green foliaceous or leaf-like but may 

be brilliantly colored, hence the term petaloid (like the petals) is 

applied. Examples: Tulip, Larkspur, Columbine and Aconite.

184 


In a gamosepalous calyx, when the union of sepals is incomplete, 

the united portion is called the tube, the free portion, the limb, the 

orifice of the tube, the throat. 

In form the calyx may be regular or irregular; regular, if its parts 

are evenly developed, and irregular if its parts differ in size and 

shape. The more common forms are tubular, resembling a tube; 

rotate, or wheel-shape; campanula te, or bell-shaped; urceolate or 

urn-shape; hypocrateriform, or salver-shape; bilabiate, or twolipped; 

corresponding to the different forms of corolla, under which 

examples illustrating each will be given. 

The calyx usually remains after the corolla and stamens have 

fallen, sometimes even until the fruit matures in either case it is 

said to be persistent. 

If it falls with the corolla and stamens, it is 

deciduous, and if when the flower opens, caducous, as in the Poppy 

and May-apple. It often more or less envelops the ovary or base of 

the pistil, and it is important, in plant analysis, to note the presence 

or absence of such a condition, which is indicated in a description by 

the terms inferior, or non-adherent (hypogynous), when free from 

the ovary and inserted upon the receptacle beneath it (the most 

simple and primitive position) ; half-superior, or half-adherent (peri- 

gynous), when it partially envelops the ovary, as in the Cherry; 

superior or adherent (epigynous), when it completely envelops it, 

as in the Colocynth, etc. 

Sepaline Spurs. Occasionally some or all of the sepals may 

become pouched and at length spurred as nectar receptacles or as 

receptacles for other parts that are nectariferous. Thus, 

in Cru- 

ciferce we occasionally see a slight pouching of the two lateral sepals. 

These act as nectar pouches for the nectar secreted by the knobs or 

girdles surrounding the short lateral stamens. These become deep 

pouches in Lunaria while in others the pouches become elongated 

spurs. - 

Again, in Delphinium, the posterior sepal forms an elongated 

spur into which pass the two spurred nectariferous petals. In 

Aconitumthe same sepal, instead of being spurred, forms an enlarged 

hood-like body (galea) arching over the flower like a helmet; into 

this pass the two hammer-shaped nectariferous petals. 

Sepaline Stipules. These structures are well developed and easily 

traceable in the more primitive herbaceous members of the Rose


family. Thus in Potentilla, Fragaria, Geum, etc., in addition to the 

normal calyx of five sepals, there is a supplementary epicalyx also of 

five parts. The five lobes of the epicalyx may be as large or larger 

than the sepals or smaller up to the disappearing point. Upon 

examining a few flowers of Potentilla or Fragaria, it will be observed 

that not infrequently one, sometimes two lobes of the epicalyx are 

bifid, or deeply cleft, or separated completely into two parts. The 

explanation is that the five sepals, after evolving in the flower bud, 

form at their bases two lateral swellings or sep aline stipules, which, as 

they grow, fuse in adjacent pairs, one stipule of one sepal joining 

with the adjacent stipule of another sepal to form five lobes. 

Sepaline Position. As already noted the most simple and primi- 

tive position for the sepals in relation to the floral parts is hypogy- 

nous, in which the sepals are inserted directly into the enlarged floral 

axis (receptacle) below the petals, stamens and carpels. But in the 

more primitive herbaceous Rosacece, Leguminosa, etc., the floral 

axis forms a saucer-like transverse expansion which pushes out the 

sepals, petals and stamens on its edge. Thus originates the perigynous 

insertion of the sepals. In not a few higher Rosacea, Saxifraga- 

ctce, Crassulacea, etc., the saucer-like floral axis becomes deepened 

and contracted into a cup-shaped structure (Cherry, Peach, Almond, 

Plum, etc.), and on the edge of this cup the sepals as well as the petals 

and stamens are inserted at different levels. Finally, in the Apple, 

assumes a 

Pear, Quince, etc., the greatly hollowed-out receptacle 

vase-shaped form and closes over the top of the ovary, at the same 

time lifting the sepals, petals, and stamens above the ovary. Here 

the sepals are epigynous. 

The Corolla. The Corolla -is the inner floral envelope, usually 

delicate in texture, and showing more or less brilliant colors and 

combinations of color. Its parts are called Petals, and when the 

calyx closely resembles the corolla in structure and coloring they 

are together called the Perianth. The purpose of these envelopes 

is to protect the reproductive organs within, and also to aid in the 

fertilization of the flower, as their bright colors, fragrance and sac- 

charine secretions serve to attract pollen-carrying insects. 

Forms of the Corolla and Perianth. When the petals are not 

united with each other, the corolla is said to be Choripetalous, Apo-


petalous or Polypetalous. When more or less united, it is Camopeta- 

lous, often called Synpetalous. 

When the distinct petals are four in number, and arranged in the 

form of a cross, the corolla is called Cruciform. Example: Mustard 

and other plants belonging to the family Cruciferae. 

The Papilionaceous corolla is so called because of a fancied resemblance 

to a butterfly. The irregularity in this form is very striking, 

and the petals bear special names: the largest one is the vexillum, or 

standard; the two beneath it the alee, or wings; the two anterior, the 

carina or keel. Examples: Locust, Pea, and Clover. 

Orchidaceous flowers are of peculiar irregularity, combining calyx 

and corolla. The petal in front of stamen and stigma, which differs 

from the others in form and secretes nectar, is called the Labellum. 

Examples: Cypridedium and other Orchids. 

When calyx and corolla each consist of three parts closely resem- 

bling each other in form and color, as in the Tulip and Lily, the 

flower is called Liliaceous. 

The Ligulate or Strap-shaped corolla is nearly confined to the 

family Composite. It is usually tubular at the base, the remainder 

resembling a single petal. Examples : Marigold, 

and Arnica Flowers. 

Labiate, or Bilabiate, having two lips, the upper composed of two 

petals, the lower one of three. This form of corolla gives the name 

to the Labiata, while in the family Leguminosa this arrangement is 

sometimes reversed. The corolla may be either ringent, or gaping, 

as in Sage, or personate, when the throat is nearly closed by a projec- 

tion of the lower lip, as in Snapdragon. 

Rotate, Wheel-shaped, when the tube is short and the divisions of 

the limb radiate from it like the spokes of a wheel. Example: The 

Potato blossom. 

Crateriform, Saucer-shaped, like the last, except that the margin 

is turned upward or cupped. Example: Kalmia latifolia (Mt. 

Laurel). 

Hypocrateriform, or Saker-shaped (more correctly, hypocraterimorphous), 

when the tube is long and slender, as in Phlox or Trailing 

Arbutus and abruptly expands into a flat limb. The name is 

derived from that of the ancient Salver, or hypocraterium with the 

stem or handle beneath.

PLANT ORGANS AND ORGANISMS i8 7 

FIG. 94. Illustrating various forms of the corolla, i, Personate bilabiate 

corolla of Linaria; 2, cruciform corolla of Rocket; 3, campanulate corolla of 

Harebell; 4, infundibuliform corolla of Bindweed; 5, ringent bilabiate corolla of 

Larkspur; 7, Ligulate corolla of Chrysanthemum; 8, rotate corolla of Pimpernel; 

9, papilionaceous corolla of Irish Broom; 10, urceolate corolla of Heath.


When of nearly cylindrical form, the corolla is Tubular, as in the 

Honeysuckle, and Stramonium. 

Funnel-form (Infundibuliform), such as the corolla of the common 

Morning Glory, a tube gradually enlarging from the base upward 

into an expanded border or limb. 

Campanulate, or Bell-shaped, a tube whose length 

than twice the breadth, and which expands gradually from base to 

apex. Examples: Canterbury Bell, Harebell. 

is not more 

Urceolate, or Urn-shaped, when the tube is globose in shape and 

the limb at right angles to its axis, as in the official Uva Ursi, Chima- 

phila and Gaultheria. 

Caryophyllaceous, when the corolla consists of five petals, each 

with a long slender claw expanding abruptly at its summit into a 

broad limb. Examples: 

Carnation and other members of the Pink 

family. 

The Androecium or Stamen System. The andrcecium is the single 

or double whorl of male organs situated within or above the corolla. 

It is composed of stamens or microsporophylls. 

A complete stamen (Fig. g^D) consists of a more or less slender 

stalk portion called a filament and a terminal appendage called the 

anther or microsorus. The anther is generally vertically halved by 

an upgrowth of th'e filament, called the connective, dividing the anther 

into two lobes. 

Number of Stamens. When few in number, stamens are said 

to be definite; when very numerous, and not readily counted, they 

are indefinite. 

their number: 

The following terms are in common use to express 

Monandrous, for a flower with but one stamen. 

Diandrous, with two stamens. 

Triandrous, with three. 

Tetrandrous, with four. 

Pentandrous, having five. 

Hexandrous, six. 

Polyandrous, an indefinite number. 

The most primitive flowers have numerous stamens, but passing 

from these to those of more evolved families there occurs a gradual 

reduction from many to ten, as in Caryophyllacece, Leguminosa and


some Aceracea, these being in two circles. In Malvacea, Umbelliferce 

and other Apopetalous families as well as many Sympetalae, the 

number five is typical. But in Scropkulariacece, while five are devel- 

oped and fertile in Verbascum, four with a fifth staminode (sterile 

stamen) are found in the allied genus Celsia. In Pentstemon there 

are four didynamous fertile stamens and an equally long staminode. 

In Scrophularia the fifth staminode is reduced to a petaloid flap in 

the posterior part of the flower. In Linaria this exists only as a 

small knob at the base of the back part of the corolla and there 

secretes nectar. In most Scrophulariacece the fifth stamen is entirely 

absent and the four stamens left are didynamous; but in 

Calceolaria two of these are rudimentary and thread-like, the 

other two alone being well-developed and fertile. In Veronica 

three stamens are entirely absorbed and two only are left as 

fertile representatives. 

Insertion of Stamens. As to insertion the stamens may be : 

Hypogynous, when inserted upon the receptacle below the base of 

the pistil (see Fig. 93^4). 

Perigynous, when inserted on the calyx or corolla above the base of 

and lateral to the pistil (see Fig. 93$). 

Epigynous, when inserted above the ovary (see Fig. 936"). 

Gynandrous, when inserted upon the pistil, as in Orchids and Aris- 

tolochia. 

Proportions of the Stamens. The stamens may be of equal length; 

unequal, or of different length. 

Didynamous, when there are two pairs, one longer than the other. 

Example: Snapdragon. 

Tetradynamous, three pairs, two of the same length, the third 

shorter. Example: Mustard. 

Connation of Stamens. Terms denoting connection between stamens 

are: 

Monadelphous (in one brotherhood), coalescence of the filaments 

into a tube. Example : Lobelia. 

Diadelphous (in two brotherhoods), coalescence into two sets. 

Example: Glycyrrhiza. 

Triadelphous, with filaments united into three sets. Example: 

St. John's Wort.

I QO 


Polyadelphous, when there are several sets or branched bundles. 

Example: Orange. 

Syngenesious , when the anthers cohere. Example: Composites. 

Color of Stamens. In most species the color of these organs is 

seldom pronounced owing to their delicate structure. It varies 

from greenish-yellow to yellow to white, through pink, pinkish-red, 

red, purple, purple-blue to blue. It is yellow, for instance, in Sassa- 

fras, Cucumber and Golden Club; greenish-yellow, yellow to red, in 

Maples; yellow-pink to pink and pinkish-red, in some Mallows; in 

Azalea amena the filaments are crimson-purple and the anthers, pur- 

ple-blue; in the genus Scilla both filaments and anthers are blue. 

Gross Structure and Histology of the Filament. The filament 

may be cylindric as in the Rose, awl-shaped as in Tulip, flat and with 

a dilated base as in the Harebell, three-toothed as in Garlic, appen- 

diculate, when it bears an appendage as in Chatostoma, Alyssum, etc. 

The filament is covered with a protective epidermis containing 

stomata. Beneath this is a soft, loose cellular tissue, the mesophyll, 

and in the center a small vascular bundle, the pathway of food from 

the floral axis to the anther. In some cases the single bundle may 

split into two or three bundle parts. 

Gross Structure and Histology of the Anther. Each staminal leaf 

(microsporophyll) bears a special development or appendage as a 

rule on its extremity which is the anther or microsorus. This consists, 

fundamentally, of a median prolongation of the filament equal to the 

connective or placenta. This develops on either side a quantity of 

indusial tissue that grows out to form a covering substance that 

protects and carries two microsporangia on either side. An anther 

therefore consist of a median connectine or placenta, producing on 

either side two anther lobes or indusial expansions. Each anther lobe 

encloses two pollen sacs or microsporangia, which, in some cases, 

remain distinct up to the dehiscence (splitting open) of the anther. 

Thus in Butomus, the anthers show four pollen chambers up to the 

time of dehiscence. Again in various species of Lauracece, the 

anthers remain four lobed and dehisce by four recurved lids. But in 

the great majority of Angiosperms each pair of pollen sacs fuse before 

dehiscence, owing to the breaking down of the partition- between 

them, and so, at that time, show two-celled anthers. Still more


rarely the anthers may be two-celled in their young state and by the 

breaking down of the partition become one-celled, e.g., Malvacea, 

Externally the mature anther is bounded by an exothecium or epi- 

dermis, often swollen, where lines of dehiscence occur, which may 

develop stomata, also hairs. Within it is a combined layer or set of 

one to often two or three, sometimes five or six cell layers (Agave, 

etc.). of indusial and sporangial cells, the endothecium. The outermost 

one to three layers of this become spirally, annularly or stel- 

lately thickened to form the elastic tissue of the anther, which, by 

pressure against the delicate epidermis or exothecium, causes ultimate 

rupture of the anther wall. Within the innermost endothecial 

layer, bounding each sporangium, is the tapetum, a single-celled 

layer. This, near the time of dehiscence, undergoes breaking down 

or absorption by developing pollen or microspore cells. Filling the 

cavities of the four sporangia are the mature pollen grains. The 

connective shows in or near its center a vascular bundle with xylem 

uppermost and phloem downward, surrounded by thin-walled cellular 

tissue, from which the indusial and sporangial substance has 

matured by extension. 

Anther Dehiscence. This is the breaking open of the anther to 

discharge the pollen. 

When fully ripe the dividing partition between each pair of spor- 

angia usually becomes thinned, flattened and ultimately breaks down, 

while the elastic and resistant endothecium, steadily pushing against 

the more delicate and now shrinking exothecium causes rupture 

where endothecium is absent, namely along opposite lines of the 

anther wall. Thus arises a line of anther dehiscence called longitudinal 

anther dehiscence on either side of the anther sacs. In the 

.division Solanece of the family Solanacecz which includes Belladonna, 

in some of the Ericacece as Rhododendron and Azalea, etc., the 

anthers dehisce by small apical pores from which the pollen 

is shed. 

This kind of dehiscence is called apical porous dehiscence. Again, 

in Lauracece and Berberidacecz, the anthers dehisce by recurved valves. 

This is called valvular dehiscence. 

Moreover, in Malvacea the originally longitudinal anther is divided 

internally by a partition. It gradually swings on the filament so 

that eventually the anther is transverse and the partition becomes

I Q2 


absorbed, thus becoming a one-celled anther with transverse dehis- 

cence. in its mature state. 

Development of the Anther. Each stamen originates as a knob- 

like swelling from the receptacle between the petals and carpels. 

This swelling represents mainly future soral (anther) tissue. The 

filament develops later. When such a young sorus or anther is cut 

PIG. 95. Cross-section of a mature lily anther. The pairs of pollen chambers 

unite to form two pollen sacs, filled with pollen grains; s, modified epidermal cells 

at line of splitting. (From a Text-book of Botany by Coulter, Barnes, and Cowles. 

Copyrightjoy the American Book Company, Publishers.) 

across and examined microscopically, it shows a mass of nearly simi- 

lar cellular tissue in which the first observable changes are the 

following: 

The surface dermatogen cells become somewhat flattened and regular 

to form the future epidermis or exothecium of the anther. About 

the same time some cells, by more rapid division in the middle 

of the anther substance, give 

rise to the elements of the vascular 

bundle in the connective. Then, along four longitudinal tracts, 

rows of cells remain undivided or only divide slowly as they increase 

in size and around them cells divide and redivide to form the future 

endothecial and covering tissue to the four sporangia. Next, the 

four sporangial tracts of undivided cells cut off from their outer 

surfaces a layer of enveloping^ cells, the tapetum. This consists of 

richly protoplasmic cells that form a covering to the spore mother-cells


within. Each spore mother-cell undergoes division and redivision 

into four spore daughter-cells, at the same time that reduction in the 

chromatin substance takes place in these cells. Thus originate 

tetrads (groups of four) of spore daughter-cells inside spore mother-cell 

wall. These continue to enlarge, press against the mother-cell wall 

which becomes converted into mucilage and each of the tetrad cells 

becomes in time a mature microspore or pollen grain. 

During this time the entire anther is growing in size, the cells of 

the endothecium in one or more layers becomes thickened by lignin 

deposits to form a mechanical endothecium', the tapetum gradually 

breaks down and appears only , at length, as an irregular layer around 

the maturing pollen cells. When the anther is finally ripe the parti- 

tion between each pair of microsporangia becomes narrowed, flat- 

tened and ruptured and thus numerous microspores or pollen grains 

fill two cavities, one on either side of the connective. The micro- 

spores or pollen grains at first show only a thin clear cellulose layer, 

but from this, by a differentiation of the exterior film, the exospore 

layer becomes cut off. This becomes cuticular. The cellulose inner 

layer (endospore), remains unaltered. In the development of the 

exopore, one to several deficiencies are usually left in it through 

which the endospore may protrude later as the rudiment of the 

pollen tube. 

Attachment of Anther. The attachment of the anther to the filament 

may be in one of several ways, as follows : 

Innate, attached at its base to the apex of the filament. 

Adnate, adherent throughout its length. 

Versatile, when the anther is attached near its center to the top 

of the filament, so that it swings freely. The adnate and versatile 

are introrse when they face inward, extrorse when they face outward. 

Pollen. The pollen grains or microspores vary in form for differ- 

ent species and varieties and while they are averagely constant for 

these, nevertheless many exceptions have been recorded. The* fol- 

are some of the commoner forms : 

lowing 

Four Spore Daughter-cells, hanging together as in the Cat Tail 

(Typha) forming a pollen grain. 

Elongated, simple pollen grains as in Zostera. 

Dumb-bell-shaped, as the pollen of the Pines. 

13

194 

PHAR&ACEUTICAL BOTANY 

FIG. 96. Various forms of pollen grains. Pollen from Typha latifolia (A), 

Zea mays (B), Ambrosia elatior (C), Lilium philadelphicum (D), Pinus (E), 

Ranunculus bulbosus (F), Carpinus caroliniana (G), Althcea rosea (H), Oenoihera 

biennis, (7). All highly magnified. Drawing by Hogstad.

PLANT ORGANS AND ORGANISMS IQ5 

Triangular, as in the (Enotheras. 

Echinate, as in the Malvacea. 

Spherical, as in Geranium, Cinnamon and Sassafras. 

Lens-shaped, as in the Lily. 

Spinose, as in the Composite. 

Barrel-shaped, as in Polygala. 

Under the microscope the immature pollen grain generally consists 

of two membranes, an outer firmer one called the exospore, which 

may be variously marked and which possesses deficiencies in the 

form of "pores" or "clefts," and an inner delicate cellulose mem- 

brane called the endospore, which surrounds a protoplasmic interior 

in which are imbedded a nucleus, oil droplets and frequently starch 

or protein. 

Pollinia. These are agglutinated pollen masses which are common 

to the Orchidacea and Asclepiadacece. 

The pollen of many plants, notably certain species of Composite, 

Graminece and Rosacea, has been shown to be responsible for "Hay 

Fever." At the present time serums, extracts and vaccines are 

manufactured from pollen to be used in the treatment of this disease. 

The Gyncecium or Pistil System. This is the female system of 

organs of flowering plants. It may consist of one or more modified 

leaves called carpels. Each carpel or megasporophyll is a female 

organ of reproduction. In the Spruce, Pine, etc., it consists of an 

open 

leaf or scale which bears but does not enclose the ovules. In 

angiosperms it forms a closed sac which envelops and protects the 

ovules, and when complete is composed of three parts, the ovary or 

hollow portion at the base enclosing the ovules or rudimentary seeds, 

the stigma or japical portion which receives the pollen grains, and 

the style, or connective which unites these two. The last is non- 

essential and when wanting the stigma is called sessile. The carpel 

clearly shows its relations to the leaf, though greatly changed in 

form. The lower portion of a leaf, when folded lengthwise with the 

margins incurved, represents the ovary; the infolded surface upon 

which the ovules are borne is the placenta, a prolongation of the 

tip of the leaf, the stigma, and the narrow intermediate portion, the 

style. A leaf thus transformed into an ovule-bearing organ is called 

a carpel. The carpels of the Columbine and Pea are made up of

196 


single carpels. In the latter the young peas occupy a double row 

along one of the sutures (seams) of the pod. This portion corre- 

sponds to the infolded edge of the leaf, and the pod splits open along 

this line, called the ventral suture. 

Dehiscence, or the natural opening of the carpel to let free the 

contained seeds, takes place also along the line which corresponds 

to the mid-rib of the leaf, the dorsal suture. 

The gyncecium or Pistil may consist of a number of separate 

carpels, as in the buttercup or Nymphaea flowers, when it is said to 

be apocarpous, or the carpels composing it may be united together 

to form a single structure, as in the flowers of Belladonna and Orange, 

when it is called syncarpous. 

If the pistil is composed of one carpel, it is called monocarpellary; 

if two carpels enter into its formation, it is said to be dicarpellary; 

if three; tricar pellary; if many, poly car pellary. 

Compound Pistils are composed of carpels which have united to 

form them, and therefore their ovaries will usually have just as 

many cells (locules)- as carpels. When each simple ovary has its 

placenta, or seed-bearing tissue, at the inner angle, the resulting 

compound ovary has as many axile or central placentae as there are 

carpels, but all more or less consolidated into one. The partitions 

are called dissepiments and form part of the walls of the ovary. If, 

however, the carpels are joined by their edges, like the petals of a 

gamopetalous corolla, there will be but one cell, and the placenta 

will be parietal, or on the wall of the compound ovary. 

The ovules or megasori are transformed buds, destined to become 

seeds in the mature fruit. Their number varies from one to hun- 

dreds. In position, they are erect, growing upward from the base 

of the ovary, as in the Compositae; ascending, turning upward from 

the side of the ovary or cell; pendulous, like the last except that 

they turn downward; horizontal, when directed straight outward; 

suspended, hanging perpendicularly from the top of the ovary. 

In Gymnosperms the ovules are naked; in Angiosperms they are 

enclosed in a seed vessel. 

A complete angiospermous seed ovule which has not undergone 

maturation consists of a nucellus or body; two coats, the outer and 

inner integuments; and a funiculus, or stalk. Within the nucellus


is found the embryo sac or megaspore containing protoplasm and a 

nucleus, (See Fig. 97 A). 

The coats do not completely envelop the nucellus, but an opening 

at the apex, called the foramen or micropyle admits the pollen tube. 

The vascular plexus near the point where the coats are attached to 

each other and to the nucellus is called the chalaza. The hilum 

marks the point where the funiculus is joined to the ovule, and if 

attached to the ovule through a part of its length, the adherent 

portion is called the raphe. The shape of the ovule may be ortho- 

tropous, or straight; campylotropous, bent or curved; amphitropous, 

partly inverted; and anatropous, inverted. The last two forms are 

most common. A campylotropous ovule is one whose body is bent 

so that the hilum and micropyle are approximated. 

The Placenta. The placenta is the nutritive tissue connecting 

the ovules with the wall of the ovary. The various types of pla- 

centa arrangement (placentation) are gouped according 

to their 

relative complexity as follows: (i) Basilar, (2) Sutural, (3) Parietal, 

(4) Central, (5) Free Central. 

Basilar placentation is well illustrated in the Polygonacea (Smart 

Weed, Rhubarb, etc.) in Piper and Juglans. Here, at the apex of the 

axis and in the center of the ovarian base, arises a single ovule from 

a small area of placental tissue. 

Sutural placentation is seen in the Leguminosce (Pea, Bean, etc.). 

Here each carpel has prolonged along its fused edges two cord-like 

placental twigs, from which start the funiculi or ovule stalks. 

Parietal placentation is seen in Gloxinia, Gesneria, Papaver, etc. 

Here we find two or more carpels joined and placental tissue running 

up along edges of the fused carpels bearing the ovules. 

Central or axile placentation is seen in Campanulacece (Lobelia), 

where the two, three, or more carpels have folded inward until they 

meet in the center and in the process have carried the originally 

parietal placenta with them. This then may form a central swelling 

bearing the ovules over the surface. 

Free Central placentation occurs perfectly in the Primulacece, 

Plantaginacece and a few other families. In this the carpels simply 

cover over or roof in a central placental pillar around which the 

ovules are scattered.


Style. The style is the portion of the carpel which connects the 

stigma with the ovary. It is usually thread-like but may also be con- 

siderably thickened. It frequently divides into branches in its upper 

part. These are called style arms. As many style arms as carpels 

may be present. In the one-carpelled pistil of some Leguminosa, the 

usually bent-up style is the tapered prolongation of a single flower. 

Again, in the apocarpous carpels of many flowers of the Ranunculacea, 

each carpel bears a short to long stylar prolongation. When 

the carpels, however, are syncarpous the common styles tend to 

become more or less fused but usually show lobes, clefts or style 

arms at their extremities that indicate the number of carpels in 

each case which form the gynoecium. 

In some plants remarkable variations from the typical stylar 

development may occur. Thus, in Viola, the end of the style is a 

swollen knob on the under surface of which is a concave stigma with 

a flap or peg. In the genus Canna the style is an elongate blade-like 

flattened body with a sub-terminal stigma. In forms of the Cam- 

panulacea the style is closely covered with so-called collecting hairs. 

On these the anthers deposit their pollen at an early period before 

the flowers have opened. Later, when the flowers open, insects 

remove the pollen after which the collecting hairs wither. The stigmas 

then curl apart to expose their viscid stigma tic hairs. In this 

instance there are two distinct and at separate times functioning 

hairs on the stylar prolongation, viz.: (a) collecting stylar hairs, 

functioning for pollen collection and distribution; and (b) stigmatic 

hairs for pollen reception from another flower. In Vinca the style 

swells near its extremity into a broad circular stigma and then is 

prolonged into a short column bearing a tuft of hairs that prevents 

the entrance of insect thieves into the flower. In the genus Iris 

the common style breaks up at the insertion of the perianth into 

three wide petaloid style arms. Each of these bifurcates at its ex- 

tremity. On the lower or outer face of this is a transverse flap that 

bears the stigmatic papillae. In Physostigma the style enlarges at 

its extremity into a flap-like swelling which bears a narrow stigmatic 

surface. Finally in Sarracenia the single style of the five-carpelled 

pistil enlarges above into a huge umbrella-like portion with five 

radiating ribs. At the extremity of each bifid end of each rib is a 

minute peg-like stigmatic surface.


The Stigma. This is usually a viscid papillose surface of greater or 

less expanse functioning for pollen reception. In wind-pollinated 

flowers such as the grasses, the stigmas are the numerous feathery 

hairs which cover the ends of the styles and intended to catch flying 

pollen grains. In animal-pollinated flowers, the stigmas are usually 

small restricted knobs, lines or depressions. The stigmatic papillae 

vary in size in different plants and even may vary on different 

individuals of the same species. Thus in the long styles of Primula, 

the stigmatic papillae are elongated columnar hair-like structures, 

whereas in the short styles of short-styled flowers the papillae are 

small knob-like cellular swellings. 

POLLINATION 

Pollination is the transfer of pollen from anther to stigma and 

the consequent germination thereon. It is a necessary step to 

fertilization. 

When the pollen is transferred to the stigma of its own flower the 

process is called Close or Self-pollination', if to a stigma of another 

flower, Cross-pollination. If fertilization follows, these processes are 

termed respectively, Close or Self-fertilization and Cross-fertilization. 

Close-fertilization means in time ruination to the race and happily is 

prevented in many cases by (a) the stamens and pistils standing in 

extraordinary relation to each other, (b) by the anthers and pistils 

maturing at different times, (c) by the pollen in many cases germinating 

better on the stigma of another flower than its own. 

The agents which are responsible for cross-pollination are the 

wind, insects, water currents, small animals, and man. 

Wind-pollinated, flowering plants are called Anemophilous; their 

pollen is dry and powdery, flowers inconspicuous and inodorous, as 

in the Pines, Wheat, Walnut, Hop, etc. 

Insect-pollinated plants are called Entomophilous. These, being 

dependent upon the visits of insects for fertilization, possess bril- 

liantly colored corollas, have fragrant odors, and secrete nectar, a 

sweet liquid very attractive to insects, which are adapted to this work 

through the possession of a pollen-carrying apparatus. Example: 

Orchids.


Plants pollinated through the agency of water currents are known as 

Hydrophilous. To this class belong such plants as live under water 

and which produce flowers at or near the surface of the same. Exam- 

ple: Sparganium. 

Animal-pollinated plants are called Zoophilous. Some plants like 

the Nasturtium and Honeysuckle are pollinated by humming birds. 

Before the pollen grain has been deposited upon the stigma and 

during its germination thereon, a series of events affecting both the 

pollen grain and the embryo sac occur which result in the ultimate 

formation of the male and female gametophytes. 

Maturation of the Pollen Grain and Formation of the Male 

Gametophyte. The substance of the microspore (pollen grain) 

divides into two cells, the mother and tube cells of the future male 

gametophyte. The nucleus of the mother-cell divides to form two 

sperm nuclei. Within the tube cell is found a tube nucleus embedded 

in protoplasm. Upon germinating the partition disappears 

and the thin endospore, carrying within it the protoplasm in which 

are embedded the tube nucleus and two sperm nuclei, penetrates 

through a deficiency of the exospore. The contents of the pollen 

grain at this stage is called the male gametophyte. 

Maturation of the Embryo Sac and Formation of the Female 

Gametophyte. The nucleus of the megaspore or embryo sac under- 

goes division until eight daughter -nuclei are produced which are 

separated into the following groups: 

(a) Three of these nuclei occupy a position at the apex, the lower 

nucleus of the group being the egg or ovum, the other two nuclei 

being the synergids or assisting nuclei. 

(b) At the opposite end of the sac are three nuclei known as the 

antipodals which apparently have no special function. 

(c) The two remaining nuclei (polar nuclei) form a group lying 

near the center of the embryo sag which unite to form a single 

endosperm nucleus from which, after fertilization, the endosperm or 

nourishing material is derived. This stage of the embryo 

sac con- 

stitutes the female gametophyte. 

Fertilization in Angiosperms. After the pollen grain reaches the 

stigma, the viscid moisture of the stigma excites the outgrowth of the 

male gametophyte which bursts through the coats of the pollen

PLANT ORGANS AND ORGANISMS 2OI 

grain forming a pollen tube. The pollen tube, carrying within its 

walls two sperm nuclei and one tube nucleus, penetrates through the 

loose cells of the style until it reaches the micropyleof the ovule, then 

piercing the nucellus, it enters the embryo sac. The tip of the tube 

breaks and one of the sperm nuclei unites with the egg to form the 

oospore. The oospore develops at once into an embryo or plantlet, 

mic 

A B C 

PIG. 97. A, Immature angiospermous ovule; B, same, after embryo-sac (e.s). 

has matured to form the female gametophyte; nucellus (nuc); outer integument 

(o. int); inner integument (i. int); embryo sac (e.s.); micropyle (mic); chalaza 

(ch); funiculus (/); synergids (5); ovum (o); polar nuclei (p); antipodals (a),- C, 

fertilized and matured angiospermous ovule (seed). Note that the nucellus 

(nuc) has been pushed out by the encroachment of the embryo sac, in which 

endosperm has formed by the fusion of the two polar nuclei with the second 

sperm nucleus from the pollen tube which has later divided to form numerous 

nuclei scattered about in the protoplasm of the embryo sac and accumulated 

protoplasm and laid down walls, within which nourishment was stored; embryo 

(em) from fertilized ovum; testa (t) from outer integument; tegmen (te) from 

maturation of inner -integument; micropyle (mic); hilum or scar (h), after funiculus 

became detached. 

which lies passive until the seed undergoes germination. The other 

sperm nucleus unites with the previously fused polar nuclei to form 

the endosperm nucleus which soon undergoes rapid division into a 

large number of nuclei that become scattered about through the protoplasm 

of the embryo sac. These accumulate protoplasm about 

them, cells walls are laid down, endosperm resulting.


THE FRUIT 

The fruit consists of the matured pistil (carpel or carpels) and 

contents, or ovarian portion thereof, but may include other organs 

of the flower which frequently are adnate to and ripen with it. 

Thus in the Apples, Pears and Quinces, the receptacle becomes thick 

and succulent, surrounds the carpels during the ripening process and 

forms the edible portion of these fruits. In Dandelion, Arnica, and 

many other members of the Composite, the modified calyx or pappus 

adheres to the ovary during its maturation into the fruit and renders 

the fruit buoyant. In Gaultheria the calyx becomes fleshy, sur- 

rounds the ovary, reddens, and forms the edible part of the fruit. In 

Physalis the calyx enlarges considerably and encloses the ovary in an 

inflated colored bladder. Involucres frequently persist around and 

mature with the fruits. These may be membranous as in Anthemis, 

Matricaria and other Composite, leathery and prickly as in the Chest- 

nuts, scaly woody cups (cupules) as in the Oaks, or foliaceous cups as 

in the Filberts. Occasionally, as in the Fig, Osage Orange, Mul- 

berry, etc., the fruit may consist of the ripened 

inflorescence. 

FRUIT STRUCTURE 

flower cluster or 

The Pericarp, or seed vessel, is the ripened wall of the ovary, and in 

general the structure of the fruit wall resembles that of the ovary, but 

undergoes numerous modifications in the course of development. 

The number of cells of the ovary may increase or decrease, the 

external surface may change from soft and hairy in the flower to hard, 

and become covered with sharp, stiff prickles, as in the Datura 

Stramonium or Jamestown weed. Transformations in consistence 

may take place and the texture of the wall of the ovary may become 

hard and bony as in the Filbert, leathery, as the rind of the Orange, 

or assume the forms seen in the Gourd, Peach, Grape, etc. 

Frequently the pericarp consists for the most part of other ele- 

ments than the ripened ovarian wall and is then termed a pseudocarp 

or anthocarp. 

The pericarp consists of three layers of different tex- 

ture, viz.: epicarp, mesocarp and endocarp. The epicarp is the 

outer layer. The mesocarp the middle, and the endocarp the inner

layer. 


When the mesocarp is fleshy, as in the Peach, it is called the 

sarcocarp. 

When the endocarp within the sarcocarp is hard, forming a shell or 

stone, this is termed a putamen. 

Sutures. The ventral suture is a line formed by the coherent 

edges of a carpellary leaf. The dorsal suture is the mid-rib of the 

carpel. 

Parietal sutures are lines or furrows frequently visible on the 

walls of fruits, formed by the ripening of a compound ovary. They 

occur between its dorsal sutures and indicate the places of union 

between adjacent septa or of two parietal placentae. 

Valves. These are the parts into which the mature fruit separates 

to permit the escape of the seeds. Depending upon the number of 

these the fruit is said to be univalved, bivalved, trivahed, etc. 

ABC 

FIG. 98. Diagrams illustrating three forms of valvular dehiscence. A, Loculicidal 

dehiscence showing each carpel split along its midrib or dorsal suture, 

the dissepiments remaining intact; B, septicidal dehiscence, in which splitting 

took place along -the partitions; 

broke away from the partitions. 

C, septifragal dehiscence, in which the valves 

Dehiscence. This is the opening of the pericarp to allow the 

seeds to escape. 

Fruits are either Dehiscent or Indehiscent according as they open to 

discharge their seeds spontaneously when ripe (dehiscent), or decay, 

thus freeing the seeds, or retain their seeds, the embryo piercing 

the pericarp in germination (indehiscent) . Dehiscent fruits open regularly, 

or normally, when the pericarp splits vertically through the 

whole or a part of its length, along sutures or lines of coalescence of 

contiguous carpels. Legumes usually dehisce by 

both sutures. 

Irregular or abnormal dehiscence has no reference to normal sutures, 

as where it is transverse or circumscissile, extending around the cap-

2O4 


sule either entirely or forming a hinged lid, as in Hyoscyamus, or 

detached. 

Dehiscence is called porous or apical when the seeds escape through 

pores at the apex, as in the Poppy; valvular, when valve-like orifices 

form in the wall of the capsule. Valvular dehiscence is septicidal, 

when the constituent carpels of a pericarp become disjoined, and 

then open along their ventral suture. Example: Colchicum; loculi- 

cidal, dehiscence into loculi, or cells, in which each component carpel 

splits down its dorsal suture, and the dissepiments remain intact. 

Example: Cardamon; septifragal dehiscence, a breaking away of the 

valves from the septa or partitions. Example: Orchids (Fig. 98). 

Classification of Fruits (according to structure). Simple fruits 

result from the ripening of a single pistil in a flower. 

Aggregate fruits are the product of all the carpel ripenings in one 

flower, the cluster of carpels being crowded on the ripened receptacle 

forming one mass, as in the Raspberry, Blackberry, and Strawberry. 

Multiple fruits are those which are the product of the ripening 

of a flower cluster instead of a single flower. 

Simple and Compound fruits are either Dry or Fleshy. The first 

may be divided into Dehiscent, those which split open when ripe; 

and Indehiscent, those which do not. 

Simple Fruits: 

f I. Capsular (dehiscing). 

n. 

Dry< II. Schizocarpic (splitting). 

[III. Achenial (indehiscent). 

f IV. Baccate (berries). 

Succulent , 

\ r ^ r ., N 

[ V. Drupaceous (stone fruits). 

The capsular fruits include all of those, whether formed of one 

or more carpels, which burst open to let their seeds escape. 

Schizocarpic or splitting fruits are those in which each carpel or 

each half carpel (in Labiatae) splits asunder from its neighbor and 

then falls to the ground. The split portion is one-seeded. 

Achenial fruits are dry, one-celled, one-seeded and indehiscent 

at the time of final ripening. 

Baccate fruits are such in which the endocarp always 

and the 

mesocarp usually becomes succulent and so the seeds lie in the pulp 

formed by the endocarp or endocarp and mesocarp combined.


Drupaceous fruits are those in which the endocarp is always 

fibrous or stony in consistence, while the mesocarp is more or less 

succulent. The endocarp may become cuticularized as in the 

Apples. The mesocarp may form stone cells lying in the midst of 

FIG. 99. Types of capsular fruits, i, Pod of Aconite; 2, Capsule of Colchicum 

showing septicidal dehiscence; 3, capsule of poppy, having porous dehiscence; 

4, pyxis of Henbane ;5, dehiscing regma of Geranium; 6, siliqua of Celandine 

showing valves opening from below upward; 7, silicule of Cochlearia; 8, 

egume of Pea. 

soft parenchyma cells as in Pears; it may become hardened and 

thickened by lignin deposits to form fibers as in the Cocoanut, or 

it may become swollen and soft-succulent as in Peaches, Cherries, etc. 

I. Capsular Fruits. These may be simple, when composed of one 

carpel as the follicle and legume, or compound, when composed of 

two or more carpels as the capsule, pyxis, regma, siliqua or silicule


The Follicle or pod is a dry, simple capsular 

fruit formed of a 

single carpel which dehisces by one suture. This is usually the 

ventral suture as in Aconite, Staphisagria, Larkspur and some other 

Ranunculacece, but may be the dorsal suture as in Magnolia, 

Fig. 99 (i). 

A Legume is a dry simple capsular fruit formed of a single carpel 

and dehiscent by both ventral and dorsal sutures. Examples: Peas, 

Beans, etc. The legume is typical of most Leguminosa, Fig. 99 (8). 

A Capsule is a fruit formed of two or more carpels which dehisce 

longitudinally or by apical teeth or valves. Examples: Cardamon, 

Poppy, Iris, etc., Fig. 99 (2 and 3). 

A Pyxis or Pyxidium is a capsular fruit formed of two or more 

carpels that dehisce transversely. Examples: Hyoscyamus, Portu- 

laca. The upper portion forms a lid which fits upon the lower pot- 

like portion, Fig. 99 (4). 

A Regma is a capsular fruit of two or more carpels that first splits 

into separate parts and then each of these dehisces. This type of 

fruit is typical of Euro, crepitans (Sandbox), Pelargonium and 

Geranium, Fig. 99 (5). 

A Siliqua is a long slender one or two-celled capsule, often with a 

spurious membranous septum (when two-celled) and two persistent 

parietal placentae, the valves opening from below upward. Ex- 

amples: Chelidonium and Wallflower, Fig. 99 (6). 

A Silicule is a short siliqua in which the length is never much 

greater than the breadth. Example: Cochlearia. Fig. 99 (7). 

II. Schizocarpic Fruits. A Carcerulus or Nutlet is the typical 

fruit of the Labiatae but is also seen in the Borraginaceae. The ovary 

that has become four-celled at the time of flowering matures into 

four little pieces which split asunder lengthwise. Each split part 

is composed of one-half of a ripened carpel, Fig. 100 (2). 

A Cremocarp is the characteristic splitting fruit of the 

Umbelliferse family. It consists of two inferior akenes or 

mericarps separated from each other by a forked stalk called a 

carpophore. These mericarps usually cling to the forks of the 

carpophore for a time after the cremocarp splits, 

later fall, Fig. 100 (i). 

but sooner or


A Samara is a winged splitting fruit. It may be one-carpelled 

as in the Elm, Ash, Tulip Poplar and Wafer Ash or two-carpelled 

as in the Maples, Fig. 100 (3 and 4). 

A Lomentum or Loment is a splitting fruit that splits transversely 

into one-seeded portions. Seen in Crucifera, in Entada scandens, 

Cathartocarpus Fistula, Desmodium, etc. of Leguminosa, Fig. 100 (5). 

FIG. 100. Schizocarpic fruits. I, Cremocarp of Fennel, composed of two 

mericarps (m) and a split carpophore (c); 2, carcerulus of the Bugle; 3, one-carpelled 

samara of the Ash; 4, double samara of the maple; 5, loment of purging 

cassia, a portion of the pericarp being removed to show chambers, each containing 

a single seed. 

III. Achenial Fruits (all indehiscent). The Akene is a dry one- 

chambered, indehiscent fruit, in which the pericarp is firm and may 

or may not be united with the seed, the style remaining in many 

cases as an agent of dissemination, Fig. 101 (i). The latter may 

be long and feathery as in Clematis or be hooked. Examples of 

akenes: Fruits of the Composite, Anemone, etc. The Hip 

of the


Roses consists of a number of akenes in a ripened concave 

receptacle. 

The Utricle is like the akene, except that the enveloping calyx is 

loose and bladder-like. Example: Chenopodium, Fig. 101 (3)'. 

A Caryopsis or Grain is similar to an akene but differs from it by 

the pericarp being always fused with the seed coat. This fruit is 

- Ctt 

FIG. 101. Achenial fruits, i, Akene of Pulsatilla cut vertically, showing 

adherent feathery style (si), pericarp (p), testa (0, endosperm (e), hypocotyl (h) 

and cotyledons (cot) the last two structures making up the embryo; 2, caryopsis 

of wheat showing beard of hairs above and position of embryo of seed below; 

3, utricle of Chenopodium cut vertically to show calyx (c), pericarp (p) and seed 

(s) regions; 4, nut of an oak consisting of a glans (g) and cupule (CM). 

more likely than any other to be mistaken for a seed. Examples: 

Wheat, Corn, Barley, Oats and other members of the Graminea, 

Fig. 101 (2). 

A Nut or Glans is a one-celled, one-seeded fruit with a leathery 

or stony pericarp. Examples: Oaks, Beeches, Chestnuts, Alders 

and Palms, Fig. 101 (4). 

IV. Baccate Fruits (Succulent fruits in which the endocarp is 

always succulent and the mesocarp sometimes). The Berry 

small fleshy fruit with a thin membranous epicarp and a succulent 

is a 

interior in which the seeds are imbedded. Examples: Capsicum, 

Tomato, Belladonna, Grape, Currant, etc.


An Uva is -a berry from a superior ovary. Examples: Bella- 

donna, Egg-plant, Tomato, Fig. 102 (i). 

A Bacca is a berry from an inferior ovary. Examples: Goose- 

berry, Honeysuckle, Currant. 

The Pepo or Gourd Fruit is a baccate fruit of large size which has 

developed from an inferior ovary. It is fleshy internally and has a 

tough or very hard rind. Examples: Fruits of the Cucurbitacea 

and the Banana, Fig. 102 (2). 

PIG. 1 02. Baccate fruits, i, berry (uva) of Belladonna with adherent 

calyx; 2, Pumpkin, cut transversely illustrating a pepo fruit; (h), a locule; 3, 

hesperidium fruit of the Orange cut transversely showing epicarp (e) , mesocarp 

(m), endocarp (en}, pulp (p), and seed (s) . 

The Hesperidium is a large 

thick-skinned succulent fruit with 

seeds embedded in the pulp but from a superior ovary. Examples: 

Orange, Grape-fruit, Lemon, etc. In each of these there is to be 

noted a glandular leathery epicarp, a sub-leathery mesocarp and 

an endocarp in the form of separate carpels. From the endocarp 

hairs grow inward into the carpellary cavities and become filled with 

succulence. The seeds lie amid the hair cells, Fig. 102 (3). 

V. Drupaceous Fruits (Succulent fruits in which the mesocarp 

is more or less succulent, but the endocarp leathery or stony). 

A Drupe is a one-celled, one-seeded drupaceous fruit such as the 

fruit of the Plum, Peach, Prune, Sabal, Rhus, Piper, Cherry, etc., 

whose endocarp or putamen is composed wholly 

of stone cells of 

stone cells and sclerenchyma fibers, Fig. 103 (i). 

The Pome is a fleshy drupaceous fruit, two or more celled with 

fibrous or stony endocarp, the chief bulk of which consists of the 

14


adherent torus. Quince, Apple and Pear are examples. The car- 

pels constitute the core, and the fleshy part is developed from the 

torus, Fig. 103 (2). 

FIG. 103. i, Drupe of cocoanut cut vertically, showing epicarp (e), mesocarp 

(m), stony endocarp (d) seed coat (5), endosperm (end), and embryo sac cavity 

(e.s.) which in the mature seed contains a nutritive fluid. 2. Pome of an apple 

cut vertically to show core composed of 5 ripened carpels and flesh of matured 

torus. 3, Eaetrio of raspberry. 4, Same, cut vertically to show arrangement of 

the little drupes on fleshy receptacle. 

FIG. 104. Multiple fruits, i, Syconium of Fig cut vertically to show hollowed 

out receptacle (r) of ripened flower cluster; 2, strobile of the hop; 3, galbalus of 

Juniper. 

Aggregate 

Fruits. An Etaerio consists of a collection of little 

drupes on a torus of a single flower. Examples: Raspberry, Black- 

berry, etc., Fig. 103 (3 and 4). .


Multiple Fruits. The Syconium is a multiple fruit consisting of a 

succulent hollow torus enclosed within which are akene-like bodies, 

products of many flowers. Example: Fig. 104 (i). 

The Sorosis is represented by the Mulberry, Osage Orange, etc., 

the grains of which are not the ovaries of a single flower, as in the 

Blackberry, but belong to as many separate flowers. In the Pineapple 

all the parts are blended into a fleshy, juicy, seedless mass, and 

the plant is prop agated by cuttings. 

The Strobile or Cone is a scaly, multiple fruit consisting of a 

scale-bearing axis, each scale enclosing one or more seeds. The 

name is applied to the fruit of the Hop, Fig. 104 (2), and also to 

the fruit of the Conifers in which the naked seeds are borne on 

the upper surface of the woody scales. 

A Galbalus is a more or less globular multiple fruit formed of 

fleshy connate scales, as in Juniper, Fig. 104 (3). 

Histology of a Capsule, 

Vanilla. The Vanilla fruit is a one-celled 

capsule formed by the union of three carpellary leaves and dehiscing 

by two unequal longitudinal valves. 

Microscopic Appearance of a Transverse Section. Passing from 

periphery toward the center, the following structures present 

themselves : 

1. Epicarp, consisting of epidermis and hypodermis. The epidermis 

consists of a layer of thick-walled epidermal cells whose outer 

walls show the presence of a thin yellow cuticle. Stomata are present 

in this tissue. The epidermal cells contain protoplasm and 

brownish bodies. Some also contain small prisms of calcium oxalate 

and a few, vanillin crystals. The hypodermis is composed of one to 

several layers of collenchymatic cells with dark-colored contents. 

Its cells are somewhat larger than those of th e epidermis and thicker- 

walled. 

2. Mesocarp, a broad region of somewhat loosely arranged large, 

thin-walled parenchyma cells becoming smaller in the inner zone of 

this region. Most of these cells contain brownish contents but some 

possess long raphides of calcium oxalate. If the section be mounted 

in phloroglucin solution (5 per cent.) and a drop of strong sulphuric 

acid is added, a carmine-red color will be observed showing the pres- 

ence of vanillin in this region. Several closed collateral bundles will 

be seen coursing through the mesocarp.


3. Endocarp, an irregular line of inner epidermal cells which is 

differentiated into two regions, the interplacental region and the 

placental region. The interplacental (inner) epidermis shows its 

cells elongated into numerous thin-walled glandular hairs which con- 

tain an abundance of balsam; the placental region 

covers the six 

bifid placenta which extend into the cavity of the capsule. Its 

(inner) epidermis is composed of mucilaginous cells. 

4. Seeds. These are minute blackish bodies attached to the placental 

twigs of the placentae. Some of them may have been torn off 

in cutting the section. 

FIG. 105. Photomicrograph of a transverse section of a mericarp of Foeniculum 

vulgare, showing epicarp (A), mesocarp (B), endocarp (F), vitta (C), endosperm 

of seed (>), carpophore (G) and fibro-vascular bundle in primary rib (). 

(Highly magnified.) % 

Histology of a Typical Mericarp, Foaniculum. This five-angled 

fruit, in transverse section, shows a concave commissural and convex 

dorsal surface. Passing 

from the surfaces toward the center we note : 

1. Epicarp, or outer covering tissue, composed of colorless epider- 

mal cells and small stomata. The epidermal cells in cross-section 

appear rectangular, while in surface view they are both polygonal 

and rectangular. 

2. Mesocarp, of several layers of thin-walled colorless isodia- 

metric cells, beneath which are two to several additional layers of 

thicker-walled cells with brownish walls. Through the angles or 

rib portions of the mesocarp extend the fibro-vascular bundles.


Between each nbro-vascular bundle and the tip of each rib will be 

found a zone of collenchyma cells. In the mesocarp between each 

two ribs on the dorsal side occurs a single mtta or oil tube which is 

lined with a layer of brownish polygonal cells. These vittae contain 

the official oil of fennel. Two vittae generally occur in the meso- 

carp of the commissural side although four are reported to have 

been found in this region of some fennel fruits. 

3. Endocarp, a narrow layer of cells, transversely elongated, except 

over the regions of bundles where they may be seen elongated in 

several directions. 

4. Spermoderm or Seed Coat, consisting of a layer of somewhat 

broadened epidermal cells attached to the endocarp and several 

layers of collapsed cells which are only well denned in the region of 

the rap he. 

5. Endosperm, a central mass of more or less polygonal cells containing 

aleurone grains and oil globules. Each aleurone grain con- 

tains a rosette aggregate of calcium oxalate and one or two globoids. 

6. Embryo, embedded in the endosperm of the upper region of the 

seed. 

THE SEED 

A seed is a matured megasorus (ovule) borne by the sporophyte of 

a spermatophytic plant, enclosing a megaspore (embryo sac) within 

which a fertilized egg of the succeeding gametophyte generation has 

segmented to form a new sporophyte plant. The purpose of the 

seed is to insure the continuation and distribution of the species/ 

Like the ovule, it consists of a nucellus or kernel enclosed by integuments, 

and the descriptive terms used are the same. The seed 

coats, corresponding to those of the ovule, are one or two in number. 

If but one seed coat is present it is termed the spermoderm. If two 

are present, the outer one is called the testa, and the inner one the 

tegmen. The testa, or outer seed shell, differs greatly in form and 

texture. If thick and hard, it is crustaceous; if smooth and glossy, 

it is polished; if roughened, it may be pitted, furrowed, hairy, reticu- 

late, etc. 

The testa may often present outgrowths or seed appendages whose 

functions are to make the seeds buoyant, whereby they may be dis-

214 


seminated by wind currents. Examples of these are seen in the 

Milkweed, which has a tuft of hairs at one end of the seed called a 

Coma, and in the official Strophanthus, which has a long bristle-like 

appendage attached to one end of the seed and called an awn. The 

wart-like appendage at the hilum or micropyle, as in Castor Oil 

Seed, is called the Caruncle. 

The tegmen or inner coat surrounds the nucellus closely and is 

generally soft and delicate. 

A third integument, or accessory seed covering outside of the testa, 

is occasionally present and is called the Aril. Example: Euonymus 

(succulent). 

When such an integument arises from the dilatation of the micro- 

pyle of the seed, as in the Nutmeg, it is known as an Arillode. 

The Nucellus or Kernel consists of tissue containing albumen, when 

this substance is present, and the embryo. Albumen is the name 

given the nutritive matter stored in the seed. The funiculus or seed 

stalk is usually absent in the official seeds. The scar left by its 

separation 

is called the hilum. When the funiculus is continued 

along the outer seed coat, it is called the raphe. 

MODE OF FORMATION OF DIFFERENT TYPES OF ALBUMEN 

If the egg-cell within the embryo sac segments and grows into the 

embryo and, stretching, fills up the cavity without food material 

laid down around it, it happens that the nutritive material lingers in 

the cells of the nucellus, pressing around the embryo. This is called 

Peris per mic albumen. Seen in the Polygonacea. 

In by far the greater number of Angiosperms, the endosperm nu- 

cleus, after double fertilization, divides and redivides, giving rise to 

numerous nuclei imbedded in the protoplasm of the embryo sac, out- 

side of the developing embryo. Gathering protoplasm about them- 

selves and laying down cell walls they form the endosperm tissue 

outside of the embryo. Into this tissue food is passed constituting 

the Endospermic albumen. 

In the Maranlacea, Piperacea, etc., nutritive material is passed 

into the nucellar cells causing them to swell up, while to one side a 

small patch of endosperm tissue accommodates a moderate amount


of nourishing substance, thus resulting in the formation of abundant 

perisperm and a small reduced endosperm. 

Exalbuminous seeds are those in which the albumen is stored in the 

embryo during the growth of the seed. Such seeds show the fleshy 

embryo taking up all or nearly all the room within the seed coat. 

Examples: Physostigma, Amygdala, etc. 

Albuminous seeds are those in which the nourishment is not stored 

in the embryo until germination takes place. Such seeds show a 

larger nourishing tissue region and a smaller embryo region. 

Exam- 

ples: Nux Vomica, Myristica, Linum, etc. 

Gross Structure of a Monocotyl Seed (With fruit wall attached), 

Indian Corn. The ripened seed of Indian Corn is surrounded by a 

thin, tough pericarp which is firmly adherent to and inseparable from 

the Spermoderm or seed coat. Because of this fact, while in reality 

a fruit called a caryopsis or grain, this structure is sometimes erro- 

neously termed a seed. 

The mature grain of most varieties of Indian Corn is flattened and 

somewhat triangular in outline, the summit being broad and the 

base comparatively narrow. The summit is indented and often 

marked by a small point which represents a vestige of the style. 

The basal or " tip" region marks the part of the grain which was in- 

serted into the cob. Upon it maybe found papery chaff, representing 

parts of the pistillate spikelets. The groove noted on the 

broader surface indicates the position of the embryo. 

Histology of the Indian Corn Seed (With fruit wall attached). If a 

longitudinal section be cut through the lesser diameter of a soaked 

grain, the following histologic characteristics will be observed: 

1. The Pericarp or ripened ovarian wall which, alike with all other 

grains, adheres firmly to the wall of the seed forming a portion of 

the skin of the grain. The pericarp comprises an outer epicarp of 

elongated cells with thin cuticle, a mesocarp of thicker walled cells 

without, becoming thinner within, and a layer of tube cells. 

2. The Spermoderm or seed coat, a single layer of delicate elon- 

gated cells. 

3. The Perisperm, another layer directly underneath the Spermo- 

derm, difficult to distinguish without special treatment, and representing 

the ripened nucellar tissue of the ovule.

2l6 PHARMACEUTICAL BOTANY 

4. The Endosperm or nourishing tissue, consisting of: (a) The 

Aleurone Layer, for the most part a single row of cells, containing 

aleurone grains. Some of the cells may be seen to be divided by 

tangential partitions, (b) Starch Parenchyma, consisting of two 

regions: an outer horny zone composed of cells containing for the 

most part polygonal starch grains and oil droplets; and an inner 

mealy zone of cells with mostly rounded starch grains. 

5. The Embryo, consisting of a single shield-shaped cotyledon 

adjoining the endosperm, the plumule or rudimentary bud at the 

end of the caulicle or rudimentary stem and the radicle or rudimen- 

tary root, with its tip covered by a root cap. 

Continuous with the 

root cap is a root sheath or coleorhiza. The cotyledon or seed leaf 

consists of two parts : the scutellum which lies next to the endosperm, 

and is an organ of absorption; and the sheafing portion which sur- 

rounds and protects the rest of the embryo. 

The embryo contains oil and proteids, but no starch. 

If a similar longitudinal section of a soaked grain be mounted in 

dilute iodine solution, the contents of the aleurone cells will be col- 

ored yellow indicating their proteid nature, while the starch grains 

will take on a blue to violet coloration. The endosperm will be 

observed taking up most of the room within the seed coat. The con- 

tents of its cells are not baled out to the embryo until after germina- 

tion begins. Indian Corn is therefore an albuminous seed. 

A MONOCOTYL SEEDLING 

Germination. When any viable seed is planted in suitable soil, 

and furnished with oxygen and water and a certain degree of heat, 

germination takes place. 

In the presence of moisture, etc., the seed 

swells, the ferments present within the cells of the endosper n then 

change the insoluble proteid, starch, and oil to soluble materials, 

which, in the case of Indian Corn, are absorbed in solution by the 

scutellum which bales this nourishment out to other parts of the 

growing embryo, there to be used in part in constructing new tissues, 

and in part to be consumed fry oxidation or respiration. The process 

of respiration or breathing takes place when the plant takes in oxy- 

gen and gives off carbon dioxide. The oxygen oxidizes the tissues


with an accompanying release of energy, which latter is necessary to 

life and growth. 

The combined pericarp and spermoderm bursts opposite the tip of 

the radicle, and the radicle, piercing through the cotyledonary sheath, 

protrudes. The cleft in the coat lengthens to the point opposite the 

tip of the plumule, which also protrudes after bursting through the 

cotyledonary sheath. The radicle, next, grows downward into the 

soil forming the primary root, and develops upon itself secondary or 

lateral roots, all of which give rise to root-hairs just above their root 

caps. Additional lateral roots emerge above the scutellar region 

which ere long attain the size of the first or primary root. The cau- 

licle, carrying upon its tip the plumule, elongates and forms the stem; 

the leaves of the plumule spread out and turn green to function as 

foliage leaves. The perforated cotyledonary sheath grows out sur- 

rounding both the root and the stem for a portion of their length. By 

this time all or nearly all of the nourishment stored in the endosperm 

has been absorbed and assimilated by the young seedling and the 

coat and scutellum, left behind, gradually decay and disappear. 

The root-hairs absorb nourishment from the soil, the green leaves 

build up carbohydrates, prop-roots make their appearance at the 

first node (joint) above ground, and the seedling grows larger. 

Gross Structure of a Dicotyl Seed, Phaseolus lunatus (Lima Bean). 

The Lima Bean Seed shows a flattened-ovate to somewhat reni- 

form outline. Externally it exhibits a polished seed coat which is 

perforated on its basal side by a minute pore called the micropyle or 

foramen. Just below this pore will be noted the hilum or scar which 

represents the point of detachment from thefuniculus or stalk, which 

connected the seed during its growth with the wall of the fruit. 

Upon soaking the seed in water, it is possible to remove the seed coal 

or spermoderm. This done, the embryo will be exposed. The two 

fleshy cotyledons are first seen. Upon spreading these out, convex 

sides down, the rest of the embryo, consisting of a thin leafy structure 

surrounding a bud and called the plumule, 

the caulicle or rudimen- 

tary stem and in line with the latter, the radicle, or rudimentary 

root, will be seen. 

Histology of the Lima Bean Seed. In transverse sections, the 

following microscopic structures will be evident:

2l8 PHARMACEUTICAL BOTANY 

1. Spermoderm of three regions, viz.: Palisade cells, Column cells, 

and Spongy Parenchyma. The palisade cell layer is composed of 

vertically elongated thick-walled cells which are covered on their 

outer faces by a clear glistening cuticle. The cells are 60 to Son long 

and 12 to 2o/z wide. The column cells, found forming a layer 

directly beneath the palisade zone, are hour-glass-shaped and 25 to 

35/u long by 14 to 35/z wide. 

The spongy parenchyma forms a zone of several layers of thin- 

walled parenchyma cells, the cells of the outer and inner layers being 

considerably smaller than the middle layers. 

2. Embryo, the two cotyledons of which make up the greatest bulk. 

These are composed of an epidermis covering over a region of mesophyll. 

The mesophyll is constituted of moderately thick-walled 

cells which contain ellipsoidal and kidney-shaped starch grains up 

to 65/4 in length. A conspicuous branching cleft will be seen in the 

larger grains. 

In the Lima Bean, the nourishment is stored in the embryo during 

the growth of the seed. It is, therefore, exalbuminous .

CHAPTER VIII 

TAXONOMY 

DIVISION I. THALLOPHYTA 

Plants, the greater number of which, consist of a thallus, a body 

undifferentiated into root, stem or leaf. The group nearest to the 

beginning of the plant kingdom presenting forms showing rudimentary 

structures which are modified through division of labor, dif- 

ferentiation, etc., in higher groups. 

SUBDIVISION I. PROTOPHYTA (SCHIZOPHYTA) 

A large assemblage of " fission plants" comprising the bacteria 

and blue-green algae. In the simplest types no nucleus is present, 

but as we arise in scale through the bacteria and blue-green algae, 

there is to be observed an open granular, gradually growing to a 

crescentic, chromatin mass that may be called a nucleus. A common 

method of asexual reproduction is possessed by these plants whereby 

the cell cleaves or splits into two parts, each of which then becomes 

a separate and independent organism. 

I. SCHIZOMYCETES BACTERIA 

Bacteria are minute, unicellular, colorless, rarely weakly red or 

green colored, non-nucleate vegetable organisms destitute of chloro- 

phyll. They serve as agents of decay and fermentation and are 

frequently employed in industrial processes. According to the various 

phenomena they produce, they may be classified as follows (a) 

Zymogens producing fermentation; (b) Aerogens producing gas; (c) 

Photogens producing light; (d) Chromogens producing color; (e) 

Saprogens, producing putrefaction ; (/) Pathogens, producing disease. 

Physical Appearance of Bacterial Colonies and Individual Forms. 

Because of their minute size a space the size of a pinhead may 

219


hold eight billion of them the student commences his study of 

bacterial growths in colonies or cultures, each kind possessing 

characteristics by which they may be distinguished and differentiated. 

The individuals in the colony, depending upon the kind of bac- 

teria under examination, may be globular, rod-shaped, or spiral. 

Bacteria are classed according to form into the following families 

and genera. 

V //" 

O 4* 

FIG. 1 06. Types of micrococci. (After Williams.) 

Family I. Coccaceae. Cells in their free condition globular, be- 

coming but slightly elongated before division. Cell-division in one, 

two or three directions of space. 

A. Cells without Flagella. 

1. Division only in one direction of space forming an aggregation 

resembling a chain of beads Streptococcus. 

2. Division in two directions of space forming an aggregation 

resembling a cluster of grapes Staphylococcus. 

FIG. 107. Types of bacilli. (After Williams.) 

3. Division in three directions of space forming a package-shaped 

or cubical aggregation Sarcina. 

B. Cells with Flagella. 

1. Division in two directions of space Plajiococcus. 

2. Division in three directions of space Planosarcina. 

Family II. Bacteriaceae. Cells longer than broad, generally two 

to six times, straight or only with an angular bend, never curved or 

spiral, division only at right angles to axis or rod; with or without 

flagella and endospores. 

i. Flagella and endospores absent Bacterium.

2. Flagella and endospores present Bacillus. 

TAXONOMY 221 

Family III. Spirillaceae. Cells curved or spirally bent, generally 

motile through polar flagella. 

1. Cells stiff, not flexile. 

(a) Cells without flagella Spirosoma. 

(b) Cells with one, very rarely with two polar flagella Micro- 

spira. 

(c) Cells with a bundle of polar flagella Spirillum. 

2. Cells flexile, spiral very close Spirochaeta. 

Family IV. Mycobacteriaceae. Cells short or long cylindrical 

or clavate-cuneate in form, without a sheath surrounding the 

chains of individuals, without endospores, with true dichotomous 

branching. 

FIG. 108. Types of spirilla. (After Williams.) 

A. In cultures possessing the characters of true bacteria. Growth 

on solid media smooth, flat, spreading. Rod with swollen ends, 

or cuneate or clavate forms Corynebacterium. 

B. Cultures on solid media raised, folded or warty. Generally 

short slender rods, rarely short branched. Take the tubercle stain 

Mycobacterium. 

Family V. Chalamydobacteriacese. Thread-like, composed of 

individual cells, surrounded by a sheath. Simple or with true 

branching. Ordinary vegetative growth by division in only one 

direction of space, i.e., at right angles to the longer axis. 

A. Cell contents without sulphur granules, 

i. Filaments unbranched. 

(a) Cell-division only in one direction of space. 

(b) Cell-division in gonidial formation in three directions 

of space Streptothrix. 

*Marine forms with cells ^surrounded by a very delicate 

hardly discernible sheath Phragmidiothrix.


**Fresh-watej; forms with easily discernible sheath 

Crenothrix. 

2. Filaments branched. 

B. Cell contents with sulphur granules Thiothrix. 

Family VI. Beggiatoaceae. Thread-like, without a capsule, but 

with an undulating membrane. Cell contents show sulphur granules. 

A. Threads apparently not septated, septa only faintly visible 

with iodine staining. Colorless or faintly rose-colored Beggiatoa. 

Sporulation. A large number of bacteria possess the power of 

developing into a resting stage by a process known as sporulation 

or spore formation. Sporulation is regarded as a method of resisting 

unfavorable environment. This is illustrated by the anthrax 

bacilli which are readily killed in twenty minutes by a 10 per cent, 

solution of carbolic acid, and able, when in the spore condition, to 

resist the same disinfectant for a long period in a concentration of 

50 per cent. And, while the vegetative forms show little more 

resistance against moist heat than the vegetative form of other 

bacteria, the spores will withstand the action of live steam for as long 

as ten to twelve minutes or more. 

Whenever the .spores are brought into favorable condition for 

bacterial growth, as to temperature, moisture and nutrition, they 

return to the vegetative form and then are capable of multiplication 

by fission in the ordinary way. 

Reproduction. Bacteria multiply and reproduce themselves by 

cleavage or fission. A young individual increases in size up to the 

limits of the adult form, when by simple cleavage at right angles to 

the long axis, the cell divides into two individuals. 

Morphology Due to Cleavage. According to limitations imposed 

by cleavate directors, the cocci assume a chain appearance, or a 

grape-like appearance, or an arrangement in packets or cubes having 

three diameters. This gives rise to the 

Staphylococcus (plural, staphylococci), from a Greek word referring 

to the shape of a bunch of grapes. 

Streptococcus (plural, streptococci), from a Greek word meaning 

chain-shaped. 

Sarcina, package-shaped or cubical.

TAXONOMY 223 

Form of Cell Groups after Cleavage. The individual bacteria 

after cleavage may separate, or cohere. The amount of cohesion, 

together with the plane of cleavage, determines the various forms 

of the cell groups. Thus, among the cocci, diplo- or double forms 

may result giving rise to distinguishing morphological characteristics. 

Similarly among the bacilli characteristic forms result 

as single individuals and others which form chains of various 

lengths. 

Rapidity of Growth and Multiplication. 

The rapidity with which 

bacteria grow and multiply is dependent upon species and environ- 

ment. The rapidity of the growth is surprising. Under favorable 

conditions they may elongate and divide every twenty or thirty 

minutes. If they should continue to reproduce at this rate for 

twenty-four hours a single individual would have 17 million descendants. 

If each of these -should continue to grow at the same 

rate, each would have in twenty-four hours more, 17 

million off- 

spring, and then the numbers would develop beyond conception. 

However, such multiplication is not possible under natural or even 

artificial conditions, both on account of lack of nutritive material 

and because of the inhibition of the growth of the bacteria by their 

own products. If they did multiply at this rate in a few days there 

would be no room in the world but bacteria. 

Chemical Composition of Bacteria. The quantitative chemical 

composition of bacteria is subject to wide variations, dependent 

upon the nutritive materials furnished them. About 80 'to 85 

substances constitute 

per cent, of the bacterial body is water; proteid 

residue. When these are ex- 

about 50 to 80 per cent, of the dry 

tracted, there remain fats, in some cases wax, 

in some bacteria 

traces of cellulose appear, and the remainder consists of i to 2 per 

cent. ash. 

The proteids consist partly of nucleo-proteids, globulins, and 

protein substances differing materially from ordinary proteids. 

Toxic substances known as endotoxins to distinguish them from 

bacterial poisons secreted by certain bacteria during the process of 

growth, also occur.

224 


SOME BACTERIA PRODUCING DISEASES IN MAN OR THE 

LOWER ANIMALS 

Organism 

Disease 

Staphylococcus pyogenes aureus : Boils, abscesses, carbuncles 

Streptococcus erysipelatis Erysipelas 

Micrococcus meningitidis Cerebrospinal meningitis 

Micrococcus gonorrhoea .'.... Gonorrhoea 

Micrococcus melitensis . 

: Malta fever 

Micrococcus catarrhalis Catarrh 

Bacillus anthracis Anthrax 

Bacterium diphtheria? Diphtheria 

Bacillus typhosus Typhoid fever 

Bacterium influenza? Influenza 

Bacillus tetani Tetanus 

Bacillus leprae -r Leprosy 

" 

Blackleg" of cattle 

Bacillus chauvei 

Bacillus aerogenes capsulatus Emphysernatous gangrene 

Bacterium tuberculosis Tuberculosis 

Bacterium mallei Glanders 

Streptococcus pneumonia) (Diplococcus pneumonia}) Pneumonia (croupous or 

Spirillum cholera? asiatica? 

fibrinous pneumonia) 

Cholera 

Spirillum obermeieri Relapsing fever 

Streptothrix (Actinomyces) bo vis Actinomycosis in cattle 

Actinomyces Myricarum 

Some Bacteria Producing Diseases in Plants 

Tubercles upon and lesions 

within Myrica and Comp- 

tonia 

Bacterium tumefaciens Crown gall 

Bacterium savastanoi Olive knot 

Bacillus amylovorus 

Pear blight 

Pseudomonas juglandis 

Walnut blight 

Bacillus Solanacearum Wilt of Solanacea? 

Bacillus tracheiphilus 

Wilt of Cucurbits 

Pseudomonas Steward Wilt of Sweet Corn 

Mounting and Staining of Bacteria. The mounting and staining 

of bacteria may be accomplished as follows: 

1. Take the square or round cover slip, which has been previously 

cleaned, out of the alcohol pot. Dry it between filter paper. 

2. Hold it in the bacteriolgic forceps, which is so constructed that 

a spring holds the cover slip firmly while an enlargement of the wire

TAXONOMY 225 

handle permits the placing of the forceps on the table while the cul- 

ture material is obtained. 

3. Place several drops of distilled water on the cover slip and add a 

loop full of the organism secured 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. 

5. Hold the open test tube between the thumb and forefinger of the 

left hand. 

6. By means of a previously flamed platinum needle, remove a 

little of the culture from the surface of the culture media. 

7. Replace the cotton plug. 

8. Add the culture media 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 

connection circles being formed by the material affixed to the cover. 

10. Pass the cover glass three times through the Bunsen flame. 

11. Apply the stain, which should remain long enough to stain the 

objects. 

12. Wash off the stain with distilled water. 

13. Dry the cover glass above the flame. 

14. Apply a drop of balsam, turn the cover slip over and drop it on 

to the center of a glass slide previously provided and cleaned for this 

purpose. 

Gram's Method. This is a method of differential bleaching after 

a stain. The cover glass preparations or sections are passed from 

absolute alcohol into Ehrlich's anilin gentian violet, where they 

remain one to three minutes, except tubercle bacilli preparations 

which remain commonly twelve to twenty-four hours. They are 

then placed for one to three minutes (occasionally five minutes) in 

iodine potassium iodide water (iodine crystals i, potass, iodide 2, 

water 300), with or without washing lightly in alcohol. In this 

they remain one to three minutes. They are then placed in absolute 

alcohol until sufficiently bleached, after which they are cleared in 

clove oil and mounted in balsam. 

Certain organisms, when stained by this method give up the stain


and are called "Gram negative; " others retain the color and are 

called " Gram positive." Examples of the latter are B. diphtherias, 

Bacillus anthracis, and Bacillus tetani. 

Stains. One of the most useful bacteriologic stains is ZiehVs 

Carbol Fuchsia, prepared as follows: 

Fuchsin (basic), i. 

Absolute Alcohol, i. 

Carbolic Acid (5 per cent, aqueous solution), 100. 

The fuchsin should be dissolved first in the alcohol and then the 

two fluids mixed. 

Ehrlich's Anilin Water Gentian Violet. Anilin Oil Water, 75 

parts. 

Sat. Sol. Gentian Violet in Alcohol, 25 parts. 

Anilin oil water is made by adding 2 mils anilin to 98 mils distilled 

water; shake violently. Filter through filter paper several 

times. 

Loffler's Methylene-blue. 

Sat. sol. Methylene-blue in Alcohol 30 mils 

Sol. KOH in distilled water (i: 10,000) 

Mix the solutions. 

Stain for "Acid Proof" Bacteria Including 

100 mils 

B. Tuberculosis. 

i. Flood the cover glass with ZiehFs carbol fuchsin and boil over 

the flame for thirty seconds. 

2. Wash and decolorize with a 2 per cent, solution of HC1 in 

80 to 95 per cent, alcohol until the thinner portions of the film 

show no red color. 

3. Wash in water. 

4. Counter stain for contrast with Loffler's Methylene-blue. 

5. Wash and examine. 

Van Ermengem's Flagella Stain. i. Mordant: 

Osmic acid (2 per cent, aqueous solution) .... 50 

Tannin (10 to 25 per cent, in water) 

100 

Four drops of glacial acetic acid may be added to this. 

2. Silver Bath: 

Dissolve 0.25 to 0.5 per cent, nitrate of silver in distilled water in a 

clean bottle.

3. Reducing and Strengthening Bath: 

TAXONOMY 227 

Gallic acid 5 

Tannin 3 

Fused sodium acetate 10 

Distilled water 350 

The flamed cover glass is first covered with the mordant for one- 

half hour, or if in a thermostat at 5oC. for five to ten minutes. 

The mordant is then carefully removed by thorough washing in 

water, alcohol and water. The cover (film side up) is now put into 

the silver bath (a few mils in a clean beaker or watch glass) for a 

few seconds, during which time it is gently agitated. Without 

rinsing it is next put into a few mils of the reducing solution and 

gently agitated until the fluid begins to blacken. It is then washed 

in water and examined. If not stained deeply enough the cover is 

returned to the silver bath. It is finally dried and mounted in 

balsam. All the dishes must be scrupulously clean. The fluids 

must not be contaminated by the fingers nor steel instruments into them. 

by dipping iron or 

Broca's Differential Stain. 

Loffler's Methylene Blue 80 mils 

Ziehl's Carbol Fuchsin 

* 

10 mils 

Mix the solutions. 

This stain differentiates between dead and living bacteria. Dead 

bacteria take on a red coloration and living bacteria a blue color. 

2. CYANOPHYCE.E 

Plants which are sometimes termed blue-green algce. They con- 

tain chlorophyll, a green pigment, and phycocyanin, a blue pigment, 

a combination giving a blue-green aspect to the plants of this group. 

Found everywhere in fresh and salt water and also on damp logs, 

rocks, bark of trees, stone walls, etc. Examples : Oscillatoria, Glce- 

ocapsa, and Nostoc. 

Glceocapsa. This blue-green alga is commonly found on old, 

damp flower pots in greenhouses and on damp rocks and walls


near springs, where it forms slimy masses. Under the microscope 

a mount of Glceocapsa will be seen to consist of isolated protoplasts 

and groups of protoplasts, surrounded by concentric gelatinous 

3-d 

FIG. 109. A, B, C, D, E, Gloeocapsa; F, Oscillatoria showing a dead cell (d) 

which marks a place of separation into segments. (A), Gloeocapsa, parent cell 

composed of central protoplast containing scattered chromatin granules, surrounded 

by cell wall and 3 mucliaginous envelopes; (B), parent cell is shown 

elongated, the protoplast in process oi division to form two daughter protoplasts; 

(C) , daughter protoplasts, each surrounded by two gelatinous envelopes and both 

within the original parent envelopes; (D) the daughter protoplasts shown in C 

have just divided to form granddaughter protoplasts which have later separated, 

each forming envelopes of its own but all four encircled by the parent envelope. 

envelopes. Each protoplast consists of a protoplasmic mass 

which contains blue and green pigments. No definitely organized 

nucleus is apparent but chromatin in the form of granules is scat-

TAXONOMY 229 

tered through the protoplasm. The whole is surrounded by a cell 

wall which undergoes mucilaginous modification producing thus 

the- soft gelatinous envelopes which encircle parent-, daughter-, 

grand-daughter- and even great-grand- 

daughter-cells. 

Oscillatoria. Oscillatoria is a blue-green 

filamentous organism found abundantly on 

the surface of the mud of drains and ditches 

as well a's in ponds where the water is foul. 

The filament is slender and composed of 

compactly arranged disc-shaped 

cells wh'ch 

are all alike, excepting the terminal ones 

which appear rounded off. The filaments 

tend to be agglomerated in thick felts or 

gelatinous masses and each possesses peculiar 

oscillating and forward movements. 

At the time of reproduction the filament 

breaks up transversely into short segments, 

each of which, by fission occurring among 

its cells, grows into a new filament. 

Nostoc. Nostoc occurs on the damp 

ground bordering streams or in slow bodies 

of water as greenish or brownish tough gela- 

tinous masses varying in size from a pea 

to a hen's egg. When one of these masses 

is dissected and examined microscopically, 

it is seen to contain, imbedded in a gela- 

tinous matrix, numerous serpentine fila- 

ments, composed of spherical or elliptical 

cells loosely attached to each other in 

chain-like fashion. Most of the cells are of 

the blue-green vegetative kind but there 

occur at intervals larger cells, often devoid PIG II0 . Nostoc , (h ), a 

of protoplasm which are termed heterocysts. 

heterocyst. 

Frequently the filaments break apart on 

either side of the heterocyst, setting free segments of cells which 

grow into new filaments.

230 


SUBDIVISION II. MYXOMYCETES, OR SLIME MOLDS 

Terrestrial or aquatic organisms, frequently classified as belonging 

to the animal kingdom and found commonly on decaying wood, 

leaves, or humous soil in forests. Their vegetative body consists 

of a naked, multinucleated mass of protoplasm called the plasmodium, 

which has a creeping and rolling amosboid motion, putting out 

and retracting regions of its body called pseudopodia. The size of 

the plasmodium varies from a ten-cent piece to several square feet 

of surface. It is net-like, the net being of irregular dimensions. 

is clear and 

Like the amoeba the outer portion of the plasmodium 

PIG. in. 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, Enerlhema papillatum, F, unripe; G, mature sporangium, 

10/1: H, capillitium, 20/1. (C, D, after nature. A, F, G, H, after Rostafinski; 

B, E, after de Bary in Die nattirlichen Pflanzenfamilien I. i, p. 26.) 

watery and known as the ectoplasm, the inner portion is granular 

and called the endoplasm. Like the amosba and unlike other plants, 

this slimy body engulfs solid food by means of its pseudopodia in- 

stead of admitting it in solution. It is extremely sensitive to light 

being negatively heliotropic, i.e., turning away from the sun's rays. 

At the time of reproduction, the plasmodium creeps to the surface. 

The whole plasmodium then forms one or more fructifications. 

These fructifications vary from cushion-like masses (athallia)

TAXONOMY 231 

to more elevated bodies in which the net-like structure of the plasmodium 

is preserved (plasmodiocarps) to stalked sporangia (spore 

cases). All of the fructifications, however, produce spores. During 

wet weather amoeboid protoplasts (swarm spores] escape from 

the spores, each developing a single cilium and moving actively 

about. In time the cilia disappear and these swarm spores coalesce 

in smaller then larger groups to form a plasmodium. 

SUBDIVISION III. ALG^ 

Low forms of thallophytes of terrestrial and aquatic distribution 

consisting for the most part of single cells or rows of single cells 

joined end to end to form filaments. The higher forms, however, 

possess structures, which might be compared to stems and leaves of 

higher plants although more rudimentary in structure. They 

contain chlorophyll or some other pigment, and so can use the COz 

and H 2O in the same manner as higher plants, e.g., in assimilating 

and providing for their own nutrition. Archegonia are absent in 

this group. 

CLASS I. CHLOROPHYCE.E, THE GREEN ALG.E 

Green algae are unicellular (sometimes motile), filamentous, 

colonial, or sheet-like water plants' characterized by the presence of 

solitary, or numerous chloroplasts in the cells, which compose the 

thallus. These chloroplasts vary considerably in form, being in 

some cases spiral bands, in other star-shaped, in others like a napkin 

ring, and in others granular. In the chloroplasts of most green 

algae are pyrenoids, which consist of a central crystalline portion 

of protein (aleurone-like) surrounded by a starchy envelope of 

variable magnitude. These are called starch centers and the starch 

is frequently in the form of rounded, or angular grains. The nutri- 

tion of these algae is autotrophic. There is a definite nucleus present, 

but in the coenocytic forms the nuclei may be many within the 

confines of the cell wall. The motile cells have one to many cilia, 

as likewise some of the reproductive cells. Reproduction is by cell 

division, the formation of zoospores (motile cells), by zygospores

232 


produced by conjugation, by egg cell and sperm cell union (oospores) 

oogamous reproduction. Green algae live mostly in fresh water. 

Some live in brackish water and a few in the sea. Some are asso- 

ciated with fungi to form lichens. 

1. Order Protococcales or One-celled Green Algae. This order 

contains nearly all of the one-celled green algae excepting the diatoms 

arid desmids. 

Family Pleurococcaceae. Pleurococcus vulgaris is a one-celled 

green alga, millions of which, living together in colonial fashion, constitute 

the so-called "green stain" that is common on the north 

sides of tree trunks, stone walls and fences. Each organism consists 

of a protoplast surrounded by a cell wall of cellulose. The 

protoplast contains a chromatophore, cytoplasm and nucleus. 

Reproduction takes place by the protoplast dividing into two equal 

parts and laying down a cell wall forming two daughter-protoplasts. 

These may again divide to form four granddaughter-protoplasts. 

Still another division may occur as a result of which eight greatgranddaughter-protoplasts 

are formed which frequently adhere to 

one another forming colonies. 

2. Order Volvocales. This order comprises free-swimming 

aquatic forms whose vegetative cells are bi-ciliated, green, more or 

less spherical or compressed. Some of the organisms like Sphcerella 

and Chlamydomonas consist of single cells bearing a pair of cilia, 

while others like Pandorina, Eudorina and Volwx show varying de- 

grees of colony formation. Reproduction sexual or asexual. 

Volvox globator, a typical representative of this order, is found in 

fresh water pools as a tiny, hollow, spherical, green colony about 

/-io to Jô of an- incn in diameter. When examined under the 

microscope (Fig. 112), it is found to consist of hundreds of green, 

more or less spherical cells, united by fine strands of cytoplasm 

(protoplasmic bridges), the whole being enveloped by a gelatinous 

sheath. The peripheral cells are provided with cilia, in order that 

the colony may rotate and roll through the water. In a young 

colony, all of the cells are alike, each consisting of a mucilaginouslike 

cell-wall enclosing cytoplasm, a nucleus, a chloroplast and often 

a red pigment spot. In a mature colony, however, throughout the 

greater part of its existence, two kinds of cells may be discerned:

TAXONOMY 233 

FIG. 112. Fofoo* globator. Mature colony in center (i); sexual cells (20); 

endochrome of primary cell has resolved itself into a cluster of secondary cells 

(ia, a 2 and 5) antherozoids ; (6, 7) bundle ; separated into component antherozoids 

in cavity of primary cell (ia 3 ); breaking of wall of primary cell showing escape 

of antherozoids into cavity of volvox sphere (ia 4 ); egg cells (ib, 6); flask shaped 

germ (egg) cells with large vacuoles in protoplasm (i& 2 , & 2 ); globular egg cell 

prepared to pass into cavity of volvox sphere (6 3 

); antherozoids collected about 

egg cell (3); oospore (4). (Carpenter.)

234 


small, sterile, vegetative cells that do not divide and from 10 to 12 

larger vegetative ones that divide to form new colonies. The latter 

slip inward below the level of the smaller cells and through repeated 

divisions form a number of ciliated cells jointed by cytoplasmic 

threads, which in reality is a minature colony. This then escapes to 

the exterior through the rupturing of the gelatinous wall of the old 

colony. 

During autumn of the year, certain of the ordinary cells undergo 

differentiation, some to form sperm cells, others, egg-cells. When 

about three times the size of the ordinary sterile cells, the sperm cells 

divide repeatedly to form a cluster of elongated secondary cells 

[Fig. 112 (ia, a 2 and 5),] each of which contains an orange colored 

endochrome with a red corpuscle and an elongated beak, bearing a 

pair of flagella (lash-like processes). The cluster in time separates 

into motile antherozoids [Fig. 112 (6, 7)] which finally escape into 

the cavity of the volvox sphere through rupture of the investing 

wall. The flask-shaped egg cells (ib, b) increase greatly in size with- 

out dividing. Each shows vacuoles, then becomes filled with a 

dark green pigment, becomes spherical and acquires a gelatinous 

envelope. It then passes into the cavity of the sphere where it is 

surrounded by numerous antherozoids (3) and is finally fertilized. 

The product of this fertilization is an oospore (4) which ere long 

becomes covered with an internal smooth membrane and a thicker 

external spinose coat. The chlorophyll within then disappears and 

starch and a reddish- or orange-colored oil make their appearance. 

Up to 40 of these oospores have been observed in a single volvox 

sphere. Not long after the formation of these oospores the whole 

fall to the bottom of the 

parent colony breaks up and the oospores 

pool to pass the winter season. As early as February each oospore 

germinates to form another volvox colony, which repeats the life 

cycle described. 

3. Order Confervales. In this order are included a variety of 

green filamentous and membranous forms some of which show sexual 

reproduction. 

Family Ulothricaceae. Ulothrix zonata, a typical representative 

of this family, is a filamentous organism found growing on stones 

around ponds, on rocks along the shores of lakes, in slow-moving

TAXONOMY 235 

streams, etc. Each filament is unbranched and consists of a row of 

short cells, one of the terminal cells, called the rhizoid cell, being 

flongated and serving as an attachment structure. Each cell con- 

sists of a cell wall of cellulose enclosing cytoplasm, a nucleus and a 

wide band-shaped green chromatophore, more or less cylindrical in 

shape. The chromatophore lies next to the cell wall and contains 

pyrenoids or starch-forming centers. The filament grows in length 

by the fission of its various component cells. After attaining a 

certain size it reproduces either asexually or sexually. Asexual 

reproduction takes place by certain cells becoming altered in their 

protoplasmic contents, through division, to form rounded or pearshaped 

zoospores. Each zoospore contains a red pigment spot and 

FIG. 113. Vaucheria terrestris. anth, antheridium (empty) ; o, oogonia. (Gager.) 

bears four cilia (protoplasmic outgrowths). The zoospores escape 

into the water by lateral openings in the walls of cells containing 

them. They swim rapidly about, propelled by their cilia, and ere 

long attach themselves to various objects and grow into Ulothrix 

filaments. The sexual method of reproduction is effected through 

the production of many gametes, in cells of the filament, which re- 

semble the zoospores in shape but differ from them in being smaller 

and possessing but two cilia. These escape into the water, and, 

after swimming about for a short time come together in pairs and 

fuse with one another. The product of the fusion of each pair of 

these like gametes is termed a zygospore. The zygospore swims 

about but finally comes to rest, remaining quiescent for a consider- 

able length of time. It then enlarges and its protoplasmic content

236 


divides to form several zoospores which, escaping from the cell, 

swim about for a while and finally, attaching themselves to objects, 

grow into filamentous Ulothrix organisms. 

4. Order Conjugates. To this order belong the desmids and 

pond scums which are distinguished from other green algae by 

presenting no motile stages, in their life histories. fresh-water habit and reproduce by conjugation. 

They are all of 

Family Desmidacese. The desmid family includes a number of 

genera of unicellular as well as filamentous green plants that present 

a variety of shapes. Each unicellular desmid is characterized by 

being composed of two like halves frequently separated by each 

other by a constriction called the isthmus. In each half there is a 

The nucleus is found in the 

chromatophore containing pyrenoids. 

isthmus. Reproduction is accomplished either asexually by fission 

or sexually by conjugation. 

Family Zygnemaceae. This is a family of pond scums including 

the well-known genera, Spirogyra and Zygnema. 

Spirogyra or Brooksilk is a filamentous organism found suspended 

or floating in masses in quiet water. Each filament when examined 

microscopically will be found to consist of more or less elongated 

cylindrical cells arranged end to end, the terminal cells having 

rounded extremities. Each cell has a cell wall of cellulose within 

which is to be found a thin film of ectoplasm. One or more spirally 

shaped chromatophores will be seen directly within this area. Each 

chromatophore contains chlorophyll and a number of pyrenoids. 

In the center of the cell the nucleus is found. Fine strands of 

protoplasm hold it in place and run out to the ectoplasm. 

Under favorable circumstances the cells of Spirogyra increase 

rather rapidly in length. Abnormally long cells are not seen, 

however, because the elongating cells speedily divide, forming two 

daughter-cells. Under the best of conditions, division may occur 

every night. In this way the filaments are rapidly made longer. 

Sooner or later they break and in this way the plant multiplies. 

Spirogyra has also a process of sexual reproduction known as 

conjugation. This process occurs normally from March to June 

and July, but can be induced in the laboratory by allowing the water 

in the vessel in which it is growing to slowly evaporate. Two fila-

TAXONOMY 237 

ments arrange themselves side by side, and the cells lying opposite 

each other undergo internal changes so as to form gametes or sexual 

cells. Each protrudes a process or conjugation tube; these unite 

FIG. 114. Spitogyra sp. A, terminal portion of vegetative filament; B, stages 

of scalariform conjugation; C, preparation for lateral conjugation; D, zygospores 

formed by lateral conjugation. (Gager.) 

and the protoplasm from one cell passes over and coalesces with that 

in the cell opposite. The result of the process is a new cell called a 

zygospore or zygote. This is set free by decay of the walls of the old

2 3 8 PHARMACEUTICAL BOTANY 

cell and falls to the bottom of the water, there to undergo a resting 

stage until favorable conditions for growth arise. 

5. Order Diatomales. Family Diatomaceae. This family com- 

prises several thousand species of unicellular plants called Diatoms 

which are found in fresh, brackish and salt water, forming much of 

the diet of small animals. While unicellular, they frequently are 

united in colonies. They all possess chromatophores containing 

PIG. 115. Two species of Diatoms. To left, Diatoma vulgare; a, side view of 

frustule; b, frustule undergoing division. To right, Grammatophora serpentina: 

a, front and side views of single frustule; b, b, front and end views of divided 

frustule; c, frustule about to undergo division; d, frustule completely divided. 

(After Carpenter.) 

chlorophyll but this green pigment is often obscured by the presence 

also of a brown pigment. 

The most striking peculiarity of the group is the structure of the 

enclosing cell wall. This is in the form of a siliceous case consisting 

of two valves which fit into each other like the halves of a pill box. 

The valves, which are beautifully sculptured, are similar except that 

one is slightly larger than the other so as to fit over it. Diatoms 

vary in form being either circular, linear, elliptical, cylindrical, 

rhomboidal, triangular or fan-shaped, etc. Some are borne on the

TAXONOMY 239 

ends of stalks, while others are held in gelatinous masses. Their 

siliceous skeleta are deposited constantly on the floor of ponds, rivers, 

lakes and seas, often in such abundance as to form Diatomaceous 

earths or Kieselguhrs (Siliceous Earths). Huge geological deposits 

of this material have been found in various parts of the world. The 

most remarkable for extent as well as for the number and beauty of 

the species contained in it is that at Richmond, Virginia. It is in 

many places 25 to 40 feet in depth and extends for many miles. 

PIG. 116. Licmophora flabellata, a diatom with wedge-shaped frustules borne 

on the ends of stalks, producing a fan-like arrangement. (After Carpenter.} 

Many of the diatomaceous earths are useful as absorbent and polish- 

ing powders. The United States Pharmacopoeia IX recognizes, 

under the name of Terra Silicea Purificata (Purified Siliceous Earth), 

a powder consisting of the frustules and fragments of diatoms which 

has been purified by boiling with diluted hydrochloric acid, washed 

and calcined. 

Diatoms exhibit two modes of reproduction, viz., fission and for- 

mation of an auxospore. The more common method is that of

240 


fission but this is peculiar for these plants. The cell-contents within 

the siliceous case separate into two distinct masses and the valves 

separate slightly from each other. As the two daughter-masses 

become more and more developed, the valves of the parent-cell are 

pushed more widely apart. Each of the two massses secretes for 

itself a new valve on the side opposite to the original valve. When 

the process is completed the girdle of the parent-diatom separates 

FIG. 117. Fossil diatoms: a, a, a, Gaillonella procera and G. granulata; b, b, 

Surirella plicata; c, Surirella craticula; d, d, d, Gaillonella (Melosira) biseriata 

(side view) ; e, Gomphonema gracile; f, Cocconema fusidium; g, Tabellaria vulgaris; 

h, Pinnularia dactylus; i, Pinnularia nobilis; k, Surirella caledonica; I, Synedra 

ulna. (After Carpenter.) 

and the two daughter-diatoms thus become independent plants. 

Each of these possesses one of the parent valves and a second, 

which it has formed itself more or less parallel to the first. 

In a number of species, repeated fission results in the formation 

of succeedingly smaller and weaker individuals. This process, 

however, goes on only for a certain number of generations until 

the decrease of size has reached a limit for the species, when the

TAXONOMY 241 

plant is rejuvenated by the formation of an auxospore. This may 

be formed with or without the conjugation of two parent-protoplasts. 

In either case the auxospore resulting undergoes a resting stage after 

which it develops new valves. The newly formed diatom is then 

several times the size of the individual or individuals which con- 

tributed to its formation and is endowed with renewed vigor for 

growth and division. 

6. Order Siphonales (Siphon Alga). This group is characterized 

by the peculiarity that the organisms constituting it possess proto- 

of nuclei within a common filament or 

plasm containing myriads 

cell cavity not segmented by cell walls. The term ccenocyte has 

been given to such structures which consist of a many-nucleated mass 

of protoplasm surrounded by a cell wall. Some of the siphon algae 

reproduce by zoospore formation, others by conjugation 

as well as 

zoospore formation while Vaucheria, the green felt, stands out alone 

in reproducing both by the formation of a single zoospore and 

by the production also of ob'gonia and antheridia with resultant 

fertilization. (Fig. 113). 

7. Order Chorales (The Stoneworts). Family Characeae. The 

highest group of algae, possessing forms which are differentiated into 

stems, leaves and rhizoids. 

Char a, a type of this family, is a submerged fresh-water plant 

which fastens itself to the muddy bottom of ponds, ditches and slow 

streams by means of slender filaments called rhizoids. From these 

there arises a many noded (jointed) stem which bears whorls of 

slender green leaves at its nodes. Branches are also found issuing 

from some of the nodes which duplicate in appearance the main 

stem. Reproduction is either asexual or sexual. Asexual reproduction 

is accomplished by means of tuber-like bodies borne on 

branches which form rhizoids on their 

submerged parts or by special 

lower nodes and later become separated from the parent plant. 

Sexual reproduction is effected through the formation of oogonia 

(female sex organs) and antheridia (male sex organs). These in 

some species are borne on the same plant; in others, on different 

plants. In all cases the sexual organs are produced at the nodes. 

The oogonium develops within itself a large ovum or egg. The 

antheridium produces within its wall numerous motile sperms. 

16

242 


Upon the maturation of the antheridium the sperms are liberated 

into the water, and, propelled by their cilia, find their way to the 

oogonia which they enter, the one best adapted fusing with the egg 

in each case and fertilizing it. The resultant cell is cafled the 

oospore. 

This undergoes a resting stage and later germinates as a 

proembryo. The proembryo consists of a simple filament and a 

long rhizoidal cell. From this proembryo, the adult stern arises as a 

side branch. 

CLASS II. PHAEOPHYCEÊ, THE BROWN ALGÊ 

Mostly marine forms showing great diversity in the form of their 

vegetative bodies. They occur for the most part in salt water between 

the high and low tide marks. Their bodies are usually fixed 

to some support in the water by means of a holdfast, and are often 

highly differentiated both as to form and tissues. Some reach 

hundreds of feet in length as, for example, Macrocystis which grows 

in the Pacific Ocean off the coast of California. They all contain 

the brown pigment called phycophcein and the green pigment, chloro- 

phyll both of which are present in their chromatophores. A yellowish 

pigment called phycoxanthin has also been isolated from some 

of the species. Many of the kelps and rockweeds belonging to this 

class have long been sources of iodine, potash and sodium. 

A Filamentous Brown Alga, Ectocarpus Siliculosus. Ecto- 

carpus occurs as tufts of branching filaments, 

each of which is 

many-celled. These tufts are found on eelgrass or other algae as 

well as attached to pilings of wharves in salt water. It is a striking 

illustration of the simplest form of brown algae and serves to show 

the beginning of a more complex form of reproduction than that 

observed in the forms studied up to this time. On examination of a 

filament we find it to consist of many cells joined end to end. A 

single cell has a cell wall of cellulose. Just within the cell wall 

there is a layer of protoplasm. Going toward the center we find an 

irregular chromatophore containing a brown pigment called phycophaein. 

From certain cells of the filament spherical sporangia (spore 

cases) arise, which are unicellular. They contain numerous biciliate 

zoospores, which escape into the sea water, move about and later

TAXONOMY 243 

develop into new Ectocarpus plants. Along the filaments several 

branches will be seen. Some of these have undergone division into 

several cells and these again into still smaller cells until many-celled 

chambers have resulted, which are called plurilocular sporangia. 

FIG. 118. End of large branch of Fucus vesiculosus (natural size); e, receptacle; 

b, air bladder. 

Each cell of a plurilocular sporangium contains a gamete or sexual 

cell, which resembles in many details a zoospore. When the sporangium 

matures these gametes are discharged into the salt water. 

They fuse together in pairs and form zygospores. Each zygospore

244 


undergoes a resting stage and upon the advent of favorable conditions 

develops into a new Ectocarpus filament. 

Fucus Vesiculosus (The Bladder Wrack). This form, a brown 

alga, occurs as a flat thallus, which forks repeatedly, a type of 

T 

FIG. 119. Fucus vesiculosus. Receptacle cut transversely, c, conceptacle, 

T, cellular thickness. (Magnified.) 

branching called dichotomous. It grows near the surface of sea 

water, attached to rocks or to mussels along banks by means of a 

basal disc-shaped holdfast. In the upper branches of the thallus are 

FIG. 120. Fucus vesiculosus. Section of a male conceptacle lined with branched 

paraphyses which bear the antheridia (highly magnified). 

to be found air bladders which are more or less spherical and usually 

in pairs. The tips of old branches become swollen and are termed 

receptacles. They are dotted over with minute cavities called con-

ceptacles. 

TAXONOMY 245 

Within these conceptacles the antheridia, or male sexual 

organs, and the archegonia, or female sexual ogans, are produced. 

The conceptacles also contain numerous branching filaments called 

FIG. 121. Fucus vesiculosus. Section of a female conceptacle. o, osteole 

by which the eggs escape. The oval shaped dark objects represent oogonia in 

different stages of maturity (highly magnified). 

paraphyses, which arise from the cells lining the cavities. The an- 

theridia are found as outgrowths of these paraphyses and produce 

sperms or male sexual cells. The oogonium is a large, globular, 

FIG. 122. Fucus vesiculosus. Fertilization. A, egg approached by biciliated 

sperms; B, sperms attached to egg and surrounding it prior to fertilization 

(highly magnified). 

stalked cell and produces eight eggs, each of which is a fema*le sexual 

cell. The eggs and sperm escape into the sea water. The eggs 

float and are surrounded by myriads of sperms. One sperm, only,

246 


gains an entrance, after which its nucleus fuses with that of the egg 

to form an oospore. The oospore at once develops into a new Fucus 

plant. 

CLASS III. RHODOPHYCEÊ, THE RED 

A greatly diversified group comprising the majority of marine algae 

but represented also by some fresh-water forms. The marine red 

algae are generally found at or just beyond the low water mark. 

Their vegetative bodies vary from simple branching filaments 

through all gradations to forms differentiated into branching stems, 

holdfasts and leaves. It has been observed that many of the higher 

types are composed of numerous filaments which are arranged so 

closely and connected so intimately by protoplasmic processes 

as to resemble the tissues of plants higher up. Their color may be 

red, purple, violet, or reddish-brown or even green and is due to the 

presence of phycoerythrin, a red pigment, which is found in the 

chromatophores with but frequently masking the chlorophyll. 

Chondrus crispus and Gigartina mamillosa yield the official drug 

Cbondrus, Irish Moss or Carragheen. Both are purplish-red in 

color. Each consists of a dichotomously branched thallus the 

lower portion of which is differentiated as a stipe or stalk; the basal 

portion of which, called the holdfast, clings to the rock. The upper 

part is several times forked and its terminal segments appear notched 

or bilobed. Scattered here and there over the segments of the 

thallus will be noted sporangia which, when mature, contain tetra- 

s pores. In Chondrus crispus the sporangia are elliptical and em- 

bedded in the thallus near its surface, whereas in Gigartina they are 

ovate and project outward from the surface of the segments. Upon 

the ripening of these structures the spores are discharged into the 

sea water. These sooner or later germinate into new Chondrus or 

Gogartina organisms. 

The dried mucilaginous 

substance extracted -from Gracilaria 

lichenoides, Gelidium and Gloiopeltis and other species of red algae 

growing in the sea along the eastern coast of Asia constitutes the 

drug Agar, a most valuable ingredient in culture media as well as 

a laxative.

TAXONOMY 247 

Rhodymenia palmata or Irish Dulse is a purplish-red, flat, membranous, 

palmately cleft or dichotomous red alga growing on the 

tissues of other algae along northern shores of the Atlantic between 

the low- and high-tide marks. 

SUBDIVISION IV FUNGI 

This great assemblage of thallophytes is characterized by 

the total 

absence of chlorophyll and so its members possess no independent 

power of manufacturing food materials such as starches, sugars, etc., 

from CC>2 and H 2O. Consequently they are either parasites, depend- 

ing for their nourishment upon other living plants or animals, called 

hosts] or saprophytes, depending upon decaying animal or vegetable 

matter in solution. Some forms are able to live either as saprophytes 

or parasites while others are restricted to either the parasitic or 

saprophytic habit. The vegetative body of a fungus 

mycelium. It consists of interlacing and branching filaments 

called hypha, which ramify through decaying matter or invade the 

tissues of living organisms and derive nourishment therefrom. In 

is known as a 

the cases of parasites, the absorbing connections which are more or 

less specialized and definite are called haustoria. In the higher forms 

the hyphae become consolidated into false tissues, and assume definite 

shapes according to the species. Of this character are the fructi- 

fying organs which constitute the above ground parts of Puff Balls, 

Cup Fungi, Mushrooms, etc. There are four classes of Fungi, viz. : 

Phycomycetes, Ascomycetes, Basidiomycetes and Fungi Imperfecti. 

CLASS I. PHYCOMYCETES, OR ALGA-LIKE FUNGI 

The Phycomycetes represent a small group of fungi showing close 

affinity with the green algae. 

Their mycelium is composed of cceno- 

cytic hyphae, which suggests a close relation with the Siphonales 

group of green algae. Their sexual organs are likewise similar in 

structure. Transverse septa appear upon the formation of repro- 

ductive organs separating these structures from the vegetative 

hyphae.

248 


SUB-CLASS A. ZYYGOMYCETES 

(Sexual apparatus shows isogamy) 

Order i. Mucorales, the black molds, mostly saphrophytic. Ex- 

amples: Mucor Mucedo, Rhizopus nigricans, Thamnidium,Pilobolus. 

Rhizopus nigricans (Mucor stolonifer), commonly known as 

" Black Mold " or " Black Bread Mold," is frequently found on bread, 

jellies, syrups, acetic pharmaceutical extracts and other substrata, 

where it forms a dense thready mycelium bearing numerous black 

tiny spore cases. The source of this mold is the spores, which are 

found in the air or water with which the attacked substratum is 

FIG. 1 23 . Black mold (Rhizopus nigricans} . A , older plant ; myc, myc elia spx 

sporangiophore; sp, sporangium; st, stoloniferous hypha produced by A, and 

' 

giving rise at its tip to a new plant, B. 

Greatly enlarged. (Gager.) 

in contact. Each of these, upon germinating, sprouts out and forms 

three kinds of hyphae, vix. : rhizoidal or submerged hyphce, spor- 

angiophores or aerial hyphce and stoloniferous hyphce. The branching 

rhizoidal hyphae penetrate the substratum and secrete a diastatic 

ferment that changes the water insoluble carb6*hydrate ma- 

terials into a soluble sugar which passes into solution and is absorbed 

by their walls. This, upon entering the hyphae, 

is converted into 

protoplasm, and so the mold increases in size. Sporangiophores 

or aerial hyphae . 

arise vertically or obliquely from a bulged-out 

common base of the rhizoidal hyphae. Each of these when mature

TAXONOMY 249 

bears upon its summit a spheroidal sporangium containing numerous 

small brownish multinucleate spores called endospores. The wall 

of the sporangium is beset with asperites of calcium oxalate. Springing 

from the base of the sporangiophores or aerial hyphae one or more 

stoloniferous hyphae traverse a portion of the surface of the sub- 

stratum and their tips, coming in contact with the substratum, 

swell up forming an adhesive organ or appressorium which branches 

out below into a cluster of spreading submerged hyphae and above 

into several aerial hyphae bearing sporangia. This method of 

growth proceeds 

until the entire surface of the nutritive medium is 

covered with a dense fluffy mycelium. 

D ;U 

FIG. 124. Rhizopus nigricans. A, Young sporangium, showing columella 

within; B, older sporangium, with the 'wall removed, showing ripe spores covering 

the columella; C, D, views of the collapsed columella after dissemination of the 

spores. (Gager.) 

Rhizopus reproduces by two methods. The most common one is 

that of internal cell formation. In this asexual method a transverse 

wall is laid down in the sporangiophore near its tip. The terminal 

cell ftius formed swells up, becoming globular in shape and its protoplasmic 

ontents become changed to form numerous spores within 

the wall of the sporangium or enlarged terminal cell of the sporangio- 

phore. The partition wall, separating the lumen of the sporangium 

from that of the sporangiophore, bulges into the sporangium as a 

dome-shaped structure, which is termed the columella. Upon the 

ripening of the spores the wall of the spore case bursts, liberating 

them. These, falling upon moist nutrient substrata, germinate and 

ultimately form new Rhizopus plants. Under certain conditions 

Rhizopus reproduces sexually. Thicker and shorter club-shaped 

hyphae arise on opposite branches of the mycelium. A partition 

If

250 


wall is laid down in each of these a short distance from its tip and 

the contents of each end-cell then becomes a gamete or sexual cell. 

The apical cells of the tips of opposite hyphae then meet, a solution 

of the cell walls at the point of contact takes place and the gametes 

of both end-cells fuse to form a zygospore. This enlarges and devel- 

ops a highly resistant wall. After a period of rest, upon coming 

into contact with a nutrient medium, it germinates into an elongated 

sporangiophore which develops a sporangium at its summit. 

FIG. 125. Mucor mucedo, showing mycelium and sporangiophores. (Palladfy.) 

Mucor mucedo, another closely allied species, found growing on 

old nuts, fleshy fruits, bread and horse manure, resembles Rhizopus 

nigricans in many respects but differs from it by the formation of 

sporangiophores singly instead of in clusters. 

Thamnidium differs from Rhizopus and Mucor in the development 

of two kinds of sporangia, microsporangia and megasporangia. The 

sporangiophore produces a terminal large megasporangium possessing 

a columella and whorls of side branches which bear smaller 

microsporangia in which the columella is frequently wanting.

TAXONOMY 251 

SUB-CLASS B. OOMYCETES 

(Sexual apparatus heterogamous) 

Order i. Chytridiales. Example: Synchytrium, a form para- 

sitic on seed plants and forming blister-like swellings. 

Order 2. Saprolegniales. Water molds which attack fishes, 

frogs, water insects, and decaying plants and animals. Example: 

Saprolegnia. 

Order 3. Peronosporales. Mildews, destructive parasites, liv- 

ing in the tissues of their hosts and effecting pathologic changes. 

Example: Albugo, the blister blight, a white rust attacking members 

of the Cruciferce and Phytophthora, producing potato rot. 

CLASS II. ASCOMYCETES, THE SAC FUNGI 

Mycelium composed of septate filaments and life history characterized 

by the appearance of a sac called an ascus in which ascospores 

are formed. The largest class of fungi. 

Order i. Protoascales. Plants with asci borne free or at the 

ends of hyphae, definite fruiting bodies being absent. Each ascus 

usually develops four ascospores. To this order belong Exoascus, 

which is responsible for the abnormal development of tufted masses 

of branches on a number of trees and shrubs, and the yeasts (Sac- 

charomycetaceae) many of which produce fermentation. 

Yeasts are unicellular plants of spheroidal, oval, elliptical, pyriform 

or sausage shape which reproduce by budding. They occur 

either in the wild or cultivated condition and are generally found 

capable of breaking down some form of sugar into alcohol and carbon 

dioxide. 

According to the kind or kinds of sugar fermented Hansen in 

1888 classified the yeasts as follows: 

1. Species which ferment dextrose, maltose and saccharose: 

Saccharomyces cerevisice I, S. ellipsoideus I, S. ellipsoideus IT, 

S. pastorianus I, S. pastorianus II, S. pastorianus III. 

2. Species which ferment dextrose and saccharose, but not maj- 

tose: Saccharomyces marxianus, S. exiguus, S. saturanus, S. Ludwigii. 

3. Species which ferment dextrose, but neither saccharose nor 

maltose: Saccharomyces mail Duclauxii.

252 


4. Species which ferment dextrose and maltose, but not saccha- 

rose: Saccharomyces n. sp. obtained from the stomach of the honey- 

bee. 

5. Species which ferment neither maltose, dextrose nor saccharose: 

Saccharomyces anomalus var. belgicus, S. farinosus, S. hyalosporus, 

S. membranifaciens. 

The two most important yeasts in the fermentation industries are 

Saccharomyces ceremsice and Saccharomyces ellipsoideus. 

Saccharomyces cerevisia, commonly called Brewer's Yeast, is a 

cultivated species with many strains. It is used extensively in the 

brewing and baking industries and in recent years has met with 

considerable esteem by the medical profession in the treatment of 

certain skin diseases. 

When examined under the microscope it is found to be somewhat 

spheroidal to ellipsoidal in outline, 8 to i2/x long, and 8 to lOyi* broad. 

It consists of an outer cell wall of fungous cellulose enclosing cyto- 

plasm and a nucleus, the latter invisible without special staining. 

The cytoplasm is differentiated into a clear outer membrane lying 

directly within the cell wall and termed the ectoplasm and an inner 

granular region, the endoplasm. In the young condition of the 

yeast cell numerous glycogen vacuoles are found scattered more or 

less uniformly throughout this region but as the cell matures these 

coalesce, until, in a very old cell, a huge glycogen vacuole may be seen 

occupying most of the interior, with the cytoplasm and nucleus 

pushed up against the cell wall and forming there a very narrow 

layer. 

Yeast plants grow in dilute saccharine solutions containing dis- 

solved nitrogenous substances such as beerwort, Pasteur's solution, 

grape juice, etc. Here they are constantly wasting away and as 

constantly being built up. The question may well arise: "How do 

they obtain the material necessary for growth and . repair?" The 

answer, in a ' 

general way, is not difficult. The fluid in which they 

live is a solution of sugars and pf nitrogenous and other matters. 

The cell walls are readily permeable. Food substances diffuse 

through it into the cell, and by a series of changes (which, indeed, 

it is no easy matter to understand) are converted into new living 

substance. The waste products likewise diffuse readily outward.

TAXONOMY 253 

This method of nutrition is called saprophy tic, and the yeast plant 

is said to be a saprophyte. 

A striking fact must be briefly mentioned in connection with the 

metabolism of yeast. Many organisms exercise a profound effect 

on the medium in which they live. Yeast causes a wholesale destruc- 

tion of sugar in the surrounding fluid. One of the decomposition 

FIG. 126. Yeast, Saccharomyces cerevisice, the variety known as brewers' bottom 

yeast; a, spore formation; b, elongated cells. (After Schneider, Pharmaceu- 

tical Bacteriology.) 

products of sugar is alcohol. The alcohol of commerce is produced 

by the activity of this plant. 

Saccharomyces has its times of danger and stress when the cells 

perish in great numbers from cold, starvation, poisons, etc. If not 

too suddenly exposed, however, they are able to meet adverse con- 

ditions by eliminating most of their water, suspending physological

254 


processes, and becoming dormant. Sometimes they enter the rest- 

ing condition after a process of division, when each cell divides into 

four parts, each of which becomes nearly dry and is surrounded by a 

thick wall. Such cells are called ascospores, and their production 

serves both as a method of multiplying the plant and of tiding over 

adverse conditions. They can survive for a long time without 

food or water, and can endure higher temperatures than the active 

cells and almost any degree of cold. 

FIG. 127. Saccharomyces ceremsia. The form or variety known as brewers' top 

yeast. (Oberhefe.) 

The dried cells and spores float in the air as dust and so accomplish 

a dispersal of tire organism. Doubtless most of them never again 

meet suitable environment and so sooner or later perish. But some 

will fall into favorable conditions and be able to multiply enormously 

again, and so the species is continued. 

The general method of reproduction in Saccharomyces is that of 

gemmation or budding. A small protuberance of the cell wall com-

TAXONOMY 255 

mences to form on the parent-cell. This grows larger and a portion 

of the cytoplasm and nuclear material pass into it. Eventually a 

daughter-bud, which may assume the size of the parent-cell, is 

formed. This generally adheres to the parent-cell and produces one 

or more granddaughter-buds which in turn may produce great-grand- 

daughter-buds before separation from the parent-cell takes place. 

There are two varieties of brewer's yeast, viz.: Top yeasts and 

Bottom yeasts. Top yeasts grow on or near the surface of the liquid 

PIG. 128. Saccharomyces ellipsoideus. A common yeast found on grapes, 

jams, jellies, etc. Budding process is shown in many of the cells as also the vacuoles. 

(Schneider, Pharmaceutical Bacteriology.} 

and produce rapid fermentation at summer temperatures causing 

great quantities of carbon dioxide to arise to the surface and thus 

forming the froth which is characteristic of ale, stout and porter. 

Bottom yeasts grow at about 4C. at or near the bottom of the 

vat. They are used in the manufacture of lager beers. 

Compressed yeast (Cerevisiae Fermentum Compressum) N. F. 

consists of the moist, living cells of Saccharomyces cerevisia or of other 

species of Saccharomyces, combined with a starchy or absorbent 

base.

256 


Saccharomyces ellipsoideus is a wild species, several varieties of 

which are found growing on grapes especially in districts where wine 

is produced. It is termed the true wine, yeast to distinguish it 

from other wild species found in grape juice, like S. apiculatus and 

S. membranifaciens which exert a deleterious effect in wine production. 

Its cells are ellipsoidal, 6/x long, occurring singly or in rows 

of several generations, which are rather loosely joined. 

Order 2. Pezizales or cup fungi. Examples: Peziza, Lachnea 

and Ascobolus. 

FIG. 129. Saucer-shapes fruit-bodies of Peziza repanda. (Harshberger, from 

Photo by W. H. Walmsley.) 

Parasitic or saprophytic plants, whose vegetative bodies consist of 

a mycelium ramifying through 

the substratum and whose above 

ground fruiting bodies are sessile or stalked, cup or saucer-shaped 

structure termed apothecia (sing, apothecium), in which a fruiting 

membrane (hymenium) lines the concave upper surface. The asci 

are usually eight-spored and separated from each other by filament- 

ous structures called paraphyses. (Figs. 129 and 130.) 

.Order 3. Plectascales, the blue and green molds. Examples: 

Aspergillus and Penicillium.

TAXONOMY 257 

Penicillium glaucum (green mold or mildew), a type of mildew, 

belonging to the Ascomycetes class of Fungi, forms sage-green crusts 

on bread, jellies, old boots, gloves, and various pharmaceutical 

preparations. It consists of a felt-like mass of interlaced tubular 

FIG. 130. A, B, Lachnea scutelata. A, Habit; B, ascus with paraphysis; C, D, 

Lachnea hemisphcerica; C, habit; D, ascus with paraphysis; E, Sarcosphcera arenosa 

habit; F, G, Sarcosphcera coronaria; F, ascus; G, habit; H, Sarcosphcera arenicola 

ascus with paraphysis. (See Die naticrlichen Pflanzenfamilien I, i, p. 181.) 

(Harshberger.) 

hyphae called a mycelium. From the mycelium numerous hyphae 

project into the air and bear a green powder, the spores. These 

hyphae are called aerial hypha. Other hyphae grow down into the 

substratum and are called submerged hypha. 

17 

H

258 


When a small portion of the mycelium is mounted in 10 per cent, 

alcohol and observed under the high-power objective, it will be noted 

that each hypha has a transparent wall and" protoplasmic contents 

and is divided by transverse septa into a number of cells. Each 

cell contains protoplasm, which is differentiated into cytoplasm (cell 

FIG. 131. Three aerial hyphae showing the characteristic brush-like branching 

and spore formation of Penicillium glaucum. This fungus is a true saprophyte 

and is never found on living fruits or vegetables, a, Conidiophore branching 

above into secondary conidiophores; b, sterigmata; c, conidiospores. (Schneider.) 

protoplasm) and several nuclei. In the cytoplasm will be seen 

several large clear spaces. These are vacuoles and contain water 

with nutritive substances in solution, called cell sap. Each hypha 

with its branches is clearly distinct from every other one.

TAXONOMY 259 

The aerial hyphae bear brush-like branches, which become con- 

stricted on their ends into a moniliform aggregation of rounded 

spores appearing like a row of beads. Each aerial hypha is composed 

of a vertical septate branch of the mycelium called the conidiophore, 

branches of this, which are called secondary conidiophores, and 

chains of spores at the tips of sterigmata (cells bearing conidia) 

which are called conidia or conidiospores. The conidia form the 

loose green powder characteristic of Penicillium. 

PIG. 132. Penicillium Roqueforti. a, Part of a conidiophore; b, c, other types 

of branching; d, young conidiophore, just branching, e. f, conidiiferous cells; g, 

h, j, diagrams of types of fructification, k, I, m, n, geminating spores. (After 

Thorn.) 

A number of species of Penicillium are useful in the arts. Peni- 

cillium roqueforti is the principal ripening agent of Roquefort, 

Gorgonzola and Stilton cheeses. It possesses blue-green globular 

conidia 4 to 5/4 in diameter. 

Penicillium camemberti is the principal agent in the ripening of 

Camembert cheese. It possesses ellipsoidal bluish-green conidia 

4.5 to 5.5/4 in diameter.


Penicillium bremcaule grows on old moist paper and has been used 

to detect the presence of arsenic, for when grown in media contain- 

ing this element, it develops the compound, diethylarsine. It is 

yellowish-brown in color and its conidia are rough and spiny. 

Q 

PIG. 133. Penicillium Camemberti. a, Conidiophore with common type. of 

branching with conidiospores; (&), a common less-branched form; c, d, f, diagrams 

of large fructifications; g, i,j, germinating conidiospores. " 

(From Bull. 82, Bureau 

of Animal Industry, also After Thorn.) 

Penicillium expansum is often found on decaying apples where it 

produces brownish coremia. 

Aspergillus herbariorum. This green mold also named Aspergillus 

glaucus and Eurotium Aspergillus glaucus is frequently found on

TAXONOMY 261 

fleshy drugs which have not been properly dried. It has also been 

observed on dried herbarium material, old extracts, on jams, jellies, 

tobacco, cotton-seed meal, old leather, stale black bread, etc. Like 

Penicillium its vegetative body consists of a mycelium consisting of 

aerial and submerged hyphae. It differs from Penicillium, however, 

mainly in not possessing septated conidiophores and by the upper 

portion of the conidiophores being globular. Upon the globular 

extremity of the conidiophores are placed numerous elongated sterigmata 

which bear chains of grayish-green conidia. These are spherical 

and prickly and range from 7 to 30/4 in diameter. Under certain 

FIG. 134. Penicillium brevicaule. a, Conidiophores and simple chains of conidiospores; 

b,f, more complex conidial fructifications; c, two young chains of conidiospores; 

d, e, echinulate conidiospores; g, h, j, sketches of forms and habits of 

conidial fructifications; k, germinated conidiospores. (After Thorn.) 

conditions closed brownish fruit bodies called perithecia are produced. 

These arise on the surface of the substratum from spirally coiled 

hyphae and when mature possess numerous asci, each of which con- 

tains five to eight ellipsoidal ascospores. 

Aspergittus oryza is a yellowish-green to brown mold which 

secretes diastase, a valuable digestive ferment, having the power of 

converting starch into sugar and dextrin. For centuries the Japanese 

have employed this species in the preparation of rice mash for


Sake, as well as in manufacture of Miso and Soja sauce. The spher- 

ical conidiospores are 6 to 7/4 in diameter and of a yellowish-green 

color. 

PIG. 135. Aspergillus oryzoe associated with yeasts in the making of the Japanese 

beverage Sake. Vegetative hyphae (a) and spore-forming hyphae(&, c, d) are 

shown. (Schneider, Pharmaceutical Bacteriology.} 

Aspergillus fumigatus is a pathogenic species which produces a 

disease in birds, horses, cattle and even though rarely in man that is

TAXONOMY 263 

called aspergillosis. The organ most prone to infection by this 

organism is the lung, although the skin, cornea, ears and other parts 

FIG. 136. Sterigmatocystis niger (Aspergillus niger} showing conidiophores and 

conidiospores formation with stages in germination of spores. (Harshberger, after 

Henri Coupin.) 

are also subject to its parasitic influence. It produces short coni- 

diophores with sterigmata bearing long chains of rounded, colorless

264 


conidia 2.5 to 3^ in diameter. Harshberger 1 cites the presence of 

perithecia in this organism which are nut-brown, globular, 250 to 

350/4 in diameter, and inclose oval thin-skinned asci with eight red 

lenticular ascospores each of which has a diameter of 4 to 5^. 

Aspergillus niger (Sterigmatocystis niger) develops dark, brown 

mycelial masses in which are to be noted slender conidiophores bear- 

ing handle-shaped, branched sterigmata that cut off from their tips 

chains of rounded black-brown conidia 3.5 to 5/x in diameter. This 

fungus has been found to produce suppurative inflammation of the 

FIG. 137. The morel, Morchella esculenta. (Gager, from photo byW.A. Murrill.) 

external and middle portions of the human ear. It is also a cause of 

cork disease, so often imparting a disagreeable taste to bottled 

beverages. 

Order 4. Tuberales, the truffles. Fungi whose septate mycelium 

is often connected with the roots of trees forming the structure 

known as mycorrhiza. 

Several species of the genus Tuber growing 

in woods of France, Germany and Italy produce tuberous subter- 

ranean bodies called Truffles, which are highly prized as a table 

delicacy by the inhabitants of these countries. 

Order 5. Helvellales, the saddle fungi. Fleshy fungi entirely 

*" Mycology and Plant Pathology" p. 147.

TAXONOMY 265 

saprophytic, living attached to leaf mold or growing in humous soil 

or, in a few cases, on decaying wood. The fleshy fruiting bodies 

(ascocarps) are divided into stalk (stipe) and cap (pileus) portions. 

The external surface of the cap is covered, with a layer of asci and 

paraphyses which together constitute the ascigeral layer. To this 

group belong the Morels and the Earth Tongues. 

One of the Morels, Morchella esculenta, is frequently found .in 

fire-swept woods. Its fruiting body consists of a hollow, externally 

ridged stipe, bearing upon its summit a fleshy pileus whose outer 

surface is honeycombed with ridges and depressions. The depressions 

are covered with an ascigeral layer composed of asci and 

paraphyses. This species is edible. 

Order 6. Pyrenomycetales, the mildews and black fungi common 

as superficial parasites on various parts of plants. To the black 

fungi division of this order the Ergot fungus, Claviceps purpurea 

belongs. 

Life History of Claviceps Purpurea. Through the agency of 

winds or insects the spores (ascospores or conidia) of this organism 

are brought to the young ovaries of the rye (Secale cereale). They 

germinate into long filaments called hyphae, which, becoming en- 

tangled to form a mycelium, spread over the ovary, enter it superficially, 

secrete a ferment, and cause decomposition of its tissue and 

the resultant formation of a yellow-mucous substance called honeydew, 

which surrounds chains of moniliform reproductive bodies 

known as conidia. The honey-dew attracts certain insects which 

disseminate the disease to other heads of grain. 

The mycelial threads penetrate deeper and deeper into the ovary 

and soon form a dense tissue which gradually consumes the entire 

substance of the ovary and hardens into a purple, somewhat curved 

body called a sclerotium, or official ergot the resting stage of the 

fungus, Claviceps. 

The ergot falls to the ground and in the following spring sprouts 

into several long stalked, globular heads called stromata or ascocarps. 

Each (fruiting) head or ascocarp has imbedded in its surface numerous 

flask-shaped invaginations called perithecia, from the bases 

of which several sacs or asci develop. Within each ascus are 

developed eight filiform spores (ascospores) which, when the ascus

266 


PIG. 138. A, Balansia claviceps on ear oiPaspalum; B-L, Claviceps purpurea; 

E, 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 ascospore; C, conidiospores 

produced on mycelium. (See Die naturlichen Pflanzenfamilien I, i, p. 371.)

TAXONOMY 267 

ruptures, are discharged and are carried by the wind to other fields 

of grain, there to begin over a new life cycle. 

CLASS III. BASIDIOMYCETES, OR BASIDIA FUNGI 

This large class of fungi, including the smuts, rusts, mushrooms, 

the occurrence of a 

gill and tooth fungi, etc., is characterized by 

basidium in the life history. A basidium is the swollen end of a 

hypha consisting of one or four cells and giving rise to branches called 

sterigmata, each of which cuts off at its tip a spore, called a basidio- 

spore. In addition to the basidiospores, some forms also produce 

spores termed chlamydospores. 

SUB-CLASS A. PROTOBASIDIOMYCETES 

(Basidium four-celled, each bearing a spore) 

Order i. Ustilaginales, the smuts. Destructive parasites which 

attack the flowers of various cereals, occasionally other parts of these 

plants. Example: Ustilago Maydis, 

the corn smut. The basidio- 

spores in this group are borne on promycelia. 

Ustilago Maydis (Ustilago Zeae) (Corn Smut). Corn smut is a 

destructive parasite which for a long time was supposed to be confined 

to the Indian Corn, but which now is known to occur on 

Mexican Grass. It is the only smut useful in medicine. The 

mycelium of the fungus extends through all parts of the infected 

host through the intercellular-air-spaces and produces large tumor- 

like masses on the ears, tassels, husk, leaves and stem. Each mass is 

filled with spores and covered with a tightly appressed membrane 

which has a whitish appearance like German silver. The spores 

are at first a dark olive-green, but on maturity are dark brown. 

They are sub-spherical and show prominent spines. They arise 

by the division of the septate mycelium into thick-walled echinulate 

resting spores called chlamydospores or brand spores. These spores 

fall to the ground and pass the winter. In the spring each germinates 

into a three- or four-celled filament called a promycelium, from 

the cells of which basidiospores arise. The basidiospores develop a 

mycelium which penetrates the seedling of the host plant.


Order 2. Uredinales, the rusts. Obligate parasites possessing 

the inter- 

a septated branched mycelium which ramifies through 

FIG. 139. Smut boil of Ustilago zeoe on ear of corn, developed from one infected 

kernel. (After Jackson, F. S., Bull. 83, Del. Coll. Agric. Exper. Stat., 

December, 1908.) 

cellular-air-spaces 

of the host and sends haustoria into the cell 

cavities. The different stages of their life cycle are either restricted

TAXONOMY 269 

to one host or distributed between two or more hosts. An outline 

of the life history of the wheat rust will give an idea of the peculiari- 

ties of the group. 

The Wheat Rust (Puccinia Graminis). If we examine the wheat 

plant just 

before harvest we will find on the stems and leaves 

FIG. 140. 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 

/*20 Per cent, acetic acid, formation of infection threads, a, Spores; b, promycelia; 

c, basidiospores; d, infection threads; e, detached pieces of mycelia. (After 

Bull. 57, Univ. III. Agric. Exper. Stat., March, 1900.) 

some rust-red lines. The presence of the mycelium of the fungus in 

the intercellular spaces of the host does not kill the host directly 

or appear to stunt its growth, but the effect of the parasite on the 

host is seen when the grains mature. The grains are small and 

mushy, due to the fact that the nutrition of the host had been disturbed 

and the formation of starch in the grains inhibited, The

270 


mycelium is localized and gives rise underneath the epidermis to 

rounded egg-shaped spores attached to it by short pedicels. The 

spores are produced in such numbers that the space beneath is too 

confined. As the long epidermal cells of grasses run longitudinally, 

the pressure of the spore masses from within causes the epidermis to 

crack and its edges become turned back. Through the resultant 

cleft the summer spores or uredospores are thrust out. These uredo- 

spores are orange-brown in color and covered with minute spines. 

The mass of them has been called a uredinium. These spores are 

FIG. 141. Spore forms of wheat rust, Pucainia graminis. A, Section through 

barberry leaf showing pycnia on upper surface and aecia on lower; B, two uredinio 

spores; C, germinating urediniospore; D, teliosorus showing several teliospores; 

E, single two-celled teliospore; F, germirating teliospore with four-celled 

basidium and two basidiospores; G, basidiospore growing on barberry leaf. 

(Harshberger, adapted from deBary.) 

detached from the pedicels and blown by the wind to healthy plants. 

After summer is over and dry weather comes on, an examination of 

stubble in the field (blades of grass and stems of wheat left carelessly), 

these rust-red lines are replaced by brownish-black spores called 

teleutospores (teliospores). A mass of these is known as a telium. 

The summer stage on wheat is known as Uredo linearis. 

The autumn stage on wheat is known as Puccinia graminis.

TAXONOMY 2 7 I 

The teleutospores are two-celled and have thick walls and persistent 

pedicels. They remain attached to the stubble until the 

following spring and then either one or both cells composing them 

produce an outgrowth known as a promycelium (nothing 

but a 

basidium divided transversely into four cells). Each cell of the 

basidium is capable of producing a branch, at the tip of which a 

basidiospore is formed. These basidiospores are blown to the 

Barberry (Berberis) and infect the leaves of this plant. The 

mycelium runs in the intercellular-air-spaces and causes the appearance 

of a number of small depressions on the upper side of the leaf. 

These in section are a rich chocolate brown and known as spermagonia. 

In the center of a spermagonium are produced hyphae, which 

project out to its orifice and obstrict off minute spores called spermacia. 

On the opposite side of the leaf cup-shaped depressions 

are formed, each with a limiting membrane (peridium). Within 

the cup-shaped depression thousands of spores are formed in chains 

closely packed together. These are the aecidiospores (aeciospores) . 

The cluster cup is called an Êcidium (Êcium). These aecio- 

spores are conveyed to wheat and cause infection, thus completing 

the life cycle. It has been observed that in America the uredospores 

or summer spores may winter over and infect healthy plants, so that 

the Barberry phase is completely eliminated from the life cycle. 

Order 3. Auriculariales. The so-called "ear fungi" which occur 

on the bark of many plants, on wooden fences, etc., as auriculate 

growths which when young are jelly-like and brilliantly colored, 

when old, hard, grayish and considerably wrinkled. The ear- 

like fruiting body is known as the sporophore. Its internal surface 

is lined with a hymenium or fruiting body consisting of numerous 

four-celled basidia, each of which cuts off at its tip a basidiospore. 

Order 4. Tremellales. Saprophytes which live on decaying 

wood as moist, soft, quivering, gelatinous growths becoming later dry 

and horny. 

SUB-CLASS B. AUTOBASIDIOMYCETES 

(Mostly fleshy forms characterized by one-celled basidia with generally four, 

occasionally six, eight or two sterigmata each of which cuts off a basidiospore at 

its tip.)

272 


Division a. Hymenomycetes 

(Hymenium or spore-bearing surface exposed) 

This division of Autobasidiomycete or higher basidiomycete 

fungi comprises the following orders: Dacromycetales, Exobasidiales, 

Thelephorales, Clavariales and Agaricales. 

Order i. Dacromycetales. This order includes the "weeping 

fungi." One of the most common is Dacromyces deliquescens which 

FIG. 142. Coral-like fruit-bodies of Clavaria flava. (Harshberger, from Photo 

by W. H. WalmsJey.} 

occurs as a gelatinous body of bright red color on dead wood. The 

basidiospores are formed during a wet period and the fungus swells 

up in the water forming a slimy mass. In addition to basidiospores 

the mycelium may break up into oidiospores, if the wet period is 

prolonged. In consisting of slimy gelatinous masses the "weeping 

fungi" approach the Tremellacea but are distinguished from them

in the basidium being 

TAXONOMY 2 73 

undivided in the former and divided in the 

latter. 

Order 2. Exobasidiales. This group is found growing parasitically 

on shrubs especially those of the heath family. The mycelium 

lives in the tissues of the stems, leaves, sepals and petals and pro- 

duces spongy fleshy yellowish or brownish galls which are popularly 

called "Azalea apples." The galls are edible. They are covered 

with a hymenium. 

FIG. 143. Boletus felleus in three stages of development. (After Patterson, 

Flora W. and Charles, Vera K., Bull. 175, U. S. Dept. Agric., pi. xxxi, Apr. 29, 

I9IS.) 

Order 3. Thelephorales, forms appearing on tree trunks as 

leathery incrustations or as bracts on the ground, old logs, etc. 

Order 4. Clavariales, the coral or fairy club fungi. Fleshy coral 

or club-shaped forms, all of which are saprophytes found in woods 

growing in bunches out of leaf mold. They are all edible and of a 

white, yellow or some other brilliant color. (See fig. 142.)

274 

PHARMACEUTICAL B OTANY 

Order 5. Agaricales, the mushroom or toadstool alliance. Alike 

with the other members of the Basidiomycetes, the plant body con- 

sists of the mycelium, ramifying through the substratum, but the 

part which rises above the surface (the Sporophore) is in most cases 

differentiated into a stalk-like body called a stipe bearing upon its 

bummit a cap or pileus, the latter having special surfaces for the 

hymenium. 

Family I.^-Hydnaceae, or tooth fungi. This group is characterized 

by the hymenium being placed over purple-like, spiny or 

long digitate projections of the pileus. Many of the species of the 

genus Hydnum are edible. 

Family II. Polyporaceae, or pore fungi. The sporophores or 

fruiting bodies of these fungi are various. They may be entirely 

supinate with pores or shallow depressions on their upper surfaces 

(Merulius), or mushroom-like (Boletus), or of the nature of woody 

(Fomes) or fleshy (Fistulina) brackets. In all cases the hymenium 

or basidial layer lines the inner surface of pores. 

The sporophore of Polyporus officinalis, when deprived of its outer 

rind and dried, constitutes the official N.F. drug AGARICUS. This 

species grows abundantly on various species of pines, spruces and 

larches. 

Family III. Agaricaceae, the gill family, in which the hymenium 

covers blade-like plates of the pileus, called gills, generally occurring 

on the under surface of the same. Examples: Agaricus campestris, 

the common edible mushroom of fields; Amanita muscaria and 

Amanita phalloides, both of which are poisonous. 

Agaricus Campestris (Common Mushroom). This plant is an 

edible gill fungus which grows in open, grassy fields during late summer 

and early autumn. It is never found in the forest or on trees or 

fallen trunks, seldom in the mountains. The cultivated form grows 

in specially constructed houses made of boards. A corridor runs 

through these houses so that the mushroom beds can be easily 

reached. In the growth of mushrooms tons of horse manure are 

used. This is covered with loamy soil i^ inches thick. The whole 

mass is compacted together. Into the resultant beds is introduced 

English-grown spawn, which comes in flat brick-shaped masses (horse 

manure through which mycelium has grown). Pieces of these

TAXONOMY 275 

"bricks" are put in the horse manure bed only after the heat has 

first disappeared. The beds are then watered well and in a short 

time the sporophores or fruiting bodies of the fungus spring up. 

The mycelium or vegetative body of Agaricus which develops 

in the soil from spores (basidiospores) is white and thready. On this 

mycelium develops little buttons, first about the size of a pin head, 

becoming later pea size and then assuming a pear-shaped form. At 

this stage the sporophore consists of a cylindrical solid stipe or stalk 

and a pileus or cap. The border of the pileus is joined to the stipe 

FIG. 144. Meadow mushroom (Agaricus campestris L.). A, view showing 

under side of pileus; g, gills; a, annulus, or remains of the veil attached to^the 

stipe; B, side view; s, stipe; a, annulus; p, margin of pileus, showing at intervals 

the remains.of the veil. (Gager, after W. A. Murrill.) 

by means of a "partial veil." Within this veil is found a circular 

cavity, into which the gills grow. At first the stipe grows faster than 

the rest of the fruiting body. The pileus expands transversely and 

the gills keep pace. After a while the veil ruptures, leaving a portion 

attached to the stipe. This constitutes the annulus or ring (true 

annulus). The hyphae in the pileus form the Tela contexta. If we 

make a section through a gill, the hyphae are seen to run longitudinally. 

The central part is called the trama] next and outside trama 

is the sub-hymenium; next, hymenium, consisting of basidia (hence a

276 

basidial layer). 

PHARMACEUTICAL BOTAN\ 

Each basidium bears one or two little points known 

as sterigmata. Each sterigma bears a purplish-brown basidiospore. 

The basidiospores falling to the ground germinate into hyphae and 

these become interlaced to form a mycelium. 

In the wild mushroom the gills are at first pink, in cultivated, 

fawn-colored. Ultimately in the wild form the gills turn brownish. 

The spores are purplish-brown. The color of the stipe and upper 

surface of the pileus varies from whitish to a drab color. 

PIG. 145. Deadly amanita (Amanita muscaria) 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.) 

The Amanitas (Poisonous Fungi). Amanita muscaria and Aman- 

ita phalloides, commonly known as the "fly agaric" and the "deadly 

agaric" respectively, are very poisonous forms. Amanita muscaria 

is common in coniferous forests, although may occasionally be found 

in grassy places. It occurs singly and not in groups. Amanita 

phalloides is found in woods and borders of fields and, like the fly 

agaric, occurs singly and not in groups. 

Each of these have fruiting bodies (sporophores), which begin at

TAXONOMY 277 

the surface of the ground as a button similar to that of the edible 

mushroom. This enlarges and assumes a dumbbell shape. The 

whole button is covered by an ouster veil, known as the velum universale, 

which encloses the pileus, gills and stipe. As the stipe lengthens 

more rapidly than the pileus, the upper part of the veil is stretched 

and finally breaks in its middle portion. The lower part remains as 

FIG. 146. The deadly amanita, Amanita phalloides. Note the cup at the base 

of the stipe. (Gager, from photo by E. M. Kittredge^ 

a cup, out of which the stipe grows. The upper part is carried up as 

shreds adhering to the margin of the pileus. The lower part is 

called the volva or death cup. The annulus present is a false annulus, 

for it represents a peeling down of the upper part of the stipe. 

have chalk-white gills, a white stipe, and white spores. 

Both

278 


The pileus of Amanita muscaria is yellow, or orange-red; the 

surface is smooth, with prominent warty scales. 

The pileus of Amanita phalloides varies from dull yellow to olive 

to pure white. It does not possess the warty scales found in the 

Amanita muscaria, but occasionally has a few membranous patches. 

Division b. Gasteromycetes 

(Hymenium inclosed) 

Order i . Lycoperdales, or puff ball alliance . This order includes 

a number of interesting parasites and saprophytes the most common 

of which are the earth stars belonging to the genus Geaster and the 

FIG. 147. A colony of Puff Balls, Lycoperdon, growing saprophytically upon a 

portion of a rotten log. (Photograph by author.) 

puff balls, the most common form being Lycoperdon. In these, the 

fruiting sporophore consists for the most part of a shell-like covering 

called the peridium, composed of an outer layer or exoperidium and 

an inner layer or endoperidium. The peridium in the unripe con- 

dition of the sporophore covers a mass of soft cellular tissue called 

the gleba. Upon the ripening of this mass, the interior is seen to be 

divided into many-branched compartments that are separated from 

each other by walls made up of branched hyphae. These walls are 

lined with a hymenium composed of many basidia, each of which

TAXONOMY 279 

constricts off usually four basidiospores. The earth stars differ 

from the puff balls in possessing an outer wall or exoperidium which 

splits in star-shaped fashion. 

Order 2. Nidulariales, the nest fungi. A group of Gasteromy- 

cetes whose sporophores are crucible- or crater-like. These arise 

from a subterranean mycelium and show 

an" outer and inner peridial layer. 

The 

outer peridium is roughened at its base. 

The inner peridium is leathery and may 

or may not be continued over the top. 

When mature the crucible-like body 

shows black seed-like bodies inside which 

resemble eggs in a bird's nest. .Each 

one of these is connected with the inner 

peridium by a cord which resembles the 

umbilical cord of an animal. These 

inner bodies are called periodiola (sing. 

peridiolum). Each peridiolum consists 

of a hard glistening outer layer and a 

spongy inner layer surrounding a cavity 

into which basida and basidiospores pro- 

ject. These fungi are found in stiff 

clayey soil. 

Order 3. Phallales, the carrion or 

stink-horn fungi. This, the highest 

group of the Autobasidiomycetes, con- 

sists of highly and characteristically 

colored forms which, when mature, emit 

most vile and penetrating odors. The 

fruiting body, in each instance, begins 

, , , . , 

as an egg-shaped structure which starts 

FIG. 148. Mature stink- 

horn> Diayophora duplicate 

its growth from a widely spread under- 

, ,. , , -,-. i 

(Harshberger, from photo, 

W.H. Walmsley.) 

by 

ground mycelium of chalky-white color. 

As the "eggs" enlarge they push above the surface of the ground. 

The central portion, elongating, then breaks through the outer or 

peridial portion, which remains as a cup or volva at the base of the 

mature fruit body. Upon the summit of the central stalk rests the


cup-like many-chambered gleba. 

The basidiospores are imbedded 

in a greenish fetid slime formed by a mucilaginous disintegration 

of the substance of the hymenium. This fetid green material is 

attractive to carrion flies which visit the plants and remove the 

material with its embedded spores. The latter will riot germinate 

until after passing through the alimentary canal of these flies. 

CLASS IV. FUNGI IMPERFECTI 

An assemblage of varied forms, the life histories of most of which 

are imperfectly understood. In this group are included numerous 

parasites which produce diseases in crop plants. 

SUBDIVISION V. LICHENS, THE LICHENS 

Lichens are variously colored, usually dry and leathery plants, 

consisting of symbioses of algae and fungi. In each case the fungus 

FIG. . 149. A foliaceous lichen, Physica sldlaris (L.) Nyb. t growing on a rock. 

The cup-shaped structures are the fruiting bodies (apothecia). 

seen two very young plants. (Gager.} 

At the left are 

derives its food from materials manufactured by the algae and in 

return extracts water from the substratum and shares it with the 

algae. The association is therefore mutually beneficial. Blue- 

green and Protococcus forms of Green Algae and Ascomycete Fungi 

are for the most part concerned in lichen formation.

TAXONOMY . 

28l 

Lichens are found on the bark of trees, on rocks, logs, old fences, 

etc. The body of a lichen shows a differentiation into two regions: 

a more or less compact row of cells on both surfaces, called the epider- 

mis; and an inner portion composed of the mycelium of the fungus. 

The alga is imbedded in this portion. In most cases the spores are 

borne in asci, which are themselves found in closed or open Apothecia. 

Scales or soredia are found on many lichens. These consist of a 

network of hyphae enclosing algal cells. By becoming detached 

from the parent plant, they develop new lichens and so constitute 

a means of vegetative propagation. 

PIG. iô.Cetraria islandica. (Sayre.) 

According to the manner of growth of the thallus and nature of 

attachment to the substratum, three different sub-groups of lichens 

may be distinguished, viz.: (i) Foliaceous where the thallus is flat, 

leathery and leaf-like and attached to the substratum at different 

points. To this group belong Physica and Parmelia. (2) Crus- 

taceous, where the thallus closely adheres to rocks and bark of trees. 

To this group belong Gr aphis and Pertusaria. (3) Fruticose, where 

the thallus is upright and branching. To the last group belong 

Cetraria islandica, species of Cladonia, and Usnea. 

To the pharmacist and chemist lichens are chiefly of interest 

because of the coloring principles which they contain. Species of 

Lecanoraand Rocella tinctoria yield, when subjected to fermentation,


the dyes orcein and litmus. Litmus is one of the best indicators in 

volumetric analysis. Cudbear, a purplish-red powder, used exten- 

sively for coloring pharmaceutical preparations in the form of tincture, 

is prepared by treating species of Rocella, Lecanora or other 

lichens with ammonia water. Other lichens, such, as Cetraria 

islandica, various species of Parmelia, Usnea and Alectoria, have 

been used in medicine because of demulcent principles which they 

contain. 

DIVISION II. BRYOPHYTA 

Plants showing a beginning of definite alternation of genera- 

tions, i.e., gametophyte (sexual phase) alternating with sporophyte 

gon 

FIG. 151. PIG. 152. 

FIG. 151- Section of thallus of Cetraria islandica through an apothecium. as, 

Asci, three of which contain ascospores. gon, Gonidia. The inner (central portion 

shows the mycelial threads of a fungus entangling the alga. (Sayre.) 

FIG. 152. A liverwort (Lunularia). Below, portions of the thallus, showing 

the lunar-shaped cupules, with brood-buds, or gemmae. Above a single gemma, 

greatly magnified. (Gager.) 

(asexual phase of development) in their life history, the two phases 

being combined in one plant. The female sexual cell is always 

lodged in an archegonium (a multicellular female sexual organ).

TAXONOMY 283 

SUBDIVISION I HEPATIC^ OR LIVERWORTS 

Plants of aquatic or terrestial habit whose bodies consist of a 

rather flat, furchate branching thallus or leafy branch which is 

dorsoventral (having distinct upper and lower surface) ; the upper 

surface consists of several layers of cells containing chlorophyll, 

which gives the green color to the plants; the lower surface gives 

origin to hair-like outgrowths of the epidermal cells serving as absorptive 

parts and called rhizoids. Upon the dorsal surface of this thalloid 

body (the gametophyte) cup-like structures are produced called 

cupules which contain special reproductive bodies called gemma, 

these being able to develop into new gametophytes. The sex organs 

are of two kinds, male and female. The male organs are termed 

antheridia, the female, archegonia. The antheridia are more or less 

club-shaped, somewhat stalked organs consisting of an outer layer of 

sterile cells investing a mass of sperm mother-cells from which are 

formed the spirally curved biciliate antherozoids, or male sexual cells. 

The archegonia are flask-shaped organs consisting of an investing 

layer of sterile cells surrounding an axile row of cells, the neck-canal 

cells, ventral-canal cells and the egg or female sexual cells. Every 

ell of the axial row breaks down in the process of maturation with 

the exception of the egg which remains in the basal portion. Both 

antheridia and archegonia generally arise on special stalks above the 

dorsal surface. After the egg is fertilized by a antherozoid, the 

young embryo resulting grows into a sporogonium (the sporophyte) 

consisting of a stalk portion partly imbedded in the archegonium 

surmounting a sporangium or capsule in which spores are produced. 

When mature the capsule splits open discharging the spores. The 

spores on germination develop into a protonema or filamentous 

outgrowth which later develops the thallus. 

Order i. Marchantiales, including Marchantia and Riccia. 

Order 2. Jungermanniales, the leafy liverworts, including 

Porella. 

Order 3. Anthocerotales, having the most complex sporophyte 

generations among liverworts, including Anthoceros, and 

Megaceros.

284 


SUBDIVISION II. MUSCI OR MOSSES 

Plants found on the ground, on rocks, trees and in running water. 

Their life histories consist of two generations, gametophyte and 

FIG. 153. Sphagnum acutifolium, Ehrb. A, prothallus (pr), with a young leafy 

branch just developing from it; B, portion of a leafy plant; a, male cones; ch, 

female branches; C, male branch or cone, enlarged with a portion of the vegetative 

branch adhering to its base; D, the same, with a portion of the leaves removed so 

as to disclose the antheridia; E, antheridium discharging spores; F, a single sperm; 

G, longitudinal section of a female branch, showing the -archegonia (ar); H, 

the foot; PS, pseudopodium; c, 

longitudinal section through a sporogonium; sg l , 

calyptra; sg, sporogonium, with dome of sporogenous tissue; ar, old neck of the 

archegonium; J. Sphagnum squarrosum Pers. ; d, operculum; c, remains of calyptra; 

qs, mature pseudopodium; ch, perichaetium. (Gager, from Schimper.) 

sporophyte similar to the liverworts but differ from liverworts, 

generally, by the ever-present differentiation of the gametophyte

TAXONOMY 285. 

body into distinct stem and simple leaves, and the formation of the 

sexual organs at the end of an axis of a shoot. They are either monoe- 

cious, when both kinds of sexual organs are borne on the same plant, 

FIG. 154. Hair-cap moss (Polytrichum commune). A, male plant; B, same, 

proliferating; C, female plant, bearing sporogonium; D, same; g, gametophyte; s, 

seta; c, capsule; o, operculum; a, calpytra; E, top view of male plant. (Gager.),


or dioecious, in which case the antheridia and archegonia arise on 

different plants. 

Order i. Sphagnales, or Bog Mosses, including the simple genus, 

Sphagnum. Pale mosses of swampy habit whose upper extremities 

repeat their growth periodically while their lower portions die away 

gradually and form peat, hence their frequent name of Peat Mosses. 

A number of species of Sphagnum have been recently employed 

in surgery as absorbents in place of gauze. -For this purpose they 

must be thoroughly cleaned and sterilized. 

Order 2. Andreaeales, including the single, genus Andreaea, of 

xerophytic habit, occurring on siliceous rock. 

Order 3. Bryales, or true mosses, comprising^ the most highly 

evolved type of bryophytes. Examples: Polytrichum, Funaria, 

Hypnum, and Mnium. 

Life History of Polytrichum Commune (A Typical True Moss). 

Polytrichum commune is quite common in woods, forming a 

carpet-like covering on the ground beneath tall tree canopies. It is 

dioecious, the plants being of two kinds, male and female. 

Beginning with a spore which has fallen to the damp soil, we note 

its beginning of growth (germination) as a green filamentous body 

called a protonema. This protonema soon becomes branched, giving 

rise to hair-like outgrowths from its lower portion called rhizoids 

and lateral buds above these which grow into leafy stems commonly 

known as "moss plants. 75 

At the tips of some of these leafy stems 

antheridia (male sexual organs) are formed while on others arche- 

gonia (female sexual organs) are formed. These organs are surrounded 

at the tips by delicate hairy processes called paraphyses 

as well as leaves for protection. The antheridia bear the antherozoids, 

the archegonia, the eggs or ova, as in the liverworts. When 

an abundance of moisture is present, the antherozoids are liberated 

from the antheridia, swim through the water to an archegonium 

and descend the neck canal, one fertilizing the egg by uniting with 

it. This completes the sexual or gametophyte generation. The 

fertilized egg now undergoes division until an elongated stalk 

bearing upon its summit a capsule is finally produced, this being 

known as the sporogonium. The base of the stalk remains imbedded 

in the basal portion of the archegonium, at the tip of the leafy

TAXONOMY 287 

stalk, and forms a foot or absorbing process. In growing upward 

the sporogonium ruptures the neck of the archegonium and carries 

it upward as the covering of the capsule, or calyptra. The calyptra 

is thrown off before the spores are matured within the capsule. 

The upper part of the capsule becomes converted into a lid or operculum 

at the margin of which an annulus or ring of cells forms. The 

cells of the annulus are hygroscopic and expand at maturity, throw- 

ing off the lid and allowing the spores to escape. This completes 

PIG. 155. Protonemata of a moss bearing young gametophyte bud. (Gager.} 

the asexual or sporophyte generation. The spores falling to the 

damp soil germinate into protonemata, thus completing 

the life 

cycle in which is seen an alteration of generations, the two phases, 

gametophyte alternating with sporophyte. 

DIVISION III. PTER1DOPHYTA 

The most highly developed cryptogams showing a distinct alter- 

nation of generations in their life history. They differ from the 

Bryophytes in presenting independent, leafy, vascular, root-bearing 

sporophytes.


SUBDIVISION 1. LYCOPODINEÊ OR CLUB MOSSES 

Small perennial, vascular, dichotomously branched herbs with 

stems thickly covered with awl-shaped leaves. The earliest forms of 

vascular plants differing from ferns in being comparatively simple in 

structure, of small size, leaves sesssile and usually possessing a single 

vein. Except in a few instances the sporangia are borne on leaves, 

crowded together and forming cones or spikes at the ends of the 

branches Homosporous. 

FIG. 156. Lycopodium clavatum. (Gager.) 

Family I. Lycopodiaceae, including the single genus Lycopodium 

with widely distributed species. The spores of Lycopodium clavatum 

are official. 

Family II. Selaginellaceae, including the single genus Selaginella 

with species for the greater part tropical. Plants similar in habit to 

the Lycopodiaceae but showing heterospory. 

Family III. Isoetaceae, including the single genus Isoetes whose 

species are plants with short and tuberous stems giving rise to a tuft

'TAXONOMY 289 

of branching roots below and a thick rosette of long, stiff awl-shaped 

leaves above Heterosporous. 

SUBDIVISION II. EQUISETINE.E 

(The Horsetails or Scouring Rushes) 

The Equisetineae, commonly known as the Horsetails or Scouring 

rushes, are perennial plants with hollow, cylindrical, jointed and 

FIG. I5i.3elaginella Martensii. a, vegetative branch; b, portion of the 

stem, bearing c^nes (x) ; c, longitudinal section of a cone, showing microsporangia 

(mic. sp.) in the axils of microsporophylls, and megasporangia in the axils of megasporophylls; 

d, microsporangium with microsporophyll; e, microspores; /, portion 

of wall of sporangium, greatly magnified; g, megaspore; h, microsporangium 

opened, and most of the microspores scattered; i, megasporangium, with megasporophyll; 

k, same, opened, showing the four megaspores. (Gager.) 

fluted stems, sheath-like whorls of united leaves and terminal cone- 

like fructifications. Their bodies contain large amounts of silicon, 

hence the name scouring rushes. 

19

2 90 


In some varieties the fruiting cone is borne on the ordinary 

stem, in others on a special stem of slightly different form. In the 

latter the spores are provided with elaters, which, being hygroscopic, 

coil and uncoil with increase or decrease in the amount of moisture 

present, thus aiding in the ejection of spores from the sporangia. 

The number of species is small and included under one genus, 

Equisetum. (See fig. 158.) 

SUBDIVISION III. 

The group Filicineae is the largest among the vascular cryptogams 

and includes all the plants commonly known as Ferns. The main 

axis of a typical fern is a creeping underground stem or rhizome 

which at its various nodes bears rootlets below and fronds above. 

These fronds are highly developed, each being provided with a 

petiole-like portion called a stipe which is extended into a lamina 

usually showing a forked venation. Some ferns possess laminae 

which are lobed, each lobe being called a pinna. If a pinna be 

further divided, its divisions are called pinnules. The unfolding of a 

frond is circinate and it increases in length by apical growth. On 

the under surface of the laminae, pinnae, or pinnules may be seen 

small brown patches each of which is called a sorus, and usually 

covered by a membrane called the indusium. Each sorus consists 

of a number of sporangia (spore cases) developed from epidermal 

cells. In some ferns the entire leaf becomes a spore-bearing organ 

(sporophyll) . Most sporangia have a row of cells around the margin, 

the whole being called the annulus. Each cell of the annulus has a 

U-shaped thickened cell wall. Water is present within these cells and 

when it evaporates it pulls the cell walls together, straightening the 

ring and tearing open the weak side. The annulus then recoils and 

hurls the spores out of the sporangium. Upon coming into contact 

with damp earth each spore germinates, producing a green septate 

filament called a protonema. This later becomes a green heart- 

shaped body called a prothallus. It develops on its under surface 

antheridia or male organs and archegonia or female organs as well 

as numerous rhizoids. Within the antheridia are developed motile 

sperms, while ova are produced within the archegonia. The many 

ciliate sperms escape from the antheridia of one prothallus during a

TAXONOMY 2 9 I 

wet season, and, moving through the water, are drawn by a chemotactic 

influence to the archegonia of another prothallus, pass down 

the neck canals of these and fuse with the ova, fertilizing them. 

The fertilized egg or oospore divides and redivides and soon becomes 

differentiated into stem-bud, first leaf, root, and foot. The foot 

FIG. 158. Equisetum arvense. P, sterile branch; P 1 , fertile branch with 

strobilus, or cone; R, rhizome (underground); T, cross-section of cone, showing 

1 insertion of sporophylls in a whorl; N, N , sporophylls with pendant sporangia; 

S, S l , S 2 , spores with coiled elaters (el). (Gager.) 

obtains nourishment from the prothallus until the root grows into 

the soil, when it atrophies, and the sporophyte becomes independent. 

Unequal growth and division of labor continue until a highly differentiated 

sporophyte results, the mature "fern plant."


ORDER i. FILICALES OR TRUE FERNS (HOMOSPOROUS) 

Family Polypodia cea. Sporangia with annulus vertical and 

incomplete. 

The rhizomes and stipes of Dryopteris filix-mas and Dryopteris mar- 

ginalis 

are official in the U. S. P. The fibro-vascular bundles of 

these are concentric in type but differ from the concentric fibro- 

vascular bundles of some monocotyledons in that xylem 

most and phloem surrounds the xylem. 

is inner- 

FIG. 159. Cyrtomyim falcalum. Under (dorsal) surface of a portion of a 

sporophyll, showing the numerous sori on the pinnae. (Gager.) 

ORDER 2. HYDROPTERALES OR WATER FERNS 

(HETEROSPOROUS) 

Family Sahiniacea. Floating ferns with broad floating leaves and 

submerged dissected leaves which bear sporocarps. 

mnia and Azolla. 

Examples: Sal- 

DIVISION IV. SPERMATOPHYTA (PHANEROGAMIA) 

Plants producing real flowers and seeds. The highest evolved 

division of the vegetable kingdom. 

SUBDIVISION I. GYMNOSPERM^: THE GYMNOSPERMS 

The Gymnosperms comprise an ancient and historic group of seed 

plants which were more numerous in the Triassic and Carboniferous

TAXONOMY 293 

periods than now. They differ from the Angiosperms in several 

respects, viz.: they bear naked ovules on the edges or flat surfaces 

of leaves called carpels, while Angiosperms bear covered ones; each 

megaspore produces within itself a bulky prothallus, in the upper 

portion of which originate one or more archegonia, while in Angiosperms 

no recognizable prothallus has been proven to exist; the 

stored food tissue within their seeds is prothallial tissue loaded with 

starch, etc., while that in Angiosperm seeds (endosperm) is developed 

from the endosperm nucleus; the mode of growth of their stems is 

always indefinite while that of Angiosperms is either indefinite or 

definite. 

FIG. 160. Cycas revoluta, showing terminal bud of foliage-leaves just opening. 

(Gager.) 

The groups still extant are the Cycads or Fern Palms, the Gne- 

and the Conifers. Of these the 

tums, the Ephedras, the Ginkgos 

Conifers comprising over 300 species are the most numerous. Many 

of them yield valuable products to pharmacy and the arts. 

The Conifers include the pines, spruces, hemlocks, cedars, firs, 

arbor vitae, chamaecyparis, and larches. All of their number are 

evergreen except the larches, which drop their foliage upon the 

advent of winter.

294 


I. Order Coniferales. Trees with a single upright stem which 

develops side branches that spread out horizontally and taper to a 

point at the summit, giving the crown of the tree the appearance of 

a huge cone, rarely shrubs. 

Pinacea (Conifera) or Pine Family. Trees or shrubs with resin- 

ous juice whose wood is characterized by being composed largely of 

tracheids with bordered pits. 

Leaves entire, awl- or needle-shaped 

frequently fascicled, exstipulate, usually evergreen. Flowers, 

monoecious or rarely dioecious, achlamydeous, in cones. Staminate 

PIG. 161. Inflorescences of the pine, i, Terminal twig; 2, ovulate cone; 3, 

staminate cone; 4, two-year-old cone. (Hamaker.) 

cone of a large number of microsporophylls (stamens) closely packed 

together and arranged spirally around a central axis, each stamen 

bearing usually two pollen sacs. Carpellate cone composed of 

spirally arranged scales, each of which bears a pair of naked ovules 

(megasori) near the base of its upper face, or, ovules springing from a 

cupuliform disc. Fruit a cone with woody or fleshy scales (Pinus, 

Thuja, Abies, Picea, etc.), a galbalus (Juniperus) or a drupe composed 

of the thickened and fleshy disc surrounding an erect seed 

(Taxus). Seeds albuminous. Embryo with two or more cotyledons .

Official drug 

Part used 

Terebinthina N.F. Concrete oleoresin 

Resina 

Oleum Tere- 

binthinae 

Resin 

Volatile oil 

Fix Liquida . . Product of destruc 

tive distillation 

Pinus Alba N.F. Inner bark 

Oleum Pini Pumi- Volatile oil 

lionis 

Terebinthina Oleoresin 

Laricis N.F. 

Juniperus N.F. Fruit 

Oleum Juniperi 

Volatile oil 

Oleum Cadinum Empyreumatic oil 

Thuja N.F. Leafy young twigs 

Oil of Cedarwood Oil from wood 

Unofficial drug 

Sabina Tops 

Pix Burgundica Resinous exudate 

Terebinthina Liquid oleoresin 

Canadensis 

Sandaraca Resinous exudate 

Dammar Resinous exudate 

Succinum (Amber) Fossil resin 

Bordeaux Concrete oleoresin 

Turpentine 

Pix Canadensis Oleoresin 

Oregon Balsam Oleoresin 

Spruce Gum Gum 

TAXONOMY 295 

Botanical origin 

Pinus palustris 

and other species 

of Pinus 



of Pinus 



of Pinus 

-Pinus palustris 


of Pinus 

Pinus Strobus 

Pinus montana 

Larix europsea 

Juniperus | 

communis 

Juniperus 

communis . 

J 

I North 

Habitat 

Southern United 

States 


States 


States 


States 

United States and 

Canada 

Tyrolese Alps 

Alps and 

Carpathians 

America, 

Europe and Asia 

Juniperus Oxycedrus So. Europe 

Thuja occidentalis United States and 

Juniperus 

Virginiana 

Canada 

North America 

Juniperus Sabina Europe 

Abies excelsa Europe and Asia 

Abies balsamea 'Northern United 

States and Canada 

Callitris 

quadrivalvis 

Agathis 

loranthifolia 

Pinites succinifer 

Pinus maritima 

Africa 

E. India 

Basin of Baltic 

France 

Tsuga canadensis North America 

Pseudotsuga 

mucronata 

Picea canadensis 

Picea mariana 

Picea rubra

296 


SUBDIVISION II ANGIOSPER1VLE OR ANGIOSPERMS 

(Plants yrith covered seeds) 

CLASS A. MONOCOTYLEDONEÊ 

A class of Angiosperms characterized by the following peculi- 

arities: 

FIG. 162. Morphology of the typical monocotyledonous plai-t. A, leaf, 

parallel-veined; B, portion of stem, showing irregular distribution of vascular 

bundles; C, ground plan of flower (the parts in 3's) ; D, top view of flower; E, seed, 

showing monocotyledonous embryo. (Gager.) 

One cotyledon or seed leaf in the embryo. 

Stems endogenous with closed collateral or concentric fibro-vascu- 

lar bundles, which are scattered. 

Leaves generally parallel veined.

TAXONOMY 297 

PIG. 163. Wheat plant showing the general habit of grasses. (Rabbins.)

298 


Flowers trimerous (having the parts 

multiple thereof). 

Secondary growth in roots generally absent. 

Medullary rays generally absent. 

of each whorl in threes or 

PIG. 164. Pistillate and staminate inflorescences of corn (Zea mays). 

(Robbins.) 

I. Order Graminales. Graminea or Grass Family. Mostly 

herbs with cylindric, hollow jointed stems whose nodes are swollen. 

The leaves are alternate, with long split sheaths and a ligule. Flow-

TAXONOMY 299 

ers generally hermaphroditic and borne in spikelets, making 

up a spicate inflorescence. Lowest floral leaves of each spikelet are 

called glumes, which are empty and paired. Fruit, a caryopsis or 

grain. Embryo with scutellum. Seeds, albuminous. Seed coat 

fused with fruit coat to form one layer. 

Official drug 

Triticum 

Saccharum 

Maltum 

Amylum 

Zea N.F. 

Part used Botanical name Habitat 

Rhizome and roots Agropyron repens Europe and Asia 

Refined sugar

300 

Official drug 

Sabal 

Unofficial 

Dragon's Blood 

Cocoanut oil 

Carnauba wax 

Areca nut 

Palm oil 


Part used 

Fruit (drupe) 

Inspissated juice 

Fixed oil 

Wax from leaves 

Seed 

Fixed oil 

Botanical name 

-Serenoa serrulata 

Calamus Draco 

Cocos nucifera 

Copernicia cerifera 

Areca Catechu 

Elaeis guineensis 

Habitat 

South Carolina to 

Florida 

East Indies 

Tropics 

Brazil 

Asia and East 

Indies 

West Africa 

FIG. 165. Sabal palmetto. This palm, which appears in the center of the 

figure, yields the official drug, sabal. In the right distance a barragona palm. 

Cuba. (Gager.) 

III. Order Arales. Aracea or Arum Family. Perennial herbs 

with fleshy rhizomes or corms, and long petioled leaves, containing 

an acrid or pungent juice. Flowers crowded on a spadix, which is 

usually surrounded by a spathe. Fruit a berry. Seeds with large 

fleshy embryo.

Unofficial drug Part used Botanical name 

Calamus Unpeeled rhizome Acorus calamus 

Skunk cabbage 

Indian turnip 

Rhizome 

Corm; 

TAXONOMY 301 

Symplocarpus 

foetidus 

Arissema 

triphyllum 

FiC. 166. Acorus calamus. (Sayre.) 

Habitat 

Europe, Asia, 

North America 

North America 

North America 

IV. Order Liliales. Liliacea or Lily Family. Herbs (Lilium), 

shrubs (Yucca), or trees (Dracena Draco), with regular and sym- 

metrical almost always six-androus flowers. Stem either short,

302 


creeping underground (Polygonatum), or, swelling up and forming 

bulbs (Hyacinth), or corms (Colchicum), or, stem may elongate 

above ground and become wiry and herbaceous or semi-shrubby as 

Smilax, or the stem may remain short giving rise to thick fleshy and 

sap-storing leaves as in Aloe. Leaves linear to lanceolate, ovate 

rarely wider, divisible into sheathing base, narrow petiole and ex- 

PIG. 167. Diagram of A, lily flower, and B, grass flower, showing homologous 

structures. A, f, bract; ax, axis; op, outer perianth; ip, inner perianth; s, stamens; 

(c) tricarpellary ovary. B, shaded structures are aborted; le, lemma 

(bract); ax, axis; p and p f 

, palet (outer perianth); / and I', lodicules (inner peri- 

anth); ^ and s', two whorls of stamens; c, tricarpellary ovary. (A, Robbins. B, 

after Shuster.) 

panded blade. Venation, parallel, becoming in some ovate leaves 

parallel with oblique connections, reticulate or highly reticulate as 

in Smilax, etc. The perianth is parted into six segments, the calyx 

and corolla being alike in color. Anthers introrse. Ovary three- 

locular with a single style. 

B 

Fruit a three-Jocular, loculicidally de- 

hiscent capsule (Lilium, etc.) or rarely a berry (Asparagus, etc.). 

Seeds usually numerous, albuminous.

Official drug 

TAXONOMY 33


Iridacea or Iris Family. Perennial herbs with equitant two- 

ranked leaves and regular or irregular flowers which are showy. 

Fruit a three-celled, loculicidal, many-seeded capsule. Rootstocks, 

tubers, or corms mostly acrid. 

Official drug

TAXONOMY 305 

VI. Order Orchidales. Orchidacece or Orchid Family. Perennial 

herbs of terrestrial or terrestrial saprophytic or epiphytic growth, 

FIG. 168. Smilax officinalis Portion of vine and rhizome. (Sayre.) 

having grotesque flowers. Roots fibrous or tuberous often saprophytic 

in relation, or aerial and with velamen. 

20 

Stems and branches

306 


upright, in epiphytic types, often forming pseudobulbs. Leaves 

alternate, entire, parallel-veined, sheathing at base, rarely reduced 

to yellowish or pale scales in saprophytes. Flowers irregular, usu- 

FIG. 169. Zingiber officinale. (Sayre.) 

ally attractive, entomophilous, arranged in elongated spikes or 

racemes, trimerous. Sepals, three usually similar; petals three of 

which two often resemble sepals, third is variously, often greatly

TAXONOMY 307 

modified and fused with two outer petaloid stamens as a labellum 

or lighting-board for insects. Third stamen of outer whorl fertile 

(Orchidea) or a barren knob (Cypripedice) ; pollen of fertile anther agglutinate 

as pollinia. Three stamens of inner circle barren and 

petaloid or one absent (Cypripedia). Stamens all epigynous and 

FIG. 170. Floral organs of an orchid (Cattleya sp.). A, the entire flower; sep, 

sepal; pet, petal; B, column, showir.g s, stigma and r, the rostellum (beak), with 

the small glands at the tip; to the glands are attached the four strap-shaped 

caudicles of the pollinia; C, pollinia, with the four caudicles; below, the gland; D, 

longitudinal section of the column; p, pollinium; E, the same, enlarged. (Gager.) 

often three are fused with the style as gynandrium. Carpels three, 

syncarpous, with inferior, three rarely four, one (usually)- celled 

ovary. Fruit a capsule, three-valved and one-celled. Seeds minute, 

abundant and wind disseminated. 

p 

r

308 


Official drug Part used Botanical name Habitat 

Vanilla N.F. Fruit Vanilla Planifolia Mexico 

Cypripedium 

Cypripedium N.F. Rhizome and roots , United States 

Cypripedium 

parviflorum 

FIG. 171. Vanilla planifolia Branch showing leaves and flowers. (Sayre.) 

CLASS B. DICOTYLEDONE^ 

Plants having the following characteristics: 

Two-seed leaves (cotyledons) in embryo. 

Netted-veined leaves.

TAXONOMY 309 

Stems, and roots of secondary growth with open collateral fibrovascular 

bundles, radially arranged about pith. 

Exogenous stems. 

Medullary rays present. 

Cambium. 

PIG. 172. Morphology of a typical dicotyledonous plant. A, leaf, pinnatelynetted 

veined; B, portion of stem, showing concentric layers of wood; C, groundplan 

of flower (the parts in 5's); D, perspective of flower; E, longitudinal section 

of seed, showing dicotyledonous embryo. (Gager.) 

Roots developing secondary structure. 

Flowers tetra- or pentamerous (parts of each whorl, four or five or 

multiple thereof). 

SUB-CLASS A. ARCHICHLAMYDEÊ 

Those dicotyledonous plants in which the petals are distinct and 

separate from one another or are entirely wanting. That group of 

> 

-, 

' 

:

3 io PHARMACEUTICAL BOTANY 

the Archichlamydeae whose flowers show the absence of petals and 

frequently of sepals is called the Apetalae. The group whose plants 

have flowers showing the parts of their corolla (petals) separate and 

distinct is called the Choripetalae. 

FIG. 173. Piper cubeba Fruiting branch and fruit (enlarged). (Sayre.) 

I. Order Piperales. Piperacece or Pepper Family. A family of 

aromatic herbs and shrubs with jointed stems, opposite,verticillate, 

or sometimes alternate leaves without stipules, and spiked incon- 

spicuous, wind-pollinated flowers. The characteristic fruit is a berry

TAXONOMY 

enclosing a 'single upright seed with abundant perisperm (from 

megasporangial 

embryo sac). 

tissue) and reduced endosperm (from matured 


Cubeba 

Piper 

Kava N.F. 

Matico N.F. 

Unripe fruit 

Piper Cubeba 

Borneo, Java 

Sumatra 

Unripe fruit Piper nigrum Cochin- China, 

India 

Rhizome and roots Piper methysticum South Sea Islands 

Leaves Piper angustifolium Peru, Bolivia 

PIG. 174. Willow (Salix). Leafy branch, bearing two pistillate catkins, 

Staminate flower above, at the left; pistillate flower below, at the right. (Gager 

after Britton and Brown.) 

II. Order Salicales. Salicacece or Willow Family. Shrubs or 

trees of temperate or cold regions, with upright woody stems. Bark 

often containing bitter principles (Salicin etc.). 

^Leaves alternate, simple, entire, stipulate; stipules rarely green, 

persistent, usually functioning as winter bud-scales and falling in 

spring.

312 


Inflorescences dioecious spikes, so on separate plants. 

Staminate 

spikes forming deciduous catkins of yellowish flowers, pistillate 

as persistent spikes of green flowers, at length maturing fruit. 

Flowers of catkins numerous, each of two to five (Willow) or 

six to fifteen (Poplar) stamens in axil of a small bract leaf, sometimes 

with small nectar knob or girdle at base; pollen abundant, hence 

plants anemophilous, rarely entomophilous. Pistillate flowers 

green, each of a bicarpellate pistil in axil of bract, ovary one-celled 

with parietal placentation, style simple, stigma bilobed. 

Fruit a capsule dehiscing longitudinally. Seeds small, exal- 

buminous, surrounded by a tuft of hairs for dissemination. 


Salicin Glucoside Several species of Europe, North 

Salix and Populus America 

nigra 

Populus > North America\ 

balsamifera 

{Populus 


Salix Bark Salix alba Europe 

III. Order Myricales. Myricacecz or Bayberry Family. Dice- 

cious or sometimes monoecious, aromatic shrubs or trees with watery 

juice and possessing underground branches which arch downward 

then upward producing many suckers. Roots fibrous and bearing 

many short rootlets upon which are frequently found coralloid 

clusters of tubercles containing the Actinomyces Myricarum Young- 

ken. Leaves alternate, revolute in vernation, serrate, irregularly 

dentate, lobed or entire, rarely pinnatifid, pinnately and reticulately 

veined, pellucid punctate, evergreen or deciduous, generally exstipulate, 

rarely stipulate. Flowers naked, unisexual, monoecious 

or dioecious, in the axils of unisexual or androgynous aments from 

scaly buds formed in the summer in the axils of the leaves of the 

year, remaining covered during the winter and opening in March or 

April before or with the unfolding of the leaves of the year. 

Staminate flowers in elongated catkins, each consisting of two to 

eight stamens inserted on the torus- like base of the oval to oval- 

lanceolate bracts of the catkin, usually subtended by two or fotir 

or rarely by numerous bracteoles; filaments short or elongated,

TAXONOMY 313 

filiform, free or connate at the base into a short stipe; anthers ovoid, 

erect, two celled, extrorse, showing longitudinal dehiscence. Pistil- 

late flowers in ovoid or ovoid-globular catkins. Gynsecium of two 

united carpels on a bract. Ovary sessile, unicellular, subtended 

by two lateral bracteoles which persist under the fruit, or by eight 

PIG. 175. Two Myrica cerifera trees growing in a field near a brackish swamp 

at Rio Grande, N. J. Photographed by author July 26, 1914. 

linear subulate bracteoles, accrescent, and forming a laciniate 

involucre inclosing the fruit; styles short and dividing into two 

elongated style arms which bear stigmatic surfaces on their inner 

face; ovule orthotropous, solitary, with a basilar placenta and 

superior micropyle. Fruit an akene or ceriferous nut. Pericarp

314 


covered with glandular emergences which' secrete wax or fleshy 

emergences, smooth and lustrous or smooth, glandular. Seed 

FIG. 176. Fructiferous branches of Myrica Caroliniensis (to left) and Myrica 

cerifera (to right) as they appear in early winter. The former species is deciduous, 

while the latter is evergreen. Collected by author at Wildwood, N. J., Dec. 16, 

1914. 

erect, exalbuminous, covered with a thin testa. Embryo straight, 

cotyledons thick, plano-convex; radicle short, superior. 

There are two distinct genera of this family, e.g., Myrica and 

Comptonia.

Official drug 

Myrica N.F. 

Part used 

Bark of root 

TAXONOMY 315 


Myrica cerifera 

Habitat 

Eastern United 

States 

FIG. 177. Fructiferous branch of Myrica Gale, showing mature pistillate 

catkins and nature of leaves. Collected at the south end of Peak's Island in 

Casco Bay, Maine, September, 1913. 

IV. Order Juglandales. JUGLANDACEÊ. A family of apetalous 

exogenous trees the walnut family with alternate odd-pinnate 

leaves and monoecious flowers, the sterile in catkins, the fertile 

solitary or in a small cluster or spike. The fruit is a dry drupe with

316 

PHARMACEUTICAL BOTANY* 

a bony nut-shell and a four-lobed seed. It embraces five genera, 

of which Carya (Hicoria) and Juglans are represented in the United 

States, and about 35 species. 


Juglans N. F. Inner root bark Juglans cineiea United States 

Betulacea or Birch Family. A family of aromatic trees or shrubs 

distinguished by monoecious flowers with scaly bracts and astringent 

resinous bark. Differs from Fagacea by superior ovary and absence 

of cupule. To this family belong the hazelnuts, birches, alders, the 

ironwood, and the hornbean. 


(Oleum Betulae) Volatile oil Betula lenta North America 

Methylis salicylas 

V. Order Fagales. Fagacece or Beech Family (Cupuliferce) . 

Beeches, Chestnuts, Oaks, the trees of this family are found in the 

temperate forests of the eastern and western hemispheres and comprise 

about 368 species. North America has over 50 species of oaks; 

2 species of Chestnuts; i species of beech and'i species of golden- 

leaved chestnut. The most important American oaks used for 

building, for furniture, for cooperage, for wagons, for tanning leather 

etc. are white oak, Quercus alba; chestnut oak, Q. prinus, black oak, 

Q. velutina] live oak, Q. mrginiana\ swamp white oak, Q. platanoides', 

cow oak, Q. Michauxii, and the two Pacific coast oaks, Q. chyrso- 

lepis and post oak, Q. garryana. The uses of the fast disappearing 

Chestnut, Castanea dentata, are well known. The wood of the beech, 

Fagus grandifolia, is used for chairs, tool handles, plane stocks, shoe 

lasts and for fuel. The nuts (mast) fatten hogs and feed wild ani- 

mals and birds. The cork of commerce is obtained from the bark of 

Quercus Suber and Quercus occidentalis , plants indigenous to Spain 

and France. 

The above trees are all monoecious, that is the staminate (male) 

and pistillate (female) flowers are distinct from each other, but borne 

on the same tree. Most of the species are trees, a few oaks are 

shrubs. The leaves are simple, netted-veined and alternate. A 

pair of deciduous stipules are found at the base of the leaf-stalk

TAXONOMY 317 

(petiole). The margins of the chestnut and beech leaves are sharply 

cut with large teeth. The leaves of the oaks divide the genus into 

two groups, 

viz. the white oaks with rounded lobed leaves and 

PIG. 178. Quercus infectoria Branch and nutgall. (Sayre.) 

annual acorn production, and the 'black oaks with sharp bristle- 

tipped lobes and biennial acorn production. The male flowers are 

in dangling heads (beech), or in catkins (chestnut and oaks). The


male flowers have a united perianth, which is 4-6 parted and en^ 

closes an indefinite number of undivided stamens. The female 

FIG. 179. Hop (Humulus lupulus). A, portion of plant showing pistillate 

inflorescences; B, staminate inflorescence; C, rachis of pistillate inflorescence 

("hop"). (Robbins.) 

flowers have a superior 6-parted perianth; the pistil consisting of 3 

carpels with a corresponding number of stigmas. The ovary is

TAXONOMY 319 

3-6 celled and each cell has 2 pendulous ovules. The fruit is a oneseeded 

nut. The cup, or cupule, in the beech is 4 sided and covered 

externally wth weak spines and encloses two 3 sided seeds. The 

cupule in the chestnut forms the spiny bur, which splits into 4 valves 

at maturity, enclosing 3 nuts. The cupule. in the oak is saucer, or 

cup-shaped, and encloses a single rounded nut, or acorn. The seeds 

are exalbuminous and the cotyledons are thick and fleshy, edible in 

the beech, chestnut and a few of the oaks. 

O facial drug 

Part used 

Galla Excrescence 

Castanea N.F. Leaves 

Quercus N.F. Bark


flowers and oily seeds. Many are parasitic 

plants. 

Official drug Part used 

Oleum Santali Volatile oil 

Santalum Album N.F. Heartwood 

on the roots of other 

Botanical name Habitat 

Santalum album 

FIG. 1 80. Aristolochia serpentaria. (Sayre.) 

f India and East 

I Indies 

VIII. Order Aristolochiales. A ristolochiacea or Birthwort Family. 

Herbs or twining semi-woody or woody plants, having more or less 

swollen nodes from which spring cordate or reniform or ovate leaves. 

Flowers regular (Asarum, etc.) or irregular (Aristolochia) often 

offensively smelling. Fruit a capsule. Seeds with copious albumen 

and minute embryo.

Official drug 

Serpentaria 

Asarum N.F. 

Part used 

Rhizome and roots 

TAXONOMY 3 2I 


Aristolochia 

Serpentaria 

Aristolochia 

reticulata 

Habitat 

United States 

Rhizome and roots Asarum canadensis United States 

FIG. 181. Serpentaria Cross-section of rhizome. (25 diam.) A, parenchyma 

of cortex; B, medullary ray; C, xylem; D, phloem; E, medulla. (Sayre.) 

IX. Order Polygonales. Polygonacea Family. Usually herbs 

(Polygonum, Rumex, etc.) rarely trees (Coccoloba uvifera and C. 

platyclada) or shrubs (Muhlenbeckia, Brunnichia) having strong 

vertical tap roots and spreading secondary roots more or less pro- 

vided with tannin compounds. Stems elongate, green, to woody, 

rarely flattened, leathery, phylloidal (Muhlenbeckia platyclada} still 

more rarely tendriliform (Antigonum leptopus). Leaves alternate 

rarely opposite or whorled (Eriogonum), entire, rarely lobed (Rheum 

palmatum, Rumex acetosella), petiolate, rarely sessile, and stipu- 

late. Stipules fused and forming a greenish membranous upgrowth 

(ocrea) which sheaths the stem. Inflorescence racemose with 

many dense scorpioid or helicoid cymes, which in some forms condense 

into single flowers. Flowers regular, pentamerous, with 

simple calyx, becoming trimerous with two whorls of three sepals 

each. Stamens varying from fifteen or twelve to nine or six more 

rarely to five, four, three to one (Kcenigia), hypogynous, more 

21

322 


rarely by enlargement of receptacle and slight fusion of sepals, 

perigynous. Pistil tri- to bicarpellate often three- to two-sided, 

ovary one-celled with one ovule. Styles .three, rarely two, radiating 

penicillate in wind-pollinated inconspicuous flowers, becoming condensed 

knob-like in conspicuous insect-pollinated flowers. Fruit a 

triangular or biconvex akene often crowned by persistent styles and 

surrounded by persistent closely applied sepals. Seeds solitary, 

albuminous, with straight embryo, or in Rumex, curved embryo. 

PIG. 182. Rhubarb (Rheum) flower, external view, median lengthwise section, 

and with perianth and stamens removed. (Robbins, after Lurssen.) 

Official drug 

Rheum Rhizome 

Rumex N.F. Root 

Unofficial 

Bistorta Rhizome 

Part used 


Rheum officinale 

Rheum palmatum 

and the variety 

tanguticum and 

probably other 

species 

Rumex crispus 

Rumex 

obtusifolius 

Polygonum 

Bistorta 

Habitat 

China and 

Thibet 

Europe 


X. Order Chenopodiales (Centrosperma). Chenopodiacea or 

Goosefoot Family. Usually herbaceous halophytes or shore growers,

TAXONOMY 3 2 3 

growing also in any alkaline soil, more rarely shrubs (Atriplex) or 

low trees (Haloxylon). 

Among them are several garden vegetables (Spinach, Beets, 

Mangels) and a number of weeds. Leaves alternate to opposite 

sometimes reduced to teeth, entire or lobed. Inflorescence spikes 

FIG. 183. Phytolacca decandra (in foreground), growing in damp woodland. 

or short racemes of condensed cymes. Flowers regular, usually 

small and greenish. They are either perfect (Beta), monoecious 

(Chenopodium), dioscious (Atriplex sp.), or polygamous (Kochia). 

Fruit a utricle. Seed albuminous.

324 



Oleum Chenopodii Volatile oil Chenopodium United States 

anthelminticum 

Saccharum Refined sugar Beta vulgaris Europe 

Unofficial 

Chenopodium Fruit Chenopodium United States 

anthelminticum 

Phytolaccacea. A family of apetalous trees, shrubs, or woody 

herbs the pokeweed family vvith alternate entire leaves, and 

flowers resembling those of the goosefoot family (Chenopodiacea) , 

but differing in having the several-celled ovary composed of carpels 

united in a ring, and forming a berry in fruit. It embraces 2 1 genera, 

and 55 species, tropical and sub-tropical. 


Phytolacca N.F. Root Phytolacca North America 

decandra 

XI. Order Ranales. Magnoliacece or Magnolia Family. Trees 

and shrubs having alternate leaves and single large flowers with 

calyx and corolla colored alike. Sepals and petals deciduous, an- 

thers adnate. Carpels and stamens numerous. Bark aromatic 

and bitter. Fruit a collection of follicles dehiscing dorsally. 

Official drug 

Oleum anisi 

Unofficial 

Winter's Bark

TAXONOMY 3 2 5 

(Columbine, Larkspur, Peony) or rarely a berry as in Baneberry 

(Actad). Seeds albuminous. 

FIG. 184. Above ground portion of AconUum Napellus showing palmatelydivided 

leaves and hooded flowers. 

Official drug 

Hydrastis 

Aconitum 

Staphisagria 

Cimicifuga 


Rhizome and roots Hydrastis canaden- Eastern United 

sis States- and Canada 

Tuberous root Aconitum 

Napellus 

Seed Delphinium 

Staphisagria 

Rhizome and root Cimicifuga 

racemosa 

Europe, Asia, 

Western North 

America 

South Europe, 

Asia Minor 

North America, 

Europe, Asia

326 


PIG. 185. Cimicifuga racemosa Plant and rhizome. Note the pinnately decompound 

leaf and the wand-like racemes which bear white flowers. (Sayre.)

Official drug 

Part used 

Pulsatilla N.F. Entire herb 

Coptis N.F. 

Adonis N.F. 

.Entire herb 

Entire herb 

Delphinium N.F. Seeds 

Unofficial 

Aconiti Folia 

Helleborus 

TAXONOMY 327 


Anemone 

Pulsatilla 

Anemone Europe 

Ludoviciana 

Anemone pratensis 

Coptis trifolia United States and 

Canada 

Adonis vernalis Europe 

Delphinium 

Consolida 

Europe 

Delphinium 

Ajacis 

Leaves and flower Aconitum 

tops Napellus 

Rhizome and roots Helleborus niger 

Europe, Asia, 

western North 

America 

Alps 

Berberidacea or Barberry Family. Herbs and woody plants with 

watery juices and alternate, or radical, simple or compound leaves 

often bearing spines or barbs, which give them a barbed appearance. 

Fruit a berry or capsule. 


Berberis N.F. Rhizome and roots Berberis species of Western North 

the Sect. 

Odostemon 

America 

Podophyllum Rhizome and roots Podophyllum Eastern North 

peltatum 

America 

Caulophyllum Rhizome and roots Caulophyllum Eastern United 

N.F. thalictroides States 

Menispermacea, or Moonseed Family. Choripetalous woody, 

climbing, tropical plants with alternate, exstipulate, simple often 

peltate leaves. Flowers green to white. Fruit a one-seeded succu- 

lent drupe. Seeds albuminous. They usually contain tonic, nar- 

cotic or poisonous bitter principles.

328 


PIG. 1 86. Podophyllum peltatum. Entire plant, above ground portion, and 

fruit. Note the flowering sterns bearing in each instance two one-sided leaves 

and a nodding flower from the forks. This plant also sends up from its rhizome 

flowerless stems each of which terminates in a 7 -9 lobed peltate leaf. 


Calumba Root Jateorhiza East Africa 

Pareira N.F. Root 

palmata 

Chondodendron 

tomentosum 

Peru and Brazil 

Cocculus N.F. Fruit 

' 

Anamirta cocculus Asia 

Unofficial 

Menispermum Rhizome and roots Menispermum 

canadense 


Canada

TAXONOMY 329 

Myristicacea or Nutmeg Family. An order of apetalous trees 

comprising the single genus Myristica, of about 80 species. 

Myristica. A large tropical genus of fragrant, apetlaous trees 

the nutmegs coextensive with the nutmeg family, having alternate, 

PIG. 187. Jateorhiza palmata Portion of vine. Note tendril for winding 

about a support as this plant climbs. (Sayre.) 

entire, often punctate leaves, small dicceious regular flowers, and a 

succulent, two-valved, one-celled fruit with a solitary seed, usually 

covered by a fleshy arillus. 

M. fragrans, a handsome tree, 20 to 30 feet high, of the Malay 

archipelago, supplies the nutmegs and mace of commerce.

330 


Myristica 

Kernel of seed 

Oleum Myristicae Volatile oil 

Macis N.F. Arillode 



Myristica fragrans 



PIG. 1 88. Myristica fragrans Branch, and fruit. (Sayre.) 

Habitat 

Molluca Islands 

Lauracecz or Spicebush Family. Shrubs or trees of sub-tropical or 

tropical, rarely of temperate regions. Bark, wood and leaves rich 

in spicy aromatic , 

hydrocarbon 

oils. Leaves alternate, simple, 

entire, often leathery shining. Inflorescenes usually cymose clus-

TAXONOMY 331 

ters. Flowers small, green, yellow or rarely whitish, hermaphrodite 

or more or less dioecious, regular calyx alone present as a floral whorl, 

of three to six small sepals. Stamens four to twelve in several rows 

of three to four each; anthers opening by recurved valves (valvular 

PIG. 189. Cinnamomum zeylanicum Branch. (Sayre.) 

dehiscence). Pistil monocarpilary, ovary superior one-celled with 

solitary pendulous ovule, style simple with usually rounded stigma. 

Fruit a succulent berry (Spicebush and "Bacca laurea") or a suc- 

culent drupe (Sassafras).

332 

Official drug 

Camphora 

Ketone 


Part used 

Sassafras Bark of root 

Sassafras Medulla Pith 

N.F. 

Cinnamomum Bark 

Zeylanicum 

' Cinnamomum Bark 

Saigonicum 

Oleum Cinnamomi Volatile oil 

Unofficial 


Cinnamomum 

Camphora 

Sassafras 

variifolium 

Sassafras 

variifolium 

Cinnamomum 

Zeylanicum 

Undetermined 

species of 

cinnamon 

Cinnamomum 

cassia 

Habitat 

Eastern Asia 

North America 

North America 

Ceylon 

China 

China

TAXONOMY 333 

numerous, anatropous, Fruit a capsule except in Platystemon 

which has follicles. Seeds richly albuminous. 

Opium 

FIG. 190. Papaver Somniferum Flowering branch and fruit. (Sayre.) 

Official drug Part used Botanical name 

Sanguinaria 

Unofficial 

Chelidonium 

Maw seed 

Air-dried milky Papaver somni- 

exudate ferum and its 

var. album 

Rhizome and roots Sanguinaria 

canadensis 

Entire flowering Chelidonium 

plant 

majus 

Seeds Papaver somniferum 

album 

: 

Habitat 

Eastern Mediter- 

ranean countries 


Canada 


Canada 

vide supra

334 


Fumariacea or Fumitory Family. Delicate herbs rarely shrubs 

containing milky watery to watery latex. Leaves more or less 

compound. Inflorescence a raceme or spike. Flowers irregular, 

zygomprphic, one or both of the petals of which having a spur. 

Fruit a one-chambered capsule. Seeds albuminous. Idioblasts 

common. 

FIG. 191. Transverse 

section of flower of 

Poppy. (Sayre.) 

PIG. 192. Gynecium 

of Poppy, with 

one stamen remaining. 

(Sayre.) 


Corydalis N.F. < canadensis 

f Tubers of Dicentra (Bicuculla) 

i Bulbs 

of Dicentra Cucullaria 

PIG. 193. Transverse 

section of ovary 

of Poppy. (Sayre.) 

Habitat 

United-Spates and 

Canada \ 

Crucifera or Mustard Family. Herbs, rarely shrubs, -mostly of 

temperate regions. Stem and branches upright or diffuse spreading 

(Arabis). Leaves alternate, simple rarely compound, exstipulate, 

entire or toothed, often more or less hairy. Inflorescence at first 

a corymb or shortened raceme, later elongating into a loose raceme. 

Bracts at base rarely reduced, usually absent. Flowers regular 

rarely irregular (Candytuft) tetramerous. Sepals four, green, 

equal or two laterals, at times pouched as nectar receptacles. Petals 

four, yellow to white or to pink or purple, cruciform, often divisible 

into claw and blade. Stamens six to four long anteroposterior, 

two short lateral and often with nectar knobs or discs, hence termed 

tetradynamous, insertion hypogynous. Pistil syncarpous bicarpellate, 

superior, carpels lateral. Ovary one-celled but . falsely 

two-celled by a placental replum, style simple, stigma rounded or 

bifid or bilobed. Ovules several, rarely few, attached to marginal

TAXONOMY 335 

placenta. Fruit a capsule rarely indehiscent bursting lengthwise 

by two valves. Seeds exalbuminous. 

Official drug 

Sinapis Alba Seed 

Sinapis Nigra 


Seed 

Semenae Rapae Seed 


Sinapis alba Europe and Asia 

Brassica nigra Europe and Asia 

Brassica napus Europe and Asia 

PIG. 194. Brassica nigra Branch. (Sayre.) 

XIII. Order Sarraceniales. Droseracea or Sundew Family 

Herbs (Drosera, Dionaea, etc.) rarely shrubs (Roridula of South 

Africa), growing in bogs or swamps or purely aquatic in habit 

(Aldrovanda). Leaves, either rosettes or more or less scattered in 

alternate fashion over stem, usually glandular-hairy or sensitive- 

hairy and insectivorous. Inflorescence a loose raceme or cymose

336 


umbel. Flowers regular, pentamerous; sepals five, aposepalous 

green; petals apopetalous, varying from white to whitish-pink, 

pink scarlet, purple to bluish-purple; stamens varying from fifteen 

to five hypogymous pistil syncarpous of five to three, rarely fewer 

carpels. Fruit a capsule. Seeds albuminous. 

PIG. 195. Drosera rotundifolia. Production of a young plant from the leaf of an 

older plant. (Gager.) 

Official drug 

Part used 

Drosera N.F. Entire plant 


Drosera rotundifolia 

Drosera intermedia 

Drosera longifolia 

Habitat 

Eastern and west- 

ern Hemispheres 

XIV. Order Resales. Saxifragacece or Saxifrage Family. 

Herbs (Saxifraga, Mitella, etc.) or shrubs (Hydrangea, Ribes, 

Philadelphus, etc.) rich in tannin content, with opposite or alter- 

x 

nate leaves usually devoid of stipules. Both stamens and petals are 

generally inserted on the calyx. Fruit a follicle, capsule, or berry. 

Seeds with copious albumen.

TAXONOMY 337 


Hydrangea N.F. Rhizome and roots Hydrangea United States 

arborescens 

Hamamelidacce or Witchazel Family. Shrubs or small or large 

trees. Leaves, simple, alternate and pinnately veined; stipules 

deciduous to caducous, paired and slightly fused at the bases of 

petioles. 

FIG. 196. Liquidambar orientalis Branch. (Sayre.) 

Flowers frequently yellow to yellowish-white, in axillary 

clusters or heads or spikes, hermaphrodite or monoecious; sepals 

and petals five to four rarely indefinite, superior (petals absent in 

Fothergilla) ; 

stamens twice as many as the petals but the outer row 

alone fertile, the inner row being more or less barren and scale-like; 

gyncecium of two carpels united below. Fruit a two-beaked, two- 

22


PIG. 197. Hamamelis virginiana. Upper figure shows this shrub as it appears 

in autumn alter the leaves have fallen. Note that the plant is in blossom. 

Lower figure shows a flowering branch from the same plant. The bright yellow 

flowers occur in axillary clusters appearing at the same time as the ripening of 

fruits from blossoms of the previous year.

TAXONOMY 339 

celled woody capsule dehiscing at the summit, with a bony seed 

in each cell, or several, only one or two of them ripening. 

Official drug

340 



Rubi Fructus Fresh fruit Rubus nigrobaccus United States 

N.F. and Rubus 

villosus 

Succus Pomorum Juice of Fruit Pyrus Malus Cultivated 

N.F. (cyltiv. var's.) 

Rubi Idsei Juice of Fruit Rubus Idseus and Europe and Asia 

Fructus N.F. Rubus strigosus 

Prunum N.F. Fruit Prunus domestica Southern Europe 

Amygdala Amara Seed Prunus amygdalus Asia Minor, Persia, 

var. amara. Syria 

FIG. 198. Quillaja saponaria Branch. (Sayre.)


Part used 

Cydonium Seed 

Rosa Centifolia Petals 

Rosa Canina Spurious Fruit 

Tormentilla Rhizome 

TAXONOMY 341 


Cydonia vulgaris 

Rosa centifolia 

Rosa canina 

Potentilla 

silvestris 

Habitat 

Cultivated widely 

Western Asia 

Europe 


FIG. 199. Prunus domestica Fruiting branch and flowering branch. (Sayre.) 

Leguminosce or Pea Family (Fabacece). Herbs, 

shrubs or trees 

of all regions, with tubercled roots. Stem usually erect, rarely 

creeping (Trifolium repens). Leaves alternate, compound rarely 

simple stipulate, sometimes tendriliform or reduced to phylloid 

petioles (Acacia sp.). Inflorescence a raceme, at times, condensed 

almost to a head or capitulum (Sp. of clover, Mimosa, etc.). Flowers 

pentamerous (rarely four), regular (Mimosea), to irregular (C&sal- 

pinea, Papilionacece). Sepals five united, green; petals five (rarely 

four) variously related, in Papilionaceae one superior, external, posterior 

standard or vexillum, two lateral forms wings or alae, two 

inferior internal and anterior slightly adherent form keel. Stamens

342 


ten to four, free or in Papilionaceae united by filaments into a mona- 

delphous (ten) or a diadelphous (nine to one) tube, inserted peri- 

gynously. Pistil monocarpellary, ovary with sutural placentation, 

style simple. Seeds exalbuminous. 

FIG. 200. Glycyrrhiza glabra Branch. (Sayre.) 


Acacia Gummy exudation Acacia Senegal 

and other African 

species 

Tragacantha Gummy exudation Astragalus gummifer 

and other 

Asiatic species 

Africa 

Asia 

Habitat

Official drug 

Balsamum 

Peruvianum 

Balsamum 

Tolutanum 

Hsematoxylon 

N.F. 

Santalum Rubrum 

Glycyrrhiza 

Senna 

Cassia Fistula 

N.F. 

TAXONOMY 343 


Balsam Toluifera Pereirae Central America 

Balsam Toluifera Columbia 

Balsamum 

Heartwood Hsematoxylon Central America 

campechianum 

Heartwood Pterocarpus Indo China 

santalinus 

and France 

Glycyrrhiza glabra | Spain 

Rhizome and roots 

^ 

Glycyrrhiza Southwestern 

Leaflets 

I glandulifera 

Asia, Russia 

( Cassia acutifolia Egypt 

I Cassia angustifolia Arabia and India 

Fruit Cathartocarpus India 

fistula 

FIG. 201. Cassia acutifolia Branch showing flower and fruit. (Sayre.)

344 

Official drag 

PHARMACEUTICAL BOTANY

TAXONOMY 345 

to irregular pentamerous (Pelargonium)', sepals five, aposepalous; 

petals five, apopetalous, varying in color from greenish-white or 

pink-red to scarlet, scarlet-crimson to crimson-purple; stamens with 

anthers ten or five, hypogynous or inserted into slightly developed 

hypogynous disc; pistil pentacarpellary, ovary five-celled with two, 

FIG. 202. Physostigma venenosum Portion of plant and fruit. (Sayre.) 

rarely one ovule in each cell, styles elongate, fused round a stylar 

column of receptacle then continued as a stylar tip which splits into 

five stigmatic surfaces. Fruit a regma rarely a simple capsule. 

Regma splits into five recurved carpels, each then dehiscing to set 

free two or one seeds. Seeds exalbuminous.

346 


Geranium N.F. Rhizome 

Unofficial 

Oil of Rose 

Geranium 


Volatile oil 


Geranium maculatum North America. 

Pelargonium 

odoratissimum 

Pelargonium Radula 

Pelargonium 

capita turn 

PIG. 203. Linum usitatissimum. (Sayre.) 

Mediterranean 

regions 

Linacecs or Flax Family. Herbs with slender stems and alter- 

nate, simple, narrow leaves. Inflorescence cymose with regular 

pentamerous flowers; pistil five-carpelled with a five-celled ovary 

containing two ovules in each cavity and having a single style with 

knob-like stigma. While the flower is still in bud condition or soon 

after, there commences an ingrowth of the mid-rib of each carpel

TAXONOMY 347 

which proceeds until, when plant is in fruit, there are formed 10 cavi- 

ties each enclosing a seed. Seeds, anatropous, mucilaginous, flat- 

tened, containing a large embryo and slight albumen. 


Linum Seeds Linum usitatissimum } Temperate 

Oeum Lini Fixed oil Linum usitatissimum / regions 

FIG. 204. Flax. A, floral diagram c, calyx; co, corolla; s, stamens; p, pistil. 

B, median lengthwise section of flower. C, calyx and corolla removed. D, 

fruit, external view. E, cross-section of fruit. (Robbins.) 

Erythroxylaceo3 or Coca Family. Shrubs (Erythroxylon) or trees 

with alternate, simple, entire, glabrous and pinnately veined leaves. 

Flowers regular, hermaphroditic, each with five sepals, five hypogynous 

petals, ten stamens and a two- to three-celled ovary subtend-

348 


ing three styles, each with a capitate stigma. 

one celled drupe containing a single seed. 

Official drug 

Cocaina 

Coca 

Unofficial 

Fruit an ovoid, angular, 

Part used Botanical origin Habitat 

Alkaloid Erythroxylon Coca Peru and Bolivia 

and its varieties 

Leaves Erythroxylon Coca Peru and Bolivia 

and its varieties 

FIG. 205. Guaiacum sanctum Flowering branch. (Sayre.) 

Zygophyllacece or Caltrop Family. Herbs, shrubs or trees (Guaia- 

cum) having jointed, often divaricate branches. Leaves usually 

opposite, stipulate and compound. Flowers regular or irregular,

TAXONOMY 349 

pentamerous, white, yellow, red or blue (G. o/icinale). Fruit a 

capsule. 

Official drug 

Guaiacum 

Guaiaci Lignum 

N. F. 

Part used 

Resin of wood 

Heartwood 

* 


( Guaiacum Guaij officinale 

I Guau 

Guaiacum sanctum 

G. officinale and G. 

sanctum 

PIG. 206. Citrus Auranlium Branch. (Sayre.) 

Habitat 

Tropical and subtropical 

America 

Tropical and sub- 

tropical America 

Rutacece or Rue Family. Herbs (Ruta, Diosma, Barosma) or 

shrubs (Xanthoxylum) or trees (Citrus). Stems upright, often wiry 

xerophytic, in sub-family Rutece, elongated and spiny in sub-family 

Zanthoxylece, woody and green in sub-family Aurantiea. Leaves 

alternate or oppostite, simple (Ruta), rarely whorled (Pilocarpus) or

350 


pinnatifid, as in Ruta graveolens, or pinnate, as lower parts of Ruta 

graveolens, becoming reduced pinnate in Citrus Aurantium. Leaves 

PIG. 207. Barosma 

Branch and flower. 

Official drug 

betulina 

(Sayre.) 

exstipulate or with spiny stipules 

(Xanthoxylum). Stems and leaves 

abound in more or less sunken 

glands. Flowers pentamerous, varying 

in color from yellow in Ruta to 

white in Citrus to pink (Barosma 

betulina) or pink crimson as in some 

Barosma and Diosma species, rarely 

to pinkish-purple (Pilocarpus} ; 

sepals five, aposepalous becoming in 

Citrus more or less synsepalous; 

petals five, apopetalus becoming 

more or less synpetalous and tubular 

(Correa grandiflora) ; stamens five, 

simple or with expanded bases, 

lobed, or lobes developed as staminal 

stipules and more or less split 

(Citrus}-, pistil of ten, five, three or 

two carpels, ovary as many-celled. 

Fruit a capsule (Dittany, Xanthoxylum}, 

berry (Citrus] or rarely a 

samara (Ptelea). Seeds albuminous 

or exalbuminous. Many of the 

plants contain volatile oils in their 

secretory cavities.

Official drug 

Xanthoxylum 

Bark 

Part used 

Oleum Bergamottae Volatile oil 

N.F. 

Oleum Aurantii Volatile oil 

Florum N.F. 

Succus Citri N.F. Juice 

Xanthoxyli 

Fnictus N.F. 

Capsules 

Unofficial 

Ruta Leaves 

Ptelea Bark of root 

Belae Fructus Unripe fruit 

TAXONOMY 351 


Xanthoxylum 

americanum 

Xanthoxylum 

Clava-Herculis 

Citrus Aurantium 

Bergamia 

Citrus Aurantium 

Habitat 

Northern United 

States 


States 

France, Italy 

Northern India 

Citrus medica acida Asia 

Xanthoxylum Northern 

americanum United States 

Xanthoxylum Southern 

Clava-Herculis United States 

Ruta graveolens 

Ptelea trifoliata 

JEgle Marmelos 

Southern Etirope 

North America 

Malabar, 

Coromandel 

Simarubacecz or Ailanthus Family. A family of chiefly tropical 

shrubs or trees containing bitter principles. The leaves are alter- 

nate and pinnate. The flowers are dioecious or polygamous and 

arranged in axillary panicles (Picrasma excels a) or racemes (Quassia 

amara). The plants are distinguished from those of the Rutacea 

by the absence of secretory cavities. 

Official drug 

Quassia 

, 

Unofficial 

Simaruba 

Wood 

Part used 

Bark of root 


Picrasma excelsa 

Quassia amara 

Habitat 

West Indies 

Surinam 

Simaruba officinalis South America 

Simaruba amara Central America, 

Bahamas and 

Florida 

Burseracece or Myrrh Family. Shrubs and trees of tropical climes 

having secretion reservoirs in their bark. Leaves alternate and 

compound. Flowers small, regular and hermaphrodite, arranged in 

racemes or panicles. 

Fruit a drupe.

35 2 

Official drug 

Myrrha 

Unofficial 

Olibanum 



Gum resin Commiphora species East Africa and 

Arabia 

Gum resin Boswellia carterii East Africa and 

Arabia 

PIG. 208. Picrasma excelsa Branch. (Sayre.} 

Meliacece or Mahogany Family. Tropical trees or shrubs with 

wood often hard, colored and odoriferous. Leaves alternate, ex-

TAXONOMY 353 

stipulate, pinnately compound, rarely simple and entire. Inflorescence 

a terminal or axillary raceme. Flowers hermaphrodite or 

rarely polygamo-dicecious, regular; sepals five to four, small; petals 

usually five to four, hypogynous; 'stamens generally ten to eight 

FIG. 209. Commiphora myrrha Branch. (Sayre.) 

rarely five, very rarely twenty to sixteen, inserted outside the base 

of the hypogynous disc; filaments united into a tube; carpels usually 

five to three; style simple; ovary free, usually five- to three-celled. 

Fruit a drupe (Melia), berry (Vavaea), or capsule (Cedrella). 

exalbuminous or with fleshy albumen. 

23 

Seeds

354 


Official drug Part used Botanical origin Habitat 

Cocillana N.F. Bark Guarea Rusbyi Bolivia 

Polygalacea or Milkwort Family. Herbs or shrubs with upright, 

herbaceous to woody stems often branching profusely, the branches 

PIG. 210. Polygala senega Plant and rhizome (Sayre.) 

occasionally becoming geotropic or subterranean and bearing 

cleistogamous flowers. Leaves simple, often lanceolate or linear, 

exstipulate, alternate. Inflorescence a raceme, spike (Polygala 

'

TAXONOMY 355 

Senega) or head (P. luted). Flowers irregular, hermaphroditic 

with commonly eight stamens. Fruit a two-celled capsule (P. 

Senega), rarely a drupe or samara. Pollen grains barrel-shaped. 


Senega 

k 

,/\ . Root 

Polygala Senega United States and 

Canada 

Euphorbiacea or Spurge Family. Often herbaceous, more rarely 

shrubby, rather seldom arborescent plants. Stem, leaves and other 

parts in several genera traversed by latex canals that are either 

ramifying cells (Euphorbias) or laticiferous vessels (Manihot, 

Hevea, etc.) or rows of laticiferous sacs (Micrandra) and contain 

a white latex with acrid often poisonous contents or alkaloid or 

hydrocarbon, at times, rubber contents. Leaves alternate, exstipulate 

to stipulate, simple to pinnate or palmate. Inflorescence 

cymose. Flowers usually as in Ricinus, etc, pentamerous, diclinous; 

sepals five, green, aposepalous, becoming rudimentary or absent 

in Anthostema and Euphorbia. Petals none or five more or less 

petaloid; stamens numerous to ten to five or one(Euphorbia) ; 

pistil in pistillate flowers rarely of twenty to ten apocarpous or 

loosely syncarpous carpels (Sandbox tree), commonly of three syn- 

carpous carpels with distinct radiate styles; ovary as many-celled 

as carpels with two to one ovules in each cell. Fruit a tricoccoid 

regma or capsule, rarely winged, indehiscent, nut-like. Seeds with 

oily endosperm. Flowers at times surrounded and subtended by 

more or less petaloid and expanded bracts and bracteoles. 

Official name Part used 

Euphorbia 

Pilalifera N.F. 

Stillingia 

Oleum Ricini 

"Herb 

Root 


Habitat 

Euphorbia piluliferaTropics and sub- 

Stillingia sylvatica 

tropics

356 


XVI. Order Sapindales. Anacardiacece or Sumac Family. 

Shrubs or trees producing in stems and leaves secretion contents 

that are either acrid watery or acrid opalescent or white viscid, 

viscid acrid and poisonous. Leaves alternate, rarely opposite, 

simple (Rhus Cotinus), three-foliate (Rhus toxicodendron) or pinnate 

(Rhus glabra, R. venenata, etc.). Inflorescence frequently terminal 

and composed of racemes or cymes, often reduced to a simple raceme. 

Flowers small, clustered, green, greenish-white to greenish-yellow; 

sepals five, rarely six or four green, small; petals five smaller than 

sepals; stamens equal in number to the petals and alternate, rarely 

fewer, sometimes double in number, rarely indefinite, inserted 

hypogynously or upon an enlarged disc that surrounds or swells 

up between stamens and pistil; pistil monocarpellary more rarely 

bicarpellary, very rarely as in Spondiece of ten to five carpels, 

ovary one-celled with single ovule. Fruit a drupe. Seeds exalbuminous 

with large embryo filling seed cavity. 

Official drug

TAXONOMY 357 

purple (Euonymus atropurpureus) ; calyx four to five-lobed; 

corolla of 

four to five petals; stamens four to five, perigynous, inserted on a disc, 

which fills the base of the calyx and sometimes covers the ovary; 

ovary three- to. five-celled. Fruit a two-to five-celled capsule. 

Seeds albuminous with fleshy succulent reddish aril (Euonymus, 

Celastrus) or white membranous aril (Pachistima). 

FIG. 211. Euonymus atropurpureus. Flowering branch to left; fruiting'branch 

to right. 

Official drug 

Part used 

Euonymus N.F. Bark of root 


Euonymus 

atropurpureus 

Habitat 

United States

358 


Sapindacea or Soapwort Family. Trees, shrubs, undershrubs or 

rarely herbs of tropical climes containing the glucoside saponin. 

Stem erect or climbing (Paullinia) often provided with tendrils. 

Leaves commonly alternate and compound. Flowers in racemes 

or panicles (Paullinia), perfect or polygamo-dioscious, yellowish in 

** _ck 

PIG. 212. Cross section through root-bark of Euonymus atropurpureus. 

Note the two broad dome-shaped phloem patches, one on either side of a wedgeshaped 

primary medullary ray. 

Paullinia Cupana. Fruit a capsule (P. Cupana) } samara, drupe or 

berry. 

Seeds exalbuminous. 

Official drug 

Guarana

TAXONOMY 359 

palmately lobed, cleft (Acer) or pinnate (Negundo). Inflorescence 

a raceme condensing in some species to a capitulum of cymes. 

Plowers small, regular, polygamous or dioecious; sepals five to four 

green; petals none or five, variously colored; stamens usually eight, 

hypogynous or perigynous; nectar disc around stamens or between 

them and pistil; pistil bicarpellary with two-celled ovary. Fruit a 

samara. Seeds green, exalbuminous with coiled or folded embryo 

and long thin cotyledons. 

Unofficial drug Part used Botanical orign Habitat 

Acer Spicatum Bark Acer spicatum United States 

XVII. Order Rhamnales. Rkamnace or Buckthorn Family. 

Shrubs or low trees usually of branching or spreading habit. 

Branches either cylindric or long green or hardened, checked back 

and spinescent, occasionally, especially flowering branches developing 

tendrils for support. Leaves simple, usually alternate. Flowers 

hermaphrodite or more or less diclinous, pentamerous 

.ous, greenish to greenish-yellow to yello wish-white ; sepals 

to tetramer- 

five to 

four; petals five to four alternating with sepals; stamens five oppo- 

site the petals, perigynous; pistil either free in center of receptacular 

cup or more or less fused with it and so semi-inferior, ovary typically 

three-celled becoming rarely four-celled with two to one atropous 

ovules in each cavity. Fruit of three indehiscent cocci, each en- 

closing a single albuminous seed with straight embryo imbedded 

in albumen. 

Official drug Parts used Botanical origin Habitat 

Cascara Sagrada Bark Rhamnus Purshiana Northern Cali- 

fornia to south- 

western British 

America 

Frangula Bark Rhamnus Frangula Europe 

Rhamnus Fruit Rhamnus cathartica Asia and Africa 

Cathartica N.F. 

Vitacea or Grape Family. Rarely tall herbaceous, usually shrubby 

and climbing, more rarely shrubby upright plants. Stems rarely 

short more usually elongate, feeble, rather brittle, climbing by 

tendrils which represent modified inflorescence shoots. Leaves


alternate, simple to lobed (either pinnately or more often palmately) 

to compound-pinnate or palmate. Perfect graded series of lanceo- 

late leaves with pinnate veining to palmate veining; from pinnately 

veined to compound-pinnate; from palmately veined to compound- 

FIG. 213. Rhamnus frangula Branch. (Sayre.) 

palmate. Stipules greenish to membranous or none. Flowers in 

racemes of compressed cymes, hermaphrodite or diclinous, nearly 

always small, clustered, green to greenish-yellow or greenish-white, 

rarely otherwise; sepals five, rarely four, small to minute (mere 

rim of receptacle) more or less persistent. Petals five, deciduous to

TAXONOMY 

caducous, typically distinct, in Vitis united by their tips into calyptroform 

corolla, so in June, as Grape Vine flowers expand, corolla 

splits at base into five lobes that separate below, being attached at 

tips, while whole becomes tumbled off by lengthening stamens. 

Pistil bicarpellate. Ovary two-celled, superior at or most sub- 

inferior. Ovules two to one, erect. Style 

short often more or less 

thickened with terminal, capitate, slightly two-lobed stigmas. 

Stamens equal to petals or sepals and opposite petals. Receptacle 

internal to stamens, often expanded into nectariferous girdle or, in 

Vitis, into receptacular knobs alternating with stamens. Fruit 

a berry rarely six- to three-celled, typically two-celled and with 

two to one seeds in each cavity. Seeds like ovules, erect, with bony 

testa. Embryo small, imbedded at base of cartilaginous albumen. 

Official drug 

Vinum Xericum 

N.F. 

Part used 

Fermented juice 

of ripe fruit 


Vitis species 

(cultivated) 

Habitat 

Cultivated 

XVIII. Order Malvales. Sterculiacea or Cola Family. Rarely 

trees with soft wood and 

herbs, usually shrubs or tall, often heavy 

broad annual rings. The cambium, in developing bast, produces 

one, two, three, four, or five alternating layers of hard and soft 

bast which in some species of this as well as the Tiliacece family 

form long finger-like processes pushing out into the cortex. Leaves 

alternate, sometimes simple and pinnately veined or passing to 

palmately veined or palmately compound. Flowers hermaphrodite; 

sepals five, sometimes surrounded by bracteoles forming an epicalyx; 

petals usually five; stamens typically five hypogynous, opposite 

petals, distinct or slightly fused in monadelphous fashion (Melochia, 

Waltheria) or, stamens subdivided above into few or numerous stam- 

inal leaflets, anthers two-celled; pistil many to ten- to five- or four- 

carpelled; carpels apocarpous or more usually partially or completely 

united. Fruit either follicles, or fused to form a capsule of ten or 

more, frequently five dehiscent carpels or, carpels splitting asunder 

into cocci or, becoming a woody capsular nut (Theobroma) or, 

rarely the fruit may become succulent. Seeds globose or subglobose 

and often provided with wings, arils or similar appendages; embryo 

straight, large and .surrounded by scanty albumen.

362 

Official drug 

Oleum 

Theobromatis 



Fixed oil Theobroma Cacao Tropical America 

- FIG. 214. Theobroma cacao Branch and fruit. (Sayre.} 

Cacao Prsepara- Prepared powder Theobroma Cacao Tropical America 

turn N.F. from roasted and other species 

kernels of seeds 

Kola N.F. Cotyledons Cola acuminata and Africa, West 

other species 

Indies

TAXONOMY 363 

Tiliacea or Linden Family. Shrubs or trees, rarely herbs, having 

stellate hairs on both stems and leaves. Leaves alternate, pinnately 

more rarely palmately veined, stipulate. Inflorescence cymose. 

Flowets hermaphrodite, more rarely, by absorption, more or less 

diclinous; sepals and petals five each, more rarely four, sepals de- 

ciduous; stamens five opposite the petals or, as in Sterculiacea, 

five phalanges of stamens representing subdivided stamens (Tilia), 

pistil of ten to five or two syncarpous carpels; ovary superior. 

Fruit either a nut-like drupe or drupe, rarely baccate. 

Unofficial drug Part used Botanical origin Habitat 

Tilia Inflorescence Tilia species United States and 

Europe 

Malvaceae or Mallow Family. Herbs in temperate regions (Malva 

rotundifolia, Althcea officinalis, etc.), occasionally shrubs in temperate 

regions (Hibiscus Syriacus, etc.), frequently shrubs or tall trees in 

the tropics. Stems, as in Sterculiaceoe and Tiliaceoe, sometimes 

forming numerous layers of hard and soft bast. Leaves alternate 

and stipulate, ovate, ovate-cordate, orbicular or palmately com- 

pound; venation pinnate or palmate. Stems, roots and leaves con- 

tain mucilage cells. Inflorescence a raceme or fascicle of cymes. 

Flowers regular, pentamerous; calyx green, of five aposepalous sepals 

but frequently surrounded outside by an epicalyx. Both calyx and 

epicalyx are persistent. Corolla of five petals varying in color which 

are more or less fused with stamens at their bases; stamens mon-adel- 

phous and forming an upright column enclosing the styles; anthers 

one-celled, dehiscing transversely; pollen grains echinate; pistil 

loosely or strongly syncarpous, rarely sub-apocarpous of thirty to 

five carpels. Fruit either a set of cocci, follicles or a capsule (Gossypium). 

Seeds albuminous with oily and mucilaginous albumen.

364 


FIG. 215. Gossypium herbaceum Branch. (Sayre.)

Official drug 

Part used 

Althaea Root (peeled) 

Althaea Folia N.F. Leaves 

Gossypii Cortex N.F. Bark of root 

Gossypium 

Purificatum 

Oleum Gossypii 

Seminis 

Hairs of seed 

Fixed oil from 

seeds 

Malvae Folia N. F. Leaves 

Unofficial 

TAXONOMY 36S 

Botanical origin Habitat 

Althaea officinalis Europe and Asia 

Althaea officinalis Europe ard Asia 

Cultivated varieties of: 

possypium 

herbaceum 

Gossypium 

Barbadense 

Gossypium 

arboreum 

varieties 

of Gossypium 

herbaceum 

(Cultivated 

varieties 

of Gossypium 

{Cultivated 

species 

( Malva sylvestris ( 

I Malva rotundifolia I 

Arabia, United 

States, East 

indies 


Africa 

Egypt, Arabia and 

India 

Indies, United 

States 

{Arabia, 

\ 

East 

States, 

Asia, Africa and 

{United South America 

Althaea Flores Flowers Althaea rosea Europe 

FIG. 216. Upland'^cotton (Gossypium hirsutum). A, mature boll opened 

out; B, cross-section of young boll; C, single seed with fibers; D, young boll. 

(Robbing.)

366 


XIX. Order Parietales (Ovaries of flower have parietal placentas.) 

Theacea or Tea Family (Ternslr(zmiace(z,Cammeliace(e). Ever- 

green shrubs or low branching or tall, often heavy trees with watery 

juice. Leaves for the most part alternate, evergreen, often leathery, 

sometimes membranous; stipuleS either bud scales and caducous 

or often absent; leaf margins sinuate or serrate (Thea). Inflores- 

cence a raceme becoming by condensation terminal, one-flowered. 

Flowers regular, perfect, pentamerous; sepals five, rarely four to 

three, deciduous, occasionally subtended by bractôlar scales; petals 

five, brittle and succulent varying from greenish-white or greenish- 

yellow through yellow to white or, from whitish pink to pink, scarlet, 

crimson, very rarely a tendency toward purple; stamens typically 

five but, as they grow, they subdivide into staminal leaflets, so that 

in their mature condition they are apparently^ indefinite and mono- 

to polyadelphous; stamens inserted hypogynously or perigynously 

and opposite the petals; pistil of typically five syncarpous carpels but 

reduced in some species to four to three or two. Fruit usually a capsule 

(Thea), five- to three-celled, dehiscing longitudinally, more 

rarely a fleshy, semi-baccate, semi-drupaceous indehiscent fruit. 

Seeds with scanty or no albumen and often attached to inner angle 

of cells by projecting spongy placentae. 


Caffeina Feebly basic Thea sinensis Eastern Asia 

substance 

Unofficial 

Thea Leaves Thea sinensis var. Eastern Asia 

Bohea and vifidis 

Guttiferce or Gamboge Family. Tropical trees (Garcinia), rarely 

shrubs, containing resinous principles in resin canals found in 

cortex and pith. Leaves opposite, coriaceous. Flowers dioecious, 

generally pentamerous or tetramerous with usually five stamens 

which are subdivided. Fruit a berry (Garcinia Hanburyi), drupe 

or capsule. Seeds generally large; embryo large to huge, often 

with enlarged radicle and reduced or absorbed cotyledons. 


Cambogia Gum resin Garcinia Hanburyi 1 

Malabar coast 

and Travancore

TAXONOMY 367 

Hypericacece or St. John's Wort Family. Herbs or shrubs 

of temperate climes with opposite (Hypericum perforation) rarely 

whorled branches and balsamic, resinous juices, which, in the herbaceous 

species, are secreted by black or pellucid glands found in the 

PIG. 217. Garcinia hanburii Branch. (Sayre,} 

leaf parenchyma. Leaves entire, opposite, usually sessile, exstipu- 

late, and dotted. Flowers, regular, hypogynous and arranged in 

panicles or forked cymes. Petals usually 5. Stamens usually 

indefinite, rarely definite, often in 3-5 sets, more rarely monadelphous

368 


or free. Pistil of usually 35 carpels with 3-5 celled compound 

ovary and as many filiform styles as carpels. Fruit a capsule 

with usually septicidal dehiscence (Hypericum) or a berry. 

small, numerous, anatropous and exalbuminous. 

Seeds 


Hypericum Entire plant Hypericum perforatum Europe 

Canellacece Family. Trees the bark of which contains aromatic 

principles. Leaves alternate, pellucid-punctate. Flowers regular, 

golden-yellow, and arranged in terminal or axillary cymes. Fruit 

a berry containing two to many seeds with oily and 'fleshy albumen. 


Canella N.F. Inner bark Canella Winterana , 

Florida and West 

T ,. 

I Indies 

Bixacece Family. Tropical shrubs or trees. Leaves alternate, 

simple with minute or no stipules. Flowers hermaphrodite or unisexual, 

regular-, stamens hypogynous, mostly indefinite with anthers 

opening by slits, rarely by one or two apical pores (Bixa). Fruit 

fleshy or dry. Seeds with fleshy albumen and sometimes covered 

with a fleshy arillus (Bixa Orellana). 


, . T /-k 11 / Tropical America 

Annatto Coloring matter Bixa Orellana < 

I and Madagascar 

Chaulmoogra oil Fixed oil from Gynocardia odorata India 

seeds 

Violacece or Violet Family. Herbs or shrub. Stems upright, 

rarely creeping, spreading or acaulescent. Leaves either cauline or 

radical, stipulate, alternate, simple to pinnatifid or palmate. 

Flowers pentamerous, regular or irregular (Viola). Fruit a loculi- 

cidally dehiscent capsule (Viola) rarely baccate. Seeds albuminous. 


Viola Entire herb Viola tricolor Temperate regions 

Turneracea or Damiana Family. Tropical herbs, shrubs or 

trees. Leaves alternate, simple, petioled, exstipulate. Flowers 

perfect, regular, axillary, pentamerous with one-celled ovary. Fruit

TAXONOMY 369 

a capsule with three valves. Seeds strophiolate with albuminous 

embryo. 

Official drug 

Damiana N.F. Leaves 

Part used 


rTurnera diffusa 

Turnera 

aphrodisiaca 

Habitat 

Lower California 

and Mexico 

Passifloracea or Passion Flower Family. Herbaceous or woody 

vines climbing by tendrils. Leaves alternate, simple, entire, lobed 

or compound. Flowers perfect or imperfect, solitary; peduncles 

jointed at the flower; perianth petaloid with urceolate or tubular 

tube and four to five or eight to ten partite and two-seriate limb, the 

throat usually crowned by one or more series of subulate filaments 

which are frequently colored; gynophore elongating supporting the 

stamens and pistil. Fruit a one-celled berry (Passiflora) or three- 

to five-valved dehiscent capsule containing numerous seeds. 

Official drug 

Passiflora N.F. 

Part used 

Entire herb 


Passiflora incarnata 

Habitat 

United States 

Caricaceos or Papaw Family. A family of latex-containing trees 

composed of two genera indigenous to tropical America. Of chief 

pharmaceutic interest is the species Carica Papaya, the Papaw or 

Melon tree, the fruit of which yields Papain, a valuable digestive 

ferment. This plant is a tree about 20 feet high which bears at its 

summit a cluster of deeply lobed petiolate leaves and dioecious flow- 

ers. The fruit is a berry, the size of one's head and contains an acrid 

milky juice from which papain can be precipitated by the addition 

of alcohol. 

Cistacece or Rock Rose Family. Herbs or shrubs whose stem and 

branches are "often glandular, pubescent or tomentose, with simple 

or stellate trichomes. Leaves simple, entire, the lower ones opposite, 

upper alternate. Flowers perfect, regular, terminal, and solitary 

or in cymes or unilateral racemes; sepals five, the two external ones 

often bractiform or wanting; petals five (Helianthemum) rarely three 

or none (Lechea); stamens hypogynous, indefinite; carpels three to 

three- to five-valved 

five, ovary free, one- celled. Fruit a one-celled, 

capsule. 

24

370 



Helianthemum Herb Helianthemum Eastern United 

N.F. canadense States 

XX. Order Opuntiales. Cactacea or Cactus Family. Herba- 

ceous rarely arborescent (Cereus giganteus) more or less succulent 

PIG. 218. Daphne mezereum Fruiting branch and flowers. (Sayre.} 

plants living in warm, dry (Peireskia), usually desert situations, 

rarely becoming epiphytic and correspondingly modified. Stems 

accordingly varying from elongate, slightly enlarged, green (Peireskia) 

to flattened (Cereus and Opuntia), to condensed (Echinocactus ,

TAXONOMY 371 

Echinocereus , etc.), to greatly condensed (Mamillaria). Leaves 

alternate, stipulate or exstipulate, enlarged and more or less fleshy 

(Peireskia), becoming reduced, green and semicircular (Opuntia), 

or modified into spines, or wholly absorbed. Flowers, regular, 

solitary or fascicled in axils of leaves; sepals five; petals similar to 

sepals, petaloid, small to much enlarged, in color varying from yellow 

to white or from yellow to yellowish-pink, pink, scarlet or crimson; 

stamens indefinite, inserted at varying levels in the throat of a greatly 

expanded upgrown receptacle; pistil generally tricarpellary; ovary 

inferior, often deeply sunk in upgrown receptacular part; style threadlike, 

divided above into as many stigmas as carpels. Fruit a receptacular 

berry enclosing numerous small seeds. Seeds exalbuminous. 

Official drug 

Cactus 

Grandiflorus 

N.F. 

Part used Botanical origin 

Fresh succulent Cactus grandiflorus 

stems (Cereus grandiflorus) 

Habitat 

Tropical America 

XXI. Order Myrtales (Myrtiflorae). Thymelevcece or Mezereum 

Family. Shrubs (Daphne Mezereum) or low trees, usually of branching 

habit, the stems developing long tenacious bast fibers. Leaves 

alternate, rarely opposite, coriaceous, simple, varying from lanceolate 

to ovate. Inflorescence a condensed raceme or spike. Flowers 

perfect, polygamous or dioecious, small with calyx 

alone of the 

perianth parts developed. This is crimson-purple in Daphne 

Mezereum. Sepals usually fused to form a tube or cup-shaped 

perianth. Stamens usually eight in two rows of four longer and 

four shorter (Daphne Mezereum) inserted on the calyx tube. Pistil 

monocarpellary; ovary superior mostly one-celled with a single 

pendulous ovule. Fruit a nut, drupe, or berry (Daphne). 

Official drug 

Mezereum Bark 

Part used 


Daphne Mezereum 

Daphne Gnidium 

Daphne Laureola 

Habitat 


Punicacece (Lythracecz) or Pomegranate Family. Herbs (Cuphea), 

shrubs (Decadon) or low trees (Punica). Leaves either alternate, 

opposite (Punica) or whorled, simple, usually lanceolate to ovate, 

entire, often glandular and viscous. Inflorescence a raceme, spike,

372 


or condensed cyme. Flowers perfect, usually regular, but pass more 

or less to irregular, sometimes very irregular as in genus Cuphea; 

sepals five to four, more or less fused below in themselves and with 

calyx tube; petals commonly five, often frilled or crumpled, inserted 

on the mouth of the calyx tube; stamens fifteen, ten or five in alter- 

nate rows of five each, inserted hypogynously or perigynously; 

pistil six-, five-, four,- two--, rarely one-carpeled with as many cavi- 

ties in the ovary and numerous small ovules; style elongate with 

pointed or knobbed stigma. Flowers of Punica granatum are 

scarlet in color. Fruit a baccate capsule (Punica granatum) or 

FIG. 219. Punica granatum Branch with flowers. (Sayre.) 

capsule, dehiscing longitudinally or transversely. Seeds exalbuminous. 


Granatum Bark Punica Granatum India 

Unofficial 

Granati Fructus Rind of fruit Punica Granatum India 

Cortex 

Henna Leaves Lawsonia inermis Egypt, Arabia

TAXONOMY 373 

PIG. 220. Eucalyptus globulus Branch. (Sayre.)

374 


MyrtacecR or Myrtle Family. Rarely herbs (Careya) mostly 

shrubs or trees, some being the tallest trees known (Eucalyptus). 

Stems often tend to develop cork in flakes which separate much as 

in the Buttonwoods. Leaves rarely alternate nearly always oppo- 

PIG. 22i. Eugenia aromatica. (Sayre.) 

site, entire, often glistening, subcoriaceous to coriaceous (Eucalyptus, 

Pimenta, etc.), frequently edge-on in position upon branches. 

Inflorescence cymose, at times forming scorpioid cymes becoming 

condensed into small fascicles, or each cyme condensing into a 

solitary flower.

TAXONOMY 375 

Flowers regular or very rarely irregular from the lop-sided devel- 

opment of the stamens. Symmetry rarely hexamerous, typically 

pentamerous, not infrequently reduced to tetramerous (Clove) ; sepals 

five, six or four, aposepalous, or synsepalous at base, superior, and 

inserted around the edge of an expanded, upgrown receptacular 

disc, varying from green and more or less expanded to short, thick 

in num- 

fleshy (Clove) or reduced to teeth (Eucalyptus) ; petals equal 

ber to the sepals, more or less petaloid and enlarged, rarely reduced 

and wanting, varying in color from green through greenish-yellow 

to white (Eugenia species) or from whitish to pink, scarlet, crimson, 

purple and blue', petals sometimes synpetalous and cup-like, detach- 

ing as the flower opens; stamens usually indefinite and epigynous, 

varying in the color of their filaments as do the petals; pistil rarely 

of ten to six carpels usually of five, not infrequently, as in Clove, of 

four carpels; ovary inferior or semi-inferior, as many-celled as there 

are carpels and with central placentation; style elongate; stigma 

undivided. Fruit either a hard, woody indeshicent nut (Brazil 

Nut), a capsule dehiscing at apex (Eucalyptus) dr berry (Eugenia). 

Seeds exalbuminous. 

Official drug 

Eucalyptus 

Eucalyptol 

Caryophyllus 

Eugeinol 

Pimenta N.F, 

Oleum Cajuputi 

Unofficial 

Myrcia 

Leaves 

Part used 

Organic oxide 

Flower buds 

Aromatic phenol 

Fruit 

Volatile oil from 

leaves and twigs 

Volatile oil and 

leaves 

Eucalyptus Kino Inspissated juice 


Habitat 

Eucalyptus globulus 1 Australia, 

Eucalyptus globulus / Tasmania 

Eugenia aromatica 

Eugenia aromatica 

Pimerta officinalis 

Melaleuca 

Leucadendron 

Myrcia acris 

Eucalyptus rostrata 


Molucca Islands 

West indies; 

Central America, 

Mexico 

East Indies 

West Indies 

Australia 

Combretacece or Myrobalans Family. Mostly tropical shrubs 

and trees containing considerable tannin. Leaves exstipulate, 

alternate or opposite, simple, pinnately veined, entire or toothed. 

Inflorescence a raceme, spike or head. Flowers regular, perfect


or imperfect. Fruit a drupe, frequently longitudinally winged, 

containing a single seed. 


Combretum Leaves Combretum Sumatra 

sundaicum 

XXII. Order Umbellales or Umbelliflorse. Araliacea or Gin- 

seng Family. Herbs (Panax quinquefolium, Hedera Helix, Aralia 

nudicaulis, etc.), undershrubs (Aralia hispida, etc.), shrubs (Fatsia 

horrida), or trees (Aralia spinosa) with stems which are more or less 

hollow along internodes and solid at nodes. Leaves alternate, varying 

from simple to trifoliate or to multipinnate (tropical Aralias) or 

passing by telescoping into compound-palmate. Leaves serrate 

margined and along with stem they develop volatile oil, resin and 

gum contents in secretion reservoirs. Inflorescence varying from a 

raceme of umbels to a raceme and even to condensed racemose 

umbels. Flowers regular, generally pentamerous, small, generally 

inconspicuous, green, greenish-yellow to rarely white, usually 

hermaphrodite but sometimes polygamous or dioecious; sepals 

five, rarely four; petals five, rarely four, often greenish to greenish- 

yellow, occasionally white, seldom pink in color; stamens varying 

from indefinite to ten to commonly five, opoosite sepals, and, like 

sepals, epigynous in insertion; anthers versatile; pistil occasionally 

fifteen- to ten-, usually five-carpellate; ovary as many celled with one 

or rarely two pendulous ovules in each cavity; styles distinct 

ending in knob-shaped stigmas. Fruit a berry. Seeds albuminous. 

Official drug 

Aralia N.F. 

Unofficial 

Aralia Nudicaulis 

Aralia Spinosa 

Ginseng 

Panax Repens 

Part used 

Rhizome and 

roots 

Rhizome 

Bark 

Root 

Rhizome 


Aralia racemosa 

Aralia nudicaulis 

Aralia spinosa 

Habitat 

Eastern Uuited 

States and 

Canada 


States and 

Canada 


States 

Panax quinquefolium North America 

Panax repens Japan

TAXONOMY 377 

Umbellifera or Parsley Family. Herbs, rarely shrubs, often of 

rapid growth, and with upright, fistular (hollow at internodes, 

solid at nodes), often grooved and ridged stems. Leaves alternate, 

compound and usually much divided, exstipulate, but with expanded 

sheathing and flattened leaf base (Pericladium), that ensheathes 

the stem. Inflorescence a simple or often compound umbel sur- 

rounded by an involucre of bracts or of bracteoles. Flowers small, 

pentamerous, with inferior ovary and superior floral parts. Sepals 

minute, tooth-like, inserted above inferior ovary, or absorbed. 

Petals small, usually yellow to white, rarely pink to purple, distinct, 

each with inflexed tip. Stamens five, epigynous, inserted below a 

nectariferous, epigynous disc, incurved in bud. Carpels two, fused 

into bicarpellate pistil. Ovary two-celled, with one pendulous 

ovule in each cell, ovarian wall traversed by oleoresin canals; 

styles two, distinct above the nectar disc or stylopod. 

Fruit a dry, 

splitting fruit or cremocarp, that splits lengthwise into two mericarps 

which hang for a time by a forked carpophore. Seeds single in each 

mericarp, albuminous. 

Official drug 

Part used 

Anisum Ripe fruit 

Anethol 


Pimpinella Anisum 

Habitat 

Asia Minor, Egypt 

and Greece

378 

N.F. 

Official drug 


Part used 

Angelica Radix Rhizome and 

N.F. roots 

Apii Fructus N.F. Ripe fruit 

Unofficial 

Imperatoria Root 

Pimpinella 

Ammoniacum 

Galbanum 

Levisticum 

Roots 

Gum-resin 

Gum-resin 

Root 


ica and other species 

of Angelica 

Angelica atropurpurea 

and other 

species of Angelica 

Apium graveolens 

Tmperatoria 

Ostruthiuni 

Habitat 

and Siberia 

United States 

and Canada 

England 

Europe 

Pimpinella Saxifraga \ Central Europe 

. Pimpinella magna 

Dorema 

Ammoniacum 

Ferula galbaniflua 

Levisticum officinale 

J 

Persia 

Persia and 

Afghanistan 

Europe 

Cornacea or Dogwood Family. Herbs (Cornus canadensis, etc.), 

shrubs (Cornus sanguinea, etc.) or trees (Cornus florida, Nyssa 

sylvatica, etc.). Leaves simple, alternate (Sour Gum), or opposite 

(Dogwoods). Inflorescence an umbel or head, the whole being 

surrounded by an enlarged and more or less petal oid involucre. 

Flowers regular, rarely pentamerous, more frequently tetramerous; 

sepals usually four, small tooth-like or absorbed; petals usually 

four, small, greenish to yellowish to white (Cornus florida), rarely 

pink or crimson; stamens four or five, alternate to the petals and 

inserted with the sepals and petals epigynously around and between 

the nectar disc; pistil syncarpous, bicarpellate, rarely tricarpellate; 

ovary as many celled with one pendulous ovule in each cavity; 

style usually simple, ending in rounded or slightly bilobed stigma. 

Fruit a two-seeded drupe. Seeds albuminous. 

Official drug 

Cornus N.F. N 

Part used 

Bark of root 


Cornus florida 

Habitat 


States and 

Canada 

SUB-CLASS B. SYMPETALE (GAMOPETAL.E OR METACHLAMYDE.E) 

A division of dicotyledonous plants in which the flowers possess 

both calyx and corolla, the latter with petals more or less united 

into one piece.

TAXONOMY 379 

I. Order Ericales. Ericacea or Heath Family. Sub-herbaceous 

(Chimaphila), suffruticose (Erica), fruticose (Azaleas, Kalmias, 

etc.), rarely sub-arborescent (Arbutus unedo or Strawberry Tree) 

plants. Roots fibrous often saprophytically associated, rarely 

tuberous or more or less enlarged. Stem upright, ascending or creeping, 

more or less woody, rarely through saprophytic connection be- 

coming soft, annual and pale above ground (Monotropa uniflora) . 

FIG. 222, Arctostaphylos uva ursi^-Eranch, flower, and fruiting branch. (Sayre.) 

Leaves alternate, simple, entire, exstipulate, rarely soft, delicate, 

herbaceous (Azaleas} , usually leathery to wiry and evergreen, more 

rarely (Pterospora, Monotropa, etc.) becoming greenish-blue, bluish- 

yellow, yellowish-white to white and correspondingly saprophytic. 

Inflorescence typically a raceme (Pyrola, Andromeda, Gaylussacia, 

Erica, Arctostaphylos Uva Ursi, etc.) but raceme condensed into a 

racemose umbel (Azalea, etc.) or further reduced to a few flowers or, 

in the degraded saprophytic condition to one flower (Monotropa

3 8o PHARMACEUTICAL BOTANY 

uniflora)\ Flowers regular, passing to irregular (Rhododendron}, 

pentamerous or tetramerous; sepals five tq. four, rarely fewer, apo- 

to synsepalous, usually green, sometimes brightly petaloid; petals 

five more rarely four, slightly to deeply synpetalous, cup-shaped 

(Kalmia) to urceolate (Arctostaphylos, Andromeda, etc.), yellow to 

white or through yellow pink to scarlet to crimson to crimson- 

purple; stamens ten to eight in two circles of five to four each, be- 

coming by absorption of inner circle, five to four only, hypogynous, 

epipetalous or epigynous; anthers two-celled, dehiscing by apical 

pores (Arctostaphylos) or apical slits; pollen sometimes agglutinated 

into long viscous threads; pistil five- to four-, rarely six- to eight car- 

peled, superior, rarely semi-inferior to inferior ( Vaccinece) ; ovarv as 

many celled as there are carpels; style elongated, filiform, usually 

five- to four lobed. Fruit a capsule (Trailing Arbutus), berry \Vac- 

cinium) or false drupe (Gaultheria). 

Official drug 

Chimaphila N.F. 

Uva Ursi 

Methylis Salicylas 

Unofficial 

Gaultheria 

Part used 

Leaves 

Leaves 

Volatile oil 

Leaves 

Seeds small, anatropou* 


Chimaphila 

umbellata 

Arctostaphylos Uva 

Ursi 

Gaultheria 

procumbens 

Habitat 

United States, 

Canada, Northern 

Europe and 

Asia 

Northern United 

States and 

Canada, Europe 

and Asia 


Canada 

Gaultheria United States and 

procumbens Canada 

II. Order Ebenales. Sapotacea or Star Apple Family. Tropical 

shrubs or trees (Palaquium) characterized by the presence of laticiferous 

sacs in the pith and cortex of the stems and adjoining the 

veins of the leaves. Leaves alternate, exstipulate, evergreen and 

coriaceous. Flowers perfect, large and axillary. Fruit a berry 

(Palaquium) rarely a capsule (Ponteria). 


coagulated 

milky 

(Purified exudate 

Various species of 

Palaquium 

Indo-China and 

East Indies

TAXONOMY 381 

Styracea or Benzoin Family. Shrubs or low trees. Leaves alternate 

to opposite, entire, often- acuminate. Flowers hermaphrodite, 

regular, rarely sub-irregular, either in condensed fascicles or solitary 

in the axils of the leaves; sepals and petals typically five each; 

corolla often white, rarely pinkish or yellowish; stamens many to 

four to two, perigynous or sub-hypogynous ; pistil bicarpellary or 

four to five carpellate. 

rarely as many-celled as there are carpels. 

Fruit either fleshy or dry, often winged and 


, 

,, 

/ Styrax Benzoin East Indies and 

Benzomum Balsamic resin < . 

( and other species biam 

III. Order Contortse (Gentianales). Oleacece or Olive Family. 

Shrubs (Forsythia, Chionanthus, Syringa, etc.) or trees (Fraxinus, 

Olea, etc.) with stems possessing close white wood, and slightly 

swollen or enlarged nodes. Leaves opposite, decussate, simple, 

rarely pinnately compound (Ash). 

Inflorescence dichesial of scor- 

pioid cymes but tending constantly toward condensation and so in 

the Lilac, the inflorescence becomes a clustered raceme of cymes 

(thyrsus). Flowers regular, pentamerous or tetramerous; sepals 

small, green, rarely petaloid, synsepalo.us; petals synpetalous, elongated 

into a narrow tube, expanding above into a stellate limb; 

stamens very rarely five, rarely four to three, nearly always two, 

epipetalous and high set on corolla tube; pistil bicarpellate, rarely 

of three to four 'carpels; ovary two-celled with two to one pendulous 

ovules in each cavity. Fruit either a capsule (Lilac), drupe (Olive), 

berry (Privet) or a winged indehiscent akene (Ash). Seeds with 

moderate to scanty albumen becoming occasionally exalbuminous. 


Manna Dried saccharine Fraxinus Ornus Southern Europe 

exudate 

Oleum Olivse Fixed oil Olea Europsea Southern Europe, 

Algeria, Asia 

Chionanthus N.F. Bark of root Chinoanthus Southern United 

virginica 

States 

Fraxinus N.F. Bark Fraxinus AmericanaNorthern United 

States and 

Canada


Loganiacea or Logania Family. Herbs (Spigelia, etc.), woody 

vines (Gelsemium, etc.) or trees (Strychnos Nux Vomica, etc.) with a 

bitter juice usually containing alkaloids. Stem, rarely herbaceous, 

FIG. 223. Strychnos nux vomica Flowering branch and seeds. (Sayre.} 

usually woody, often long climbing and roperlike (Gelsemium), usu- 

ally with a bicollateral bundle system. Leaves opposite, stipulate

TAXONOMY 

or exstipulate. Inflorescence racemose or cymose (Spigelia) (scor- 

pioid cyme (Strychnos), sometimes condensed into solitary, axillary 

flowers. Flowers perfect, usually regular; calyx gamosepalous; 

corolla gamopetalous, hypogynous, rotate, campanulate or infundi- 

PIG. 224. Gentiana lutea Flowering head and dissected flower. (Sayre.) 

buliform; stamens inserted on the corolla tube or throat and with 

thread-like filaments; ovary superior, two-celled; style elongate with 

bifid stimga; ovules numerous. Fruit usually a capsule, septicidally 

dehiscent (Gelsemium sempervirens) , or loculicidally dehiscent (Spige-


lia marilandica) , sometimes a berry (Strychnos Nux Vomica) or 

drupe. Seeds numerous or solitary, sometimes winged. 


Nux Vomica

TAXONOMY 

latex tubes which ramify through the cortex and mesophyll tissues. 

Leaves alternate, opposite or verticillate, simple, entire, deciduous 

or evergreen. Inflorescence cymose. Flowers regular, pentamerous, 

rarely tetramerous; sepals five, gamosepalous, green, rarely sub- 

petaloid to petaloid; petals five, slightly to deeply gamopetalous, in 

PIG. 225. Strophanthus hispidus Branch and seed with comose awn. (Sayre.) 

shape varying from open tubular, stellate, to elongate tubular to 

elongate funnel-shaped, in color varying from greenish-yellow to 

white or from yellow to yellow-red to crimson to crimson-purple to 

nearly purple-blue; stamens five, epipetalous; pistil usually bicarpel- 

late; ovary two-celled with central placentation; style more or less 

25


Part used Botanical origin Habitat

TAXONOMY 387 

woody, perennial climbing plants with underground parts sometimes 

swollen into tuberous roots (Jalap, Sweet Potato, Wild Man of the 

Earth). Stems rarely short, upright or tufted, usually elongate 

and circumnutating in action. Vascular bundles frequently bi- 

collateral. Leaves alternate, simple, exstipulate, varying from 

FIG. 226. Convolvulus scammonia Branch. (Sayre.) 

cordate to cordate-sagittal, to broad reniform to reniform, palmately 

lobed to palmatifid to palmately-compound (Ipomaa shows all these 

transitions). Stem and leaves frequently contain a dull, viscous, 

watery to milky-white juice. Inflorescence a scorpioid cyme becoming 

reduced in some forms to a solitary flower. Flowers penta-

388 


merous; sepals five, green, gamosepalous; corolla varying in shape 

from rotate to funnel-like with expanded mouth, in color from 

greenish-yellow to white or through yellowish-pink to scarlet, 

crimson, purple or blue; stamens five, often with the bases of the filaments 

expanded; pistil bicarpellate; ovary two celled, superior, often 

surrounded by a nectar girdle; style 

filiform with bilobed or bifid 

stigma. Fruit usually a capsule (Exogonium, etc.), dehiscing septifragally, 

rarely a berry. Seeds scantily albuminous to exalbumi- 

nous-. 

Official drug 

' 


Jalapa Tuberous root Exogonium Purga Mexico 

Convolvulus \ Asia Minor, 

Scammoniae Radix Root ( ~ c . 

Scammonia ) Greece, Syria 

Unofficial 

Male Jalap Root > Ipomoea orizabensis Mexico 

Tampico Jalap Root Ipomoea simulans Mexico 

Wild Jalap Root Ipomoea pandurata United States 

Turpeth Root Root Operculina TurpethumEast Indies 

Hydrophyllacece or Water Leaf Family. Annual, herbaceous, 

rarely perennial woody plants whose stems, branches, leaves and 

sepals are often viscous and glandular hairy. Leaves alternate, 

exstipulate, from simple linear to pinnatipartite to pinnate. Inflorescence 

rarely expanded, usually scorpioid cymes. Flowers small 

to large, funnel-form in Eriodictyon calif ornicum; sepals five, green; 

petals five, regular; corolla varying from small stellate with slightly 

fused petals to large rotate, campanulate or tubular, in color varying 

from greenish-white or yellow to rarely white, often pink, purple 

or blue; stamens five, rarely with alternate staminodes; pistil bicar- 

pellate. 

Fruit a two-celled capsule dehiscing usually septicidally. 


Eriodictyon Leaves Eriodictyon California and 

californicum New Mexico 

Borraginacea or Borage Family. Herbaceous (Borraginea sub- 

family) or shrubby (Heliotropes sub-family), plants forming a primary 

root and a single or often branched shoots. Leaves often 

divisible into expanded, sometimes large basal and alternate scat- 

tered cauline leaves. Each of these simple, exstipulate, often hairy,

TAXONOMY 389 

rarely glabrous. Inflorescence a raceme of dichesial or scorpioid 

cymes, at times condensed into a dichesium of scorpioids or a simple 

scorpioid cyme. Flowers pentamerous, regular, passing to slight 

or marked irregularity (Eckium)', sepals five; green, slightly or 

FIG. 227. Atropa belladonna Branch. (Sayre.) 

deeply gamosepalous, often hairy; petals five, the corolla varying in 

shape from rotate with shallow tube (Myosotis and Borage), to 

tubular (Symphytum), to funnel-shaped in most species, in color, all 

transitions frequently purple-blue to blue; stamens five ; pistil

390 


bicarpellate, syncarpous, embryologically two-celled with two ovules 

in each cavity, but dorsal ingrowths divide ovary by time of flowering 

into four cells with one ovule in each cavity; style gynobasic. Fruit 

typically four-nutlets. Seeds solitary in each cavity and either 

scantily albuminous (Heliotropes) or exalbuminous (Borraginea) . 

PIG. 228. Hyoscyamus niger Flowering branch. (Sayre.) 

Unofficial Part used Botanical origin Habitat 

Symphytum 'Root 

(Comfrey) 

Cynoglossum Herb and root 

(Hound's tongue) 

Alkanet Root 

Symphytum officinale Europe and 

United States 

Cynoglossum 

United States 

officinale 

Alkanna tinctoria So. Europe and Asia 

Solanacea or Nightshade Family. Stem herbaceous, rarely shrubby 

or arborescent, frequently with bicollateral bundles. Leaves alter-

TAXONOMY 391 

nate, exstipulate, entire or more or less lobed, rarely compound; 

often glandular-hairy. Flowers in cymes ; regular or rarely irregular 

(Petunia, Tobacco sps.), pentamerous, perfect, synphyllous; sepals 

green (rarely petaloid), rotate to tubular, usually persistent and 

accrescent; petals rotate (Solanum), to tubular (Atropa), to funnelshaped 

(Tobacco), and so (i) open to all comers, or (2) to bees or 

wasps, or (3) to butterflies, moths; color, greenish-yellow, or 

greenish-white, to white, to pink, crimson, purple, rarely blue; stamens 

five, epipetalous, hypogynous, along with style usually forming 

nectar glands. Filaments short to long, anthers dehiscing longitudinally 

or by apical pores; pistil bicarpellate, syncarpous, with 

or without nectar girdle; superior ovary, two-celled with central 

placentation, ovules numerous, style more or less elongate with 

bilobed or bifid stigma. Fruit, a capsule (Tobacco, Thornapple, 

Henbane) dehiscing longitudinally or transversely; or a berry 

(potato, egg-plant, tomato, red pepper). Seeds albuminous. 

Official drug


Scorphulariacea or Figwort Family. Herbs (Linaria, Verbascum, 

Gerardia, Digitalis, etc.), shrubs (shrubby Veronicas, etc.), rarely 

trees (Paulownia imperialis). Stem, branches and leaves usually 

green and independently vegetating, but in Pedicularis, Gerardia, 

Euphrasia, Buchnera, Rhinanthus, etc., the stem, leaves, and 

branches are condensed from the development of a parasitic root 

habit. Stems cylindrical to frequently quadrangular, especially 

when leaves are opposite. Leaves alternate to opposite and decus- 

sate, simple, exstipulate, often hairy, but becoming by drought or 

FIG. 229. Nightshade, or bittersweet (Solatium Dulcamara). (Gager.) 

parasiticism reduced to scales or almost absorbed. Inflorescence a 

raceme of cymes (Paulownia) or a simple raceme (Foxglove, Linaria, 

often a 

etc.) or spike (Verbascum Thapsus) or, if leaves are opposite, 

whorl of axillary flowers or solitary axillary flowers. Flowers rarely 

regular, mostly irregular; calyx of five sepals condensed in Veronica 

to four through absorption of one sepal by fusion of two sepals; 

corolla of five to four petals, deeply synpetalous, varying from rotate 

(Verbascum Blattaria, etc.) to irregular tubular to elongate, irregular 

bilabiate to funnel-shaped. In color, corolla varies from greenish

TAXONOMY 393 

to greenish-yellow or white (Scrophularia) to pure white or from red 

to purple to blue (Veronica). Stamens five, fertile, equal in length 

in a few Verbascum species or unequal in other Verbascum species to 

stamens four with a long sterile staminode (Pentstemon) to four 

didynamous stamens with a short petal oid staminode (Scrophularia) 

to four didynamous stamens with a minute often nectariferous 

staminode (Linaria), to frequenty four didynamous stamens only, 

the two lateral or two anterior stamens stronger and longer (Antirrhinum) 

to two perfect stamens and two minute staminodes 

(Calceolaria) to two stamens alone developed (Veronica). 

Pistil bi- 

carpellate; ovary two-celled with central placentation; style terminal 

with bilobed stigma; ovules numerous, small. Fruit a two-celled 

and usually many-seeded capsule. Seeds richly albuminous, 

anatropous or amphitropous. 

Official drug

394 


PIG. 230. Digitalis purpur ea, var. gloxinaeflora.

TAXONOMY 395 

more rarely whorled, entire, exstipulate. Inflorescence a raceme of 

condensed cymes, becoming a simple raceme or spike, rarely con- 

densed into a solitary terminal inflorescence. Flowers hermaphro- 

dite, usually irregular; calyx five-cleft; corolla hypogynous, 

gamopetalous, more or less bilabiate, funnel-form and composed of 

five sepals; stamens usually four (Ruellia, etc.), occasionally reduced 

to two, as in genus Dianthera, didynamous or diandrous, epipetalous; 

with numerous cam- 

pistil bicarpellate ; ovary two-celled, superior, 

pylotropal ovule's; style terminal, filiform. Fruit a capsule contain- 

ing numerous curved seeds. The family is of pharmaceutic interest 

mainly because of Ruellia ciliosa, a pubescent perennial herb growing 

in th6 Eastern United States, whose rhizome and roots have fre- 

quently been admixed with or substituted for Spigelia. 

Verbenacea or Vervain Family. Herbs (Verbena), Shrubs (Clarodendron), 

rarely trees (Tectona or Teak-wood) whose stems and 

branches are usually quadrangular and rarely scented. Leaves 

generally opposite, exstipulate, simple or compound. Inflorescence 

a terminal panicle of spikes (Verbena hastata), a cyme (Callicarpa) 

or head (Lippia lanceolata). Flowers white, pink or blue (Verbena 

haslata) irregular, more or less 2-lipped ; calyx gamosepalous, 

tubular; corolla gamopetalous, hypogymous with a 4-5 fid limb; 

stamens generally 4, didynamous and inserted on the corolla tube 

or throat; pistil of 2-4 carpels, a terminal style and undivided stigma. 

Fruit a drupe or 2 to 4 celled berry, usually splitting into as many 

nutlets. Seeds exalbuminous. 


Verbena N. F. 

' 

Overground portion Verbena hastata United States 

Labiates, (Lamiacea) or Mint Family. Herbs producing creeping 

runners that spread out and root at the nodes. Stems quadrangular, 

rarely cylindrical in outline. Leaves opposite, decussate, mainly 

petiolate; leaf margin nearly always serrate, dentate or crenate. 

Stems and leaves further characterized by the presence of glandular 

hairs containing aromatic volatile oil. These hairs consist of a 

short one-celled stalk and a head (gland) of six or eight cells. Inflo- 

rescence a raceme or spike of verticillasters (double dichesial cymes) 

or, as in the Ground Ivy, a reduced verticillaster. Flowers typically

396 


pentamerous, rarely tetramerous; sepals five, synsepalous, ribbed 

and forming a tubular regular or irregular bilabiate (Scullcap, etc.) 

calyx whose upper lip is bifid and lower trifid; corolla of five to four 

gamopetalous petals, hypogynous, frequently two-lipped, the upper 

lip bifid, the lower trifid; stamens four, didynamous, rarely one pair 

alone fertile and the other pair reduced, in some cases almost or 

FIG. 231. Mentha piperita Flowering branch. (Sayre.) 

quite to disappearing point (Salvia and Monarda) ; pistil bicarpellate, 

embryologically two-celled with two ovules in each cavity, becoming, 

at time of flowering, four-celled with one ovule in each cavity. Style 

embryologically terminal, but, upon opening of flower, deeply gynobasic, 

elongate, slender with two stigmatic surfaces. Fruit four 

nutlets enclosing as many exalbuminous seeds.

Official drug 

TAXONOMY 397

398 


V. Order Rubiales. Rubiacea or Madder Family. Herbs 

(Galium, Mitchella, etc.), shrubs (Cephalanthus, etc.), or trees (Cinchona 

species) with fibrous roots, sometimes, as in Cephaelis Ipecacuanha, 

annularly enlarged. Roots, stems and to a less extent leaves 

rich in varied alkaloids, some of medicinal value. Leaves opposite, 

entire, stipulate and interpetiolate. Inflorescence a raceme of 

dichesial cymes occasionally condensing to scorpioids. Flowers 

perfect, often dimorphic, pentamerous or tetramerous; sepals five 

(Cinchona, etc.) but four in Galium, small, green, subtended with 

other flowers by one or two or more enlarged petaloid bracts; petals 

five (Cinchona, etc.) to four in Galium, stellate, varying from shallow 

rotate to elongate tubular or funnel-shaped with stellate limbs; 

stamens five to four, epipetalous; pistil nearly always bicarpellate, 

rarely of five to four carpels; ovary inferior, two-celled with central 

placentation; styles either distinct with knob-shaped stigmas or style 

elongate, filiform, ending in bilobed stigmas. Fruit varied, a capsule 

in Cinchona, a berry in Coffee, a drupe, or frequently, as in Galium, 

dry and splitting into nutlets; seeds albuminous, each with a curved 

embryo. 

Official drug 

Caffeina 

Cinchona 

Official drug 

Cinchona Rubra 

Part used 

P'eebly basic 

principle 

Bark 

Part used 

Bark 

Coffea Tosta N.F. Roasted seeds 

Gambir 

Ipecacuanha 

Prepared extract 

from de- 

coctions of leaves 

and twigs 

Root 


Coffea arabica 

Cinchona Ledgeri- 

ana, C. Calisaya 

and hybrids of 

these with other 

Cinchona species 


Cinchona succirubra 

or its hybrids 

Coffea arabica 

Coffea liberica 

Habitat 

Eastern Africa 

South America 

Habitat 

> South America 

Eastern Africa 

Ourouparia Gambir East Indies 

Cephaelis 

Ipecacuanha 

Cephaelis acuminata 

Brazil 

United States of 

Columbia

TAXONOMY 399 

Caprifoliacea or Honey Suckle Family. Shrubs or rarely herbs. 

Leaves entire, opposite, exstipulate or with delicate, attenuate or 

filiform stipules. Inflorescence varying from a raceme of shortened 

cymes to a capitulum. Flowers varying from regular and small 

PIG. 232. Cepha'&lis ipecacuanha Plant and dried root. (Sayre.) 

(Sambucus, Viburnum, etc.) to increasingly large, slightly irregular 

and ultimately very irregular in some Loniceras and in a few Weigelas 

and allies; calyx pentamerous, superior; corolla superior, gamopetalous, 

limb pentafid, small in Viburnum and Sambucus to

4OO 


elongate, tubular or irregular infundibuliform in Loniceras; stamens 

five, inserted on tube of corolla and alternating with corolla seg- 

ments; filaments equal or didynamous (in irregular flowers); ovary 

inferior, rarely five- to three-celled, usually three- or frequently two- 

celled; style terminal. Fruit a berry (Viburnum) from an inferior 

ovary, several celled, occasionally becoming one-celled with several 

to rarely one seed, or fruit a capsule (Diermlla, Weigelia). Seeds 

albuminous. 

Official drug 

Sambucus N.F. 

Viburnum 

Prunifolium 

Flowers 

Bark 

Viburnum Opulus Bark 

N.F. 

Part used 

FIG. 233. Colocynth Portion of 

vine and whole fruit. (Sayre.) 


Sambucus canadensis 

Sambucus nigra 

Viburnum . 

prunifolium 

Viburnum Lentago 

Viburnum Opulus 

var. Americanum 

~ 

Habitat 

} United States 

/ Europe 

Eastern and 

} 

central United 

States 

United States 

and- Canada 

VI. Order Campanulales. 

Cucurbitacea or Gourd Family. 

Herbaceous, very often annual 

(Colocynth, etc.), more rarely 

perennial (Bryonia, etc.), sometimes 

shrubby plants, the peren- 

nial and shrubby forms perennat- 

ing by swollen roots, some of 

which are heavy and tuberous. 

Stems very usually grooved and 

ridged, often provided with 

and barbed hairs. 

roughened 

Tendrils are frequently produced 

in the axils of leaves from tendril 

axillary buds (Pumpkin, Colo- 

cynth, Watermelon, Cucumber, 

Bryony, etc.). Leaves varying 

from entire, simple, 'usually del- 

toid to triangular through stages 

of trilobate, pentalobate, deeply palmatifid to palmatipartite to seldom 

approaching compound (Colocynth). Venation in nearly 

all cases

TAXONOMY 401 

palmate. Leaves thin, herbaceous, much expanded, often hairy. 

Vascular bundles of petioles, branches and stems, bicollateral. 

Inflorescence either of loose cymes or more frequently racemes or 

spikes or entire axillary inflorescence may become solitary axillary. 

Flowers pentamerous, very rarely tetramerous, monoecious (Bryonia 

alba) or dioecious (Bryonia dioica) ; sepals five, gamosepalous, adnate 

to ovary; corolla of five, rarely four gamopetalous petals varying 

in size and shape from small to large campanulate or broadly cup- 

shaped (Cucumber) and in color from greenish-yellow to greenish 

white to pure yellow to yellowish- white to white ; stamens typically 

five, epigynous, with anthers either joined by pairs or synantherous; 

carpels usually three; ovary inferior, one- to three-celled. Fruit a 

pepo (a berry from an inferior ovary with thick skin). Seeds flat 

and exalbuminous. 

Official drug 

Bryonia N.F. Root 

Colocynthis 

Pepo 

Elaterinum 

Unofficial 

Part used 

Pulp of fruit 

Seeds 

Watermelon Seed Seeds 

Momordica (Bal- Fruit 

sam apple) 

Principle from 

elaterium 


Habitat 

Bryonia alba 1 Europe 

Bryonia dioica 

j 

Citrullus Colocyn- Africa and Asia 

this Tropics 

Cucurbita Pepo 

(cultivated varie- 

ties) 

Ecballium 

Elaterium 

Citrullus vulgaris 

Momordica 

Balsamina 

Mediterranean 

region 

Southern Asia 

East Indies 

Campanulacea or Bluebell Family. Herbs of annual or more commonly 

perennial growth rarely sub-shrubby or sub-woody in habit, 

frequently with laticiferous tubes containing a milky juice. Stem 

upright or feeble and spreading. Leaves alternate, simple, exstipulate. 

Inflorescence primitively a racemose cyme condensing into a 

raceme, to a sub-capitulum and ultimately to a capitulum. Flowers 

regular, campanulate to campanulate-elongate to elongate and deeply 

cleft in petals; sepals five, only slightly synsepalous, epigynous; petals 

five, campanulate to campanulate- tubular to tubular elongate 

to tubular and deeply cleft; corolla varying in color from greenish- 

26

402 


yellow to yellowish-white to white or again, from yellowish-purple 

(rarely through yellowish-pink or red) to purple to pure blue; 

stamens five, epigynous, usually free from corolla; nectary epigynous; 

pistil usually tricarpellary; ovary as many celled as number of 

carpels and with central placenta; style single elongate; stigmas as 

many as carpels. 

contain inulin. 

Fruit a capsule. Seeds albuminous. The plants 

Lobeliacece or Lobelia Family. Herbs, with inulin and latex con- 

tents, corresponding with Campanulacece in their vegetative parts, 

but differing from that group by having irregular flowers (pale blue 

in Lobelia inflata), anthers always synantherous and pistil always 

bicarpellate with two-celled ovary and bilobed or 'bilabiate stigma. 


Lobelia Leaves and Lobelia inflata United States and 

flowering tops 

Canada 

VII. Order Aggregatse. Valerianacece or Valerian Family. 

Herbaceous often low succulent plants with creeping rhizomes, fre- 

quently strongly scented and possessing stimulating properties. 

Leaves frequently dimorphic; radical fascicled; cauline opposite; 

petiole dilated, exstipulate. Inflorescence a raceme of dichesial or 

scorpioid cymes. Flowers more or less irregular; calyx absent as 

such, but represented by a series of teeth that are incurved in the 

bud and flower and which expand later into a pappose crown and act 

in the fruit as a pappose disseminator; corolla pentamerous, gamopetalous, 

varying from rotate synpetalous to irregular tubular with 

petals diversely united, in color varying from greenish-white to 

white or pink (Valeriana officinalis) to crimson; stamens three to 

two or one (Valerian), epipetalous; pistil syncarpous; ovary usually 

one-celled, inferior; style filiform with three stigmatic surfaces. 

Fruit an akene from inferior ovary crowned by a persistent ex- 

panded pappose calyx rudiment. Seeds anatropous, exalbuminous. 


Valeriana Rhizome and Valeriana officinalis Europe and Asia 

roots 

Composite (Asteracea) or Daisy Family. Herbs, rarely shrubs or 

trees, of annual or perennial habit, and with watery or milky juice.

TAXONOMY 403 

Inulin is present in cell sap of parenchyma. Leaves alternate, rarely 

opposite, simple to compound, exstipulate. 

Inflorescence a capitu- 

PiG. 234. Valeriana officinalis Plant and rhizome. (Sayre.*) 

lum or a raceme of capitula, each capitulum surrounded by an in- 

volucre or protective whorl of bracts, and composed of numerous

404 


florets that may be : (a) wholly regular, tubular and hermaphrodite 

(Thistle, etc.); or (b) central florets as in (a), but marginals strapshaped 

or ligulate and usually pistillate (Daisy, Dahlia, etc.); or 

(c) florets all ligulate and hermaphrodite (Dandelion, Chicory, etc.) ; 

or (d) florets in part or in whole bilabiate (Mutisia, etc.). Flowers 

__ 6fy b 

^ ^pappus 

receptacle 

I c/v 

-ovary 

'sca\e 

- 

ovary vw// 

ovule 

FIG. 235. Capitulum of a composite Jerusalem artichoke (Helianthus 

tuberosus). A, lengthwise section of capitulum, X i; B, ray flower, X 6; C, disk 

flower, cut lengthwise, x 6. (A after Baillon, B and C, Robbins.) 

small (florets) closely crowded, pentamerous, shaped as above, with 

ovary inferior and other floral parts superior. Sepals rudimentary, 

tooth-like (Sunflower), or reduced to a pappose or hairy rudiment 

above ovary that is functionless during flowering, but that expands in 

fruit as a hairy fruit disseminator (Dandelion, Thistle, etc.); or 

sepals wholly absorbed (Daisy). 

Petals synpetalous, tubular, ligu-

TAXONOMY 405 

late or rarely bilabiate, greenish-yellow to white, or through pink- 

crimson and purple to blue (Chicory). Stamens five, epipetalous, 

filaments distinct, anthers united into an upright anther-box (so 

synantherous) into which pollen is shed before or during opening 

FIG. 236. Matricaria chamomilla Branch and dissected flowers. (Sayre.) 

of each floret. Carpels two, syncarpous, ovary inferior, one-celled 

with single ovule; style simple, at first short, later elongating and by 

collecting hairs sweeping pollen to top of anther box, then dividing 

into two stigmatic surfaces with stigmatic hairs for pollen reception.

406 


Fruit an indehiscent achene often (Dandelion, Thistle) crowned by 

the pappose, calyx rudiment. Seed single, exalbuminous. 


Lactucarium


Part used 

Santonica Unexpanded 

flower heads 

Carthamus Tubular florets 

Achillea 

Tanacetum 

Gnaphalium 

Cichorium 

Leaves and 


Leaves and 


Leaves and 


Rhizome and 

roots 

Oleum Erigerontis Volatile oil 

TAXONOMY 407 


Artemisia pauciflora Russian 

Habitat 

Turkestan 

Carthamus India 

tinctorius 

Achillea millefolium Europe and Asia 

Tanacetum vulgare Europe 

Gnaphalium 

polycephalum 

North America 

Cichorium Intybus Europe 

Erigeron canadensis North America 

FIG. 237. Chicory (Cichorium Intybus). A, portion of flowerirg branch; B, 

basal leat (runcinate-pinnatifid) ; C, median longitudinal section through a 

head, showing the insertion ot the flowers; D, individual flower; E, fruit (ripened 

ovary), showing the persistent pappus (calyx) of short scales. (Gager.)

CHAPTER IX 

ECOLOGY 

Ecology is x that department of biology which deals with the 

relations of plants and animals of various habitats to their environ- 

mental conditions. Every living thing is a creature of circumstance, 

dominated and controlled by heredity and environment. In order 

to exist and keep healthy it must adapt itself to the various factors 

of its surroundings. The environmental factors having to do with 

the existence and health of plants include soil constituents, air, 

moisture, light, range in temperature, gravity, surrounding animals 

and plants of other kinds. 

A group of plants occurring in a common habitat constitutes what 

is termed a plant association or society. Plant associations may 

be classified either from the point of view of their order of develop- 

their water 

ment, as based upon the principle of succession, or upon 

relation. The latter method, appears to be the one more generally 

adopted, because of its ready application and will now be considered. 

According, therefore, to the relation plant associations have 

assumed in regard to water, they may be grouped as follows: 

1. Hydrophytes or water plants. 

2. Helophytes or marsh plants. 

3. Halophytes or salt plants. 

4. Xerophytes or desert plants. 

5. Mesophytes or intermediate plants. 

6. Tropophytes or alternate plants. 

Hydrophytes. The effect of an aquatic environment on the structure 

of water plants is most striking. The root systems are reduced 

both in length and number of branches. The root hairs of those 

immersed in the water are absent. The supportive action of the 

water is such that the fibrovascular elements of the stems, which 

usually function both for support and conduction of crude sap, 

are greatly reduced in size and strength. The leaves, stems and 

roots possess large air-spaces. The mesophyll of the leaves is 

408

ECOLOGY 409 

spongy and the chloroplasts motile. Stomata are entirely absent 

from leaves that are submerged and only present on the upper sur- 

face of floating ones, where they are nearly always open. Some of 

these plants have broad floating leaves and dissected submerged 

ones, often with thread-like divisions. The submerged parts are 

devoid of special protective walls e.g. those containing cutin or 

suberin. The cell sap has a low osmotic pressure. The submersed 

leaves often absorb more water than the roots. The free floating 

microscopic plants (blue-green algae, bacteria, diatoms, desmids, etc.) 

form the plankton of our ponds, rivers and lakes. The free-swim- 

ming higher plants (the pleuston) comprise certain liverworts 

like Riccia and Ricciocarpus, water-ferns and such seed plants as the 

water-lettuce and water-hyacinth. The aquatic plants including 

the algae, mosses and flowering plants, which live attached to rocks 

comprise the lithophilous benthos. Another class of aquatic plants 

(benthos) include those with true roots, which attach the plant to 

the substratum, and at most possess floating leaves. This type 

includes the water-lilies, the water-chestnut, the splatter docks, 

the floating-heart and the pondweeds. 

Helophytes. To this group belong plants typical to marshes. 

A marsh is an area with wet soil, wholly or partially covered with 

water and with annual or perennial herbs (never shrubs and trees) 

the usual 

which are adjusted structurally to a mucky soil, lacking 

supplies of oxygen. These plants likewise show an adjustment to a 

partial or periodical submergence. Like hydrophytes, marsh plants 

are for the most part perennial. They produce adventitious roots 

and possess horizontal rhizomes, or runners, and frequently have 

air chambers in roots, stems and leaves, so that they are adapted 

to meet the scarcity of air in wet soils. They also show a striking 

development of erect chlorophyll-bearing organs in the shape of 

leaves, in the flags, and stems, in the rushes. 

The taller seed-like plants of the marsh-land, such as seed-grass 

(Phragmites), the bur-reed (Sparganium), the cat-tails (Typha), the 

blue flags (Iris), the sweet flag (A corns calamus) and the papyrus 

(Papyrus) form associations known as fresh-water marshes, reedmarshes 

or fens. The channels or pools of water in amongst these 

amphibious plants are filled with true aquatic plants.

410 


Halophytes. The plants of this group live in a soil which is rich 

in soluble salt, usually common salt (NaCl), and on account of the 

fact that the osmotic force of the root is nearly inadequate to overcome 

that of the concentrated solution of the soil, the soil to such 

plants is physiologically dry. A halophyte in fact is one form of 

xerophyte. The most striking feature among halophytes is that 

they are nearly all succulent plants. The leaves of such plants, for 

example, are thick, fleshy and more or less translucent. They are 

rich in concentrated cell sap by which they are able to counteract 

the osmotic pull of the concentrated saline solution of the soils 

in which they live. Anatomically they are poor in chlorophyll, 

the intercellular-air-spaces are small and the palisade tissue is 

more abundant. Coatings of wax are found and a hairy covering, 

although infrequent, sometimes occurs. Coriaceous and glossy 

leaves, especially in tropical halophytes, are noteworthy, while 

in many salt-loving plants the stomata are sunken. Halophytes 

are found in our coastal salt marshes and on saline tidal flats in tem- 

perate and tropical countries and on the alkali flats of the interior 

of continents. Notable examples of these plants are the Salt Marsh 

Samphire, Salicornia ambigua, the Mangroves (Rhizopora) and the 

Bald Cypress (Taxodium). 

Xerophytes. The plants of this group, like the halophytes, are 

adjusted to live in a soil which is physiologically dry. The soil may 

owe this condition to its physical nature, such as porosity (sand), 

or to the presence of humic acids, or by chemical action, which in- 

hibits the absorption of water. They are adapted to meet the con- 

ditions of strongest transpiration and most precarious water supply. 

To meet such conditions of physiological drought, the plants show 

various structural adaptations. In deserts, where the atmospheric 

precipitations are less than a certain limit, the plants acquire a 

xerophytic structure, such as succulency, water storage tissue, 

associated frequently with mucilage, lignified tissues, thick cuticle 

to the leaves depressed, stomata (frequently in pits), reduced transpiration 

surfaces and thorns. Mechanical tissues like wood and 

bast fibers attain their highest development in these plants. Cacti 

and the century plant (Agave) are types of xerophytes while many 

bog plants like the cranberry and Laborador tea, with leathery leaves, 

are xerophytic.

ECOLOGY 411 

Mesophytes. These are plants that grow in soil of an intermediate 

character which is neither specially acid, cold or saline, but is 

sufficiently well supplied 

with water and rich in the elements re- 

quired for plant growth. Plants which grow under such conditions 

do not have structures by which transpiration is closely controlled. 

They have large leaves frequently toothed and incised, with numerous 

stomata usually on the lower surface and small intercellular -air- 

spaces. The leaves and stems are usually of a fresh green color. 

Typical of the mesophytes are the grasses and most of the annual 

and biennial herbs of temperate regions. 

Tropophytes. This term was first introduced by Schimper in 

1898 for land plants which have deciduous leaves and whose condi- 

tions of life are, according to the season of the year, alternately those 

of mesophytes and xerophy tes. The mesophy tic condition is found in 

summer, when the trees, shrubs and perennial herbs, included in this 

group, are in full leafage, and when, owing to the regular supply of 

rain during the growing season, the soil is plentifully supplied with 

water to meet the demands of -these plants during the period of active 

transpiration. During the winter they are xerophytes. The 

cold of winter freezes the water in the soil so that the transpiration 

is reduced to a minimum, and this is associated with the fall of the 

leaves of the trees and shrubs and the death of the overground parts 

of the perennial herbs which spring up each year from their under- 

ground parts. The vegetation of cold temperate regions is mainly 

tropophytic. 

The deciduous trees and shrubs also known as the broad-leaved 

plants and the summer-green plants form the principal tropophytes. 

The deciduous forests, which include the oaks, the beeches, the 

ashes, the maples, the walnuts, the chestnuts, cover a great part of 

eastern and western China, central Europe (England, France, 

Belgium, Germany) and eastern Australia, and are coincident 

with the countries occupied by the most civilized races of man, such 

as the Americans, Europeans, Chinese and Japanese. The cold 

temperate climatic conditions which have determined the distribution 

of the forest trees have been influential also in the development 

of the energetic races of mankind.

GLOSSARY 

Abor'tion. The imperfect or non-development of an organ. 

Acaules'cent. Without an obvious aerial stem. 

Achene' (akene). A small, dry, one-celled indehiscent fruit in which the seed 

coat and pericarp (fruit wall) are not firmly attached. 

Achlamy'deous. Destitute of calyx and corolla. 

Acic'ular. Applied to crystals of calcium oxalate, etc. that are slenderly needle- 

Shaped. 

Acrop'etal. Development from outside (below) toward the inside (above). 

Acu'minate. Tapering gradually to a long point. 

Acute'. Sharp-pointed, the point being less than a right angle. 

Ad'nate. Applied to the growing together of unlike parts. 

Adventi'tious. Applied to roots and buds that are out of their ordinary position. 

Aestiva'tion. Arrangement of the parts of the flower in the bud. 

Albu'men. Nutritive material stored in the embryo, endosperm, or perisperm. 

Al'ternate. Applied to leaves, buds, etc. that are arranged singly (one after 

another) at the nodes. 

Albur'num. Sapwood. Am'ent. A scaly spike-like inflorescence. Another name for catkin. 

Amor'phous. Without definite shape. 

Amphit'ropous (ovules and seeds). Half-inverted and straight, with the 

hilum about the middle, and micropyle terminal. 

Amplex'icaul. Clasping the stem. 

Anal'ogy. Resemblance in function. 

Anastomo'sing. Applied to veins that are connected with others by cross veins, 

so forming a network, as with the marginal veins of Eucalyptus. 

Anat'ropous. Inverted ovules or seeds with micropyle adjacent to hilum. 

Androe'cium. The male system of organs in a flower. 

Androg'ynous. Applied to inflorescences composed of both staminate and 

pistillate flowers. 

Anemophi'lous. Wind pollinated. 

Angiospenn'ous. Having ovules and seeds borne within a box-like covering, 

the pericarp. 

An'nual. Producing flowers, fruit and seed within a year from the time the 

seed germinated and then dying completely. 

An'nular. Ring-like. 

Ante'rior. The front region. 

An'ther. That portion of a stamen which bears, the pollen. 

An'theridium. Male sexual organ of Thallophytes, Bryophytes and Pterido- 

phytes. 

412

GLOSSARY 413 

An'therozoid. A male sexual cell formed within an antheridium. 

An'thophore. A lengthened internode of the receptacle between calyx and 

corolla. 

Apet'alous. Without petals, as in the oaks, etc. 

Apocar'pous. Carpels separate and distinct. 

Apopet'alous. Petals separate and distinct. 

Aposep'alous. Sepals separate and distinct. 

Archego'nium. A multicellular female sexual organ. 

Ar'il. An accessory seed covering outside of the testa and arising at or about 

the hilum, as in Euonymus. 

Ar'illode. A fake accessory seed covering outside of the testa, as in Nutmeg, 

and arising from the dilatation of the micropyle. 

Aris'tate. Having a stiff bristle-like termination. 

. Auric'ulate. 

Ascend'ing. Growing obliquely upward. 

As'cus. Spore case of an Ascomycete fungus. 

Atavism. Reversion to ancestral type. 

Ear-like. 

Awn. A bristle-like structure that branches along its axis. 

Ax'il. Angle formed by branch, leaf or bud with the stem. 

Ax'illary. 

In the axil. 

Bac'cate . B er ry-lik e . 

Barb. A short bristle usually bent back. 

Bast. Applied to the phloem region but mainly to the fibrous portion thereof. 

Beard'ed. Furnished with long hairs. 

Ber'ry. A fleshy fruit wh'ose mesocarp and endocarp are fleshy and frequently 

succulent throughout, and with seeds imbedded therein, as tomato, capsi- 

cum, belladonna, etc. 

Bi. A prefix of the Latin language indicating two, twice or doubly. 

Bien'nial. Applied to plants that live for more than one year but not longer 

than two years. 

Bila'biate. Two lipped. 

Blade. Expanded part of a leaf. 

Bloom. The whitish and waxy secretion of epidermal cells, as in the stems of 

Sugar Cane or the leaves .of Cabbage. 

Bract. A modified leaf, frequently scale-like, appearing on inflorescence axes. 

Brac'teole (bracteolar leaf). A modified leaf found on pedicels. 

Bud. A rudimentary stem. 

Bulb. A very short scaly underground stem. 

Bul'bils. Small underground bulbs, as in garlic. 

Bul'blets. Small above ground bulbs, as in the tree onions. 

Cadu'cous. Falling with the opening of the flower, as the calyx of Papaver. 

Ca'lyx. The outermost whorl of floral leaves.

414 


Cam'bium. The growing meristematic layer of a vascular bundle. 

Campan'ulate. Bell shaped. 

Campylo'tropous. Applied to ovules or seeds that are curved so as to. bring the 

apex and base near together. 

Canes'cent. White or gray from a coating of fine hairs. 

Capiliit'ium. A network of filaments among spores, as in slime molds, puff 

balls, etc. 

Cap'itate. Shaped like a head. 

Caprifi cation. The process of pollinating figs artificially. 

Cap'sule. A dry, dehiscent fruit of two or more carpels. 

Car'pel. A transformed leaf bearing one or more ovules, a simple pistil; a part 

of a compound pistil. 

Car'pophore. A slender stalk, the prolongation of the receptacle, to which 

the inferior akenes (mericarps) of the Umbelliferce are attached. 

Caryop'sis. A dry, indehiscent, one seeded fruit of the grasses or cereals in 

which the fruit wall (pericarp) and seed coat firmly adhere. 

Cat'kin. A scaly spike of flowers. 

Cau'date. Tailed. 

Caules'cent. With an obvious aerial stem. 

Cau'line. Pertaining to the stem. 

Centrifugal. Applied to a flower cluster in which the terminal or central 

flower blossoms first. 

Centrip'etal. Applied to a flower cluster in which the lower or outer flowers 

bloom first. 

Chaff. The glume? and palets of grains; the scaly hairs on the stipes of ferns; 

the bracts subtending each floret in some heads "of Compositae. 

Chala'za. That portion of the ovule marked by the junction of the integuments 

with the nucellus. 

Chasmo'gamous. Pertaining to flowers that regularly open. 

Chlamy'dospore. Thick walled spore formed within the hyphae of smuts. 

Chlo'rophyll. The green coloring matter of all green plants. 

Chloroplas'tid. A protoplasmic body in the cells of green parts of plants con- 

taining chlorophyll. 

Chro'matin. That portion of the nucleus which is readily colored by a basic 

dye. The substance that carries the hereditary characters from parent to 

offspring. 

Chromoplas'tid. A protoplasmic body in the cells of certain parts of plants 

containing a pigment other than chlorophyll. 

Chro'mosome. One of the bodies into which the chromatin of the nucleus is 

resolved during indirect nuclear division. 

CiTia. Vibratory hair-like protoplasmic outgrowths of zoospores, bacteria, 

gametes, etc. 

Circumnuta'tion. The repeated bending in different directions of the growing 

tips of stems of climbing plants.

GLOSSARY 415 

Cir'cinate. Rolled inward from apex toward base, as the young leaves of ferns. 

Circumscis'sile. Applied to the splitting open of capsules transversely into 

lid and pot portions. 

Clad'ode. A flattened branch which somewhat resembles a leaf. 

Claw. The narrowed base of some petals, as those of the Pink Family. 

Cleistog'amous. Applied to flowers that never open but are self fertilized, as 

in some Polygalas and Violets. 

Coch'lea. A spirally coiled legume. 

Coe'nocyte. A multinucleate cell. 

Cohe'sion. The union of parts of the same whorl. 

CoTiort. A group of natural orders. 

Coleorhi'za. A root sheath. 

Collateral. Applied to fibrovascular bundles in which the phloem and xylem 

masses are arranged side by side. 

Collen'chyma. Tissue composed of cells thickened at their angles. 

Columeria. The end cell wall of an aerial hypha that bulges into the sporan- 

gium; also applied to the axis of a capsule. 

Col'umn. The united stamens and carpels in Orchids. 

Co'ma. A tuft of hairs, as found on the seeds of Milkweeds. 

Com'missure. The contiguous surfaces of two carpels as in the flowers and 

fruits of the Parsley Family. 

Concen'tric. Applied to several circles or whorls one within the other. Concentric 

fibrovascular bundles are those in which the xylem mass surrounds 

the phloem mass or vice versa. 

Concep'tacle. A sac bearing the fruiting organs in certain Algae and Fungi. 

Condu'plicate. Folded together lengthwise as for example the bud leaves of 

the oak or peach. 

Conid'ia. Asexual spores cut off from the ends of hyphae or sterigmata by 

Penicillium, Aspergillus, Peronospora, Claviceps, etc. 

Conid'iophore. A hypha bearing conidia. 

Conjugation. One of the sexual methods of reproduction where two like sexual 

cells unite to form a zygospore. 

Con'nate. Applied to parts that have grown together, 

as the bases of two 

opposite leaves. 

Connect'ive. The continuation of the filament of the stamen that connects the 

two lobes of the anther. 

Conni'vent. Brought close together; converging. 

Con'volute. Rolled lengthwise from one edge as the leaves in the buds of the 

Wild Cherry and Plum. 

Cor'date. Heart shaped. 

Coria'ceous. Leathery in texture. 

Corm. A solid, swollen, fleshy underground stem. 

Corol'la. The inner whorl of floral envelopes composed of petals.

416 


Coro'na. A crown like appendage in the throat of the corolla, as in the flowers 

of Narcissus and Silene^ 

Cor'tex. That region in dicotyl and gymnosperm roots of primary growth and 

in roots and stems of monocotyledons between epidermis and endcdermis, 

in dicotyl and gymnosperm roots of secondary growth or in barks between 

cork cambium and phloem. 

Cor'ymb. A flat topped or convex centripetal inflorescence with the lowermost 

pedicels the longest. 

Cos'ta. A rib. 

Cotyle'don. A seed-leaf of the embryo. 

Crem'ocarp. The peculiar fruit of Umbellifera, consisting of two inferior akenes 

(mericarps) separated from each other by a carpophore. 

Cre'nate. Applied to leaf margins having rounded teeth. 

Cren'ulate. The margin with fine rounded teeth. 

Crib'riforin. Sieve like. 

Cru'ciform. Applied to the corolla or the calyx of flowers, the parts of which are 

arranged in the form of a cross. 

Crusta'ceous. Applied to the thallus of a lichen that closely adheres to the 

substratum. 

Cryp'togam. A plant belonging to one of the divisions of the vegetable kingdom 

below the Spermatophytes. 

Crys'talloid. A protein body found in the aleurone grains of seeds or under- 

ground parts. 

Culm. A jointed stem of a grass or sedge. 

Cu'neate. Wedge-shaped. 

Cu'pule. Applied to the concave involucre enclosing the glans of an acorn but 

also to other cup shaped parts of plants. 

Cu'ticle. A thin covering of a waxy substance called cutin on the outer wall of 

epidermal cells. 

Cus'pidate. Tipped with a sharp rigid point. 

Cyme. A more or less flat topped determinate inflorescence. 

Cy'mose. Cyme-like. 

Cytol'ogy. The study of cells and their contents. 

Cy'toplasm. The cell protoplasm outside of the nucleus. 

Decan'drous. Having ten stamens. 

Decid'uous. Applied to leaves which fall in autumn, to plants bearing such 

leaves and to the calyx and corolla which fall shortly after blossoming before 

the development of the fruit. 

Dec'linate. Curved or bent downward. 

Decompound'. Several times 'compounded, as- the leaf-blades of Cimicifuga. 

Decum'bent. Erect at base, then lying on the ground, with the end rising. 

Decus'sate. Applied to opposite leaves when the pairs stand at right angles 

to each other along the stem.

GLOSSARY 417 

Dehis'cence. Splitting open. 

Deliques'cent. Applied to a tree whose trunk or main stem is lost in branches. 

Del'toid. Having the shape of the Greek letter A. 

Den'tate. Having broad acute marginal teeth pointing outward. 

Dentic'ulate. Finely dentate. 

Dermat'ogen. The generative tissue that gives rise to epidermis. 

Deter'minate. Applied to inflorescences on which flowering begins with the 

terminal bud, thus ending the elongation of the stem bearing the flowers. 

Diadel'phous. Applied to stamens whose filaments are united at their edges 

into two sets. 

Diageot'ropic. Applied to a plant organ that assumes a horizontal position. 

Dian'drous. Possessing two stamens. 

Di'astase. A ferment found in germinating seeds and fungal hyphae which 

changes starch into maltose. 

Dichlamyd'eous. Pertaining to flowers that possess both calyx and corolla. 

Dichog'amy. The maturation of one set of sexual organs before the other. 

Dichot'omous. Forked. 

Dic'linous. Pertaining to the stamens and flowers. 

carpels being found in separate 

Dicot'yle'don. A plant whose embryo possesses two seed leaves or cotyledons. 

Digitate. Referring to a compound leaf whose leaflets come off at the end of 

the petiole. 

Dimor'phism. Having two forms of flowers, one with long styles and short 

stamens, the other with short styles and long stamens; the occurrence of 

two distinct forms. 

Dipe'cious. Applied to species having two kinds of individuals, male and 

female. 

Dissect'ed. Cut deeply into numerous divisions. 

Dissep'iment. A partition separating cells in a compound ovary or fruit. 

Dis'tichous. Pertaining to the arrangement of leaves in two rows. 

Divi'ded. Segmented to the mid-rib or base. 

Dorsoven'tral. Having distinct upper and lower surfaces. 

Dor'sum. The back of an organ. The lower surface of a foliage or floral leaf. 

Down'y. Covered densely with soft hairs. 

Drupe. A one-celled, one-seeded fruit whose endocarp is stony. 

Drupe'let. A small drupe. 

Duct. A tubular element found in the xylem region of a fibrovascular bundle. 

Dura'men. Heartwood. 

E- or Ex-, A prefix meaning devoid of, outside of, or away from. 

Eccen'tric. Deviating from the center. Applied to the hila of starch grains 

which are outside of the center, also to woody plants which develop more 

rapidly on one side than on the other. 

Echin'ulate. Beset with small prickles or spines. 

7

4 1 8 PHARMACEUTICAL BOTANY 

Ech'inate. Beset with prickles or spines. 

Ec'toplasm. A clear layer of protoplasm just beneath the cell wall. 

Egg-Appara'tus. The ovum and two synergids at the micropylar end of the 

embryo sac. 

El'ater. An elastic spiral filament attached to the spores of some Liverworts 

and Horsetails and aiding in their dispersal when mature. 

Emar'ginate. Notched at the apex. 

Em'bryo. A rudimentary plant found wi f hin the seed. 

Embryol'ogy. The study of the embryo and its development. 

Em'bryo-sac. A large cell within the nucleus of the ovule in which the embryo 

is formed after fertilization. 

En'docarp. The inner layer of the pericarp. 

Endoder'mis. A layer of cells forming the innermost boundary of the cortex 

and surrounding the fibrovascular region. 

En'dogen. A Monocotyledon. 

Endogenous. Applied to the axes of Monocotyl plants that do not increase 

materially in diameter. 

En'dophyte. A plant which grows within the tissues of another. 

En dosperm. A mass of cells formed in the embryo sac of ovules a, they mature 

to form seeds. 

En'dospore The inner wall of a spore. 

Endothe'cium. A zone of one or more layers within the exothecium of an anther 

En'siform. S word-shaped. 

Entomoph'ilous. Insect pollinated. 

En'tophyte. See Endophyte. 

Ephem'eral. Lasting for a brief period (a day or so;. 

Epica'lyx. A whorl of bracts resembling the calyx but below it. 

Epi'carp. The outer layer of the pericarp. 

Epicot'yl. The portion of the embryo axis above the cotyledon or cotyledojis. 

Epider'mis. The outer covering layer of ceUs of plants, sometimes later replaced 

by cork. 

Epig'ynous. Applied to floral leaves that appear to be inserted upon the ovary. 

Epipet'alous. Upon the corolla. 

Ep'iphyte. An air plant. A plant growing on another plant but not necessarily 

nourished by it. 

Epithe'lium. A delicate layer of cells lining an internal cavity. 

Eq'uitant. Applied to leaves, as in Iris, when they all spring from a rhizome 

and are successively folded on each other toward their bases. 

Eryth'rophyll. The red coloring matter of leaves. 

Estiva'tion (Aestivation). The arrangements of the floral bud. 

organs in the flower 

Etae'rio. An aggregate fruit like the Raspberry or Blackberry, the product 

of a single flower, consisting of an aggregation of drupelets on a receptacle. 

E'tiolation. The bleaching of green parts of plants when kept in the dark for 

some time.

GLOSSARY 419 

Evolu'tion. The presumable theory that all forms of living things existing today 

have been derived from others previously existing, either by direct descent 

or by common ancestry. 

Exalbu'minous. Applied 

to a seed in which the nourishment is stored in the 

embryo during the growth of seed from the ovule stage. 

Excen'tric. See Eccentric. 

Excres'cence. A morbid outgrowth. 

Excre'tion. Getting rid of nitrogenous waste. 

Excur'rent. Applied to trees, the main stems of which do not disappear in 

branches but grow erect to the summit ending in a terminal bud. The 

opposite of Deliquescent. 

Exfo'liate. To shed layers of bark. To cast off layers of tissue. 

Ex'ine. The outer wall of a pollen grain. 

Ex'ocarp. The outer layer of the pericarp. 

Exog'enous. Applied to the axes of Gymnosperms and Dicotyledons which 

increase materially in diameter. 

Ex'ogens. Plants with exogenous axes. 

Exospor'ium. The outer wall of a spore. 

Exert'ed. Applied to stamens that protrude from the throat of the corolla. 

Exstip'ulate. Without stipules. 

Ex'tine. The outer coat of a pollen grain. 

Extrorse'. Applied to anthers which face outward, away from the gynrecium. 

Face. The free surface of an organ. 

FaTcate. Scythe or sickle-shaped. 

Fam'ily. A sub-division of an order. 

Farina'ceous. Starchy or mealy. 

Fas'cicle. A bundle or cluster. 

Fascic'ular. Belonging to a bundle. 

Fascic'ulate. Clustered. 

Fec'ula. The nutritive part of a cereal. 

Fer'tile. Producing fruit or reproductive organs. Applied 

to flowers which 

contain functionally active stamens and carpels. 

Fertiliza'tion. That method of reproduction characterized by the union of two 

dissimilar gametes. 

Fi'brous. Fiber-like. Referring to root systems composed of many slender 

rootlets. 

Fibrovas'cular Bun'dle. A stringy group of fibers, vessels and cells coursing 

through the various organs of the higher plants and serving for support 

and conduction of sap. 

FiTament. The stalk of a stamen; a thread like structure. 

Filamen'tous. Thread-like. 

Fil'ifonn. Thread-like. 

Fim'briated. Fringed.

420 


Fis'sion. A form of division in which the cell separates into two equal or nearly 

equal parts. 

Flagel'lum. A whip like protoplasmic outgrowth of certain organisms or of 

zoospores, serving as an organ of locomotion. 

Folia'ceous. Leaf-like. 

Fol'licle. A one chambered dry fruit that dehisces along one suture only. 

Fovil'la. The contents of a pollen grain. 

Frac'ture. The manner in which a root or other plant part breaks when sub- 

jected to sufficient pressure. 

Frond. The leaf of a fern. 

Fruit. A matured pistil, or ovarian portion thereof together with any closely 

adhering part. 

Fru'ticose. Shrubby. 

Fuga'cious. Falling off early. 

Fundamental Tis'sue. Ground-tissue. The tissue of plants through which 

the fibrovascular bundles course. 

Funic'ulus. The stalk of an ovule. 

Fur'cate. Forked. 

Fu'siform. Enlarged in the middle and tapering toward either end. 

Gal'balus. A berry-like cone, as in Juniperus, formed by the coalescence of 

fleshy scales. . 

Ga'leate. Helmet shaped. 

Gam'ete. A sexual cell. 

Gam'etophyte. The sexual generation. 

Gamopet'alous. Applied to a flower whose corolla is composed of petals which 

are more or less united at their edges. 

Gamosep'alous. Having the sepals more or less united at their margins. 

Gem'ma. An asexual bud-like structure found in the capules of Liverworts. 

Gemma'tion. The process of budding as seen in the yeasts. 

Gen'era. Plural of genus. 

Genic'ulate. Kneed. 

Geot'ropism. Response to the stimulus of gravity. 

Germina'tion. The sprouting of a spore or seed. 

Germ Cell. A reproductive cell as distinguished from a somatic or body cell. 

Gills. The spore bearing plates of a toadstool. 

Gla'brous. Smooth. 

Gland. A secreting structure. 

Glans. A nut. 

Glau'cous. Covered with a bloom. 

Glo'boids. Small granules of calcium-magnesium phosphate found in aleurone 

grains 

Glob'ular. Spherical. 

Glom'erule. A head-like cyme.

Glume. A floral bract of the grasses and sedges. 

Glu'ten. The proteid matter of cereals. 

GLOSSARY 421 

Gonidium. Applied to the algal cells in lichens as well as to many forms of 

asexual reproductive bodies in flower-less plants 

Gon'ophore. An upgrowth of the receptacle between the corolla and stamens, 

as in Passi flora. 

Gynoeci'um. 

Gyn'ophore. 

The female sexual system of a flower. 

An upgrowth of the receptade between gynoecium and andrcecium 

as in Geum. 

Gynoste'mium. The united stamens and style. The column of orchids. 

Hab'itat. The original home o f a plant. 

Has'tate. Shaped like the head of a halberd, the basal lobes diverging. 

Head. An indeterminate form of inflorescence, as seen in the Daisy family, in 

which the flowers are in a dense duster on the receptacle. 

Heliot'ropism. Response to the stimulus of light. 

Herba'rium; A classified collection of dried plant specimens. 

Hermaph'rodite. Applied to flowers which contain both sets of essential organs, 

not necessarily functionally active. 

Hesperid'ium. A large thick-skinned succulent fruit like the orange, lemon or 

grape-fruit. 

Heterocyst. A large cell, occurring in the filaments of Nostoc. 

Heterophyl'lous. Having more than one kind of foliage-leaves on the same 

plant. 

Heteros'porous. Producing asexual spores of more than one kind as in Selagi- 

nella and the rusts. 

Hex. A prefix of Greek origin meaning six. 

Hexag'ynous. Having six carpels or styles. 

Hexam'erous. Having the parts of the flower in 6's. 

Hexan'drous. Having six stamens. 

Hibernation. Passing the winter in a dormant state of existence. 

Hi'lum. The scar of a seed, after the stalk of the ovule has fallen ofi\ Also 

applied to the point of origin or growth of a starch grain. 

Hip. The fruit of a Rose, consisting of a number of akenes surrounded by a 

ripened concave receptacle. 

Hirsute. Covered with numerous long coarse hairs. 

His'pid. Beset with erect stiff hairs, as Borage. 

Histol'ogy. The study of tissues with the aid of the microscope. 

Homol'ogous. Having the same structural nature. 

Homos'porous. Producing asexual spores of only one kind. 

Hy'brid. A cross between two varieties or species, rarely between two genera 

of the same family. 

Hydroph'ilous. Applied to flowers that are pollinated through the agency of 

water currents.

422 


Hy'drophyte. A water-plant. 

Hydrot'ropism. The response of a plant organ to the stimulus of moisture. 

Hygroscop'ic. The property possessed by certain cells or substances of absorb- 

ing moisture with avidity. 

Hyme'nium. A spore bearing membrane of a fungus. 

Hy'pha. A filament of the mycelium of a fungus. 

Hypo. A prefix of Greek orig ; n meaning under. 

Hy'pocotyl. That part of an embryo p ] antlet below the cotyledon or cotyledons. 

Hypocrater'iform. Applied to a calyx or corolla when the tube is long and 

slender and abruptly expands into a flat limb. 

Hypoder'mis. That portion of a plant organ directly beneath the epidermis. 

Hypoge'ous. Beneath the surface of the soil. 

Hypothe'cium. That portion of a thallus of a lichen directly beneath or around 

the apothecium. 

Hypog'ynous. Applied to the insertion of various floral parts on the receptacle 

and beneath the pistil. 

Id'ioblast. A cell which differs materially in form, size, character of cell wall, 

or contents from its neighbors in a tissue. 

Imbibi'tion. The taking in of water by organic bodies in such a manner as to 

cause them to swell up. 

Im'bricate. Overlapping like shingles. 

Immersed'. Growing entirely under water. 

Imparipin'nate. Applied to a pinnately compound leaf terminating with a single 

leaflet. 

Indefinite. Applied to stamens and other organs of the flower, when too 

numerous to be conveniently counted. 

Indehis'cent Not splitting open in a definite manner when ripe. 

Indig'ftious. Native. 

Indu'sium. An outgrowth of the lower epidermis of many ferns that covers 

the cluster of sporangia. 

Inequilat'eral. Having unequal sides. 

Inflores'cence. The arrangement of the flowers on a plant. 

Infundib'uliform. Funnel shaped. 

Innate'. Applied to anthers that are attached by their base to the summit of 

the filament. 

Integ'ument. A covering. 

Intercel'lular. Between the cells. 

Interfacic'ular. Applied to a cambium layer which extends from one fibrovascular 

bundle to another in the stems of Dicotyledons and Gymnosperms. 

In'ternode. That portion of the stem between two nodes. 

Interrup'tedly-Pin'nate. Applied to a pinnate leaf that has either smaller or 

" 

larger leaflets between those of usual size.

In'tine. The inner coat of the pollen grain. 

In'tra. A prefix meaning within. 

GLOSSARY 423 

Intrapet'iolar. Applied to stipules that are between the petiole and the stem; 

also to buds that are beneath or inside of the base of the petiole. 

Introrse'. Applied to anthers that face toward the gyncecium. 

Intussuscep'tion. The formation of additional particles of protoplasm between 

those already present. 

In'ulin. A carbohydrate substance isomeric with starch found in the Compos- 

itae and soma other families. 

In'volucre. A whorl (or whorls) of bracts subtending a flower or flower cluster. 

Invol'ucel. A secondary involucre. 

In'volute. Applied to the arrangement of leaves within a bud when they are 

rolled inward from both sides. 

JrritabiTity. That property of living matter whereby it responds to a stimulus. 

Isog'amy. The union of sexual cells of similar form. 

Isom'erous. Having the same number of parts in each whorl. 

Isostem'onous. Having the stamens and petals each in one whorl and of the 

same number. 

Isth'mus. Applied to the constricted portion between the two half cells in 

certain desmids. 

Karyokine'sis. Indirect nuclear division. 

Katab'olism. Destructive metabolism. 

Keel. Applied to a longitudinal ridge or elevation of cortical tissue of Senega 

root which extends from the crown downward. Also applied to the two 

inferior petals of a papilionaceous corolla which are more or less united into 

a body resembling the keel of a boat. 

Knee. A form of knot which projects upward into the air from the roots of cer- 

tain trees that grow in wet soil notably the bald cypress. 

Label'lum. The large lip-like lower petal in the flower of an orchid. 

La'biate. Two lipped. 

La'bium. The lower lip of a labiate flower. 

Lacin'iate. Applied to the margins of leaves which are deeply cut into irregular 

narrow lobes. 

Lamel'la. A little plate. Applied to the layers of carbohydrate material in a 

starch grain which surround the growing point; also to the gills of a toad- 

stool. 

Lam'ina. The blade or expanded part of any leaf. 

La'nate. Covered with long curled wool-like hairs. 

Lan'ceolate. Lance shaped. 

La'tex. The milk juice of a plant. 

Laticif'erous. Applied to the latex carrying tissue of a plant. 

Latifo'liate. Possessing broad leaves.

424 


Leaf. An expansion of the stem or branch in whose axil one or more branches 

arise. 

Leaflet. A division of a compound leaf. 

Leaf-Trace. -A fibrovascular bundle while on its way from the stem bundle to 

the leaf. 

Leg'ume. A dry, simple capsular fruit formed of a single carpel and dehiscent 

by both ventral and dorsal sutures. 

Len'ticels. Fissures in the cork of Dicotyledons formed by the swelling up and 

rupture of secondary cortex cells beneath. 

Lentic'ular. Having the shape of a double convex lens. 

Leu'coplast. A colorless plastid found in the cells of plants not exposed to light. 

Li'ane. A woody climber or twiner of tropical forests. 

Li'ber. The inner bark or phloem region of Gymnosperms and Dicotyledons. 

Li'briform-Cells. Those cells of the xylem that are thick walled and resemble 

bast-fibers. 

Lig'neous. Woody. 

Lig'nified. Covered with deposits of lignin. 

Lig'nin. A substance that adheres to the cellulose walls of certain cells and 

which is characterized by taking on a reddish coloration with phloroglucin 

and hydrochloric acid. 

Lig'ulate. Strap shaped. 

Lig'ule. A membranous appendage at the summit of the leaf-sheath in many 

grasses and cereals; a strap shaped corolla of a Composite. 

Liguliflo'rcus. Applied to Composite flower heads, as those of Dandelion and 

Chicory, which contain ligulate florets only. 

Limb. The spreading portion of a gamosepalous calyx or a gamopetalous 

corolla. 

Line. One-twelfth of an inch. 

Lin'ear. Many times longer than broad and with nearly parallel margins. 

Lobe. A division of a leaf or other flattened organ which is larger than a tooth 

but which is not a leaflet. 

Loc'ular. Having a cavity or cavities. 

Loculici'dal. Applied to the deshiscence of a capsule when it splits open along 

the dorsal suture. 

Loc'ulus. A cell or cavity of an anther, ovary, or fruit. 

Lo'ment. A modified jointed or multilocular legume that breaks open trans- 

versely into segments when mature. 

Lu'cid. Clear. 

Lu'niform. Half-moon or crescent shaped. 

Lu'rid. Dingy-brown. 

Lutes'cent.-r-Yellowish. 

Ly'rate. Applied to a pinnatifid leaf, as that of the Turnip, in which the terminal 

lobe is the largest and the rest decreasing in size toward the base. 

Lysig'enous. Applied to the formation of a type of intercellular-air-space

GLOSSARY 425 

which originates through the breaking down of cell walls common to a group 

of cells. 

Macro. A prefix of Greek origin meaning large. 

Macrosporan'gium. A spore case containing one or more macrospores. (The 

nucellus in Spermatophytes.) 

Mac'rospores. The larger of the two different kinds of spores produced by some 

of the higher Pteridophytes and the Spermatophytes. (The embryo-sac 

in Spermatophytes). 

Macrospo'rophyll. The leaf bearing the macrosporangium. (The carpel in 

in Spermatophytes.) 

Mac'ulate. Spotted. 

Ma'millate. Bearing teat-like protuberances. 

Marces'cent. Withering but not falling, dropping off. 

Marine'. Applied to plants which grow in the sea or ocean. 

Medulla. Pith. 

Med'ullary. Pertaining to the pith. 

Med'ullary Rays. Strands of parenchyma connecting the cortex with the pith 

or a portion of the xylem with a portion of the phloem. 

Megasorus. The ovule. 

Megasporan'gium. See macrosporangium. 

Megaspore. See macrospore. 

Mem'branous. Thin, soft and flexile. 

Mer'icarp. One of the two inferior akenes which are found with the carpophore 

making up the cremocarp in Umbelliferae. 

Mer'istem. Formative tissue consisting of cells which in the living plant are 

in an active state of division. 

Meristemat'ic. Consisting of generative cells or meristem. 

Mes'ocarp. The middle layer of the fruit wall or pericarp. 

Mes'ophyll. All of the leaf parenchyma within the epidermis. 

Mes'tome. The conducting portion of a fibrovascular bundle. 

Metab'olism. The sum total of all the chemical changes which take place in 

a living plant. 

Metagen'esis. Alternation of generations. The production of 

[sexual 

individuals 

by asexual mearrs and asexual or neutral individuals by sexual means. 

Metamor'phosis. A change in the form or function of an organ or organism. 

Micro. A prefix of Greek origin meaning small. 

Mi'crobe. A minute vegetable or animal organism. 

Mi'cropyle. The opening between the coats of 

anôvuTe through which the pollen 

tube enters. The orifice or foramen in the seed coat through which the 

hypocotyl passes during germination. 

Microso'mes. Applied by Strasburger to minute particles in the protoplasm 

which have a high degree of refringency. 

Microso'rus. A lobe of the anther.

426 


Microsporan'gium. A spore case containing microspores. An anther sac. 

Mi'crospore. A small spore found in a microsporangium. The pollen grain of 

a seed plant. 

Microspo'rophyll. A leaf bearing microsporangia. The stamen of seed plants. 

Mid'dle Lamel'la. A dividing line of calcium pectate between adjoining cells. 

Mid'rib. The large main central vein of a pinnately-veined leaf which is con- 

tinuous with the leaf stalk. 

Mito'sis. Indirect nuclear division. 

Monadel'phous. AppUed to stamens which are united by their filaments into 

one set as in the Mahacea. 

Monan'drous. Possessing only one stamen. 

Monan'thous. Having only a single flower on the peduncle. 

MoniTiform. Resembling a chain of beads. 

Mono. A prefix of Greek origin, meaning one or single. 

Monocar'pellary. Of one carpel. 

Monochlamyd'eous. Possessing but one perianth whorl. 

Monoc'linous. Having both andrcecium and gynceciun. 

Monocotyled'onous. Having only one cotyledon or seed leaf. 

Monce'cious. Having separate staminate and pistillate flowers on the same 

plant. 

Monoloc'ular. One chambered. 

Monom'erous. Applied to flowers having one part running through each 

whorl. 

Monopo'dium. A plant axis which elongates at the apex and sends off lateral 

branches in acropetal sequence. 

Monos'tichous. Arranged in one vertical row. 

Mu'cronate. Terminating abruptly in a small soft point. 

Multi. A prefix of Latin origin meaning many. 

Multicel'lular. Consisting of many cells. 

Multicip'ital. Many-headed; applied to a rhizome or root from which numerous 

stems arise. 

Multifa'rious. Composed of many diverse parts. 

MuTtiloc'ular. Many celled or chambered. 

Mul'tiple Fmit. A fruit composed of many small fruits, each the product of 

a separate flower, as in the Fig or Hop. 

Myce'lium. The vegetative body of a fungus consisting of intertangled hyphse. 

Mycol'ogy. That branch of Botany that treats of the Fungi. 

Mycorrhi'za. An association between the roots of certain plants and the mycelium 

of certain fungi which form an investment about their tips. 

Na'piform. Turnip-shaped. Somewhat globular, becoming abruptly slender 

and then terminating in a conical tap root. 

Naturalized. Applied to plants that have been introduced from another 

country.

GLOSSARY 

Navic'ular. Boat-shaped. 

Nec'tar. A sweet secretion by the flower. 

Nec'tary. The part of the flower which secretes nectar. 

Nerva'tion. The arrangement of veins in a leaf. 

427 

Neu'tral. Said of flowers which possess neither stamens or carpels. Also 

applied to the asexual generation of plants. 

Niv'eous. Snow-white. 

Node. The place on the stem which normally shows outgrowths of a leaf , whorl 

of leaves or leaf modifications. 

Nodose'. Having swollen joints or knobs. 

Nod'ule. A small rounded body as a root tubercle. 

Nor'mal. Usual. 

Non Not. 

Nucel'lus. The body of an ovule. 

Nuciferous. Nut-bea ring. 

Nu'cleus. A dense region of protoplasm within the cell containing chromatin 

and usually definitely circumscribed. 

Nucle'olus. A small body of dense protoplasm within the nucleus. 

Nut. A dry, indehiscent, i-celled, i-seeded fruit with a stony or leathery 

pericarp. 

Nut'let. A small nut. The characteristic fruit of the Labiates. 

Nutri'tion. That branch of Physiology which includes the absorption, distribu- 

tion and assimilation of food stuffs. 

Ob. A prefix of Latin origin signifying inversion. 

Obcon'ical. Inversely cone-shaped. 

Obcor'date. Inversely heart-shaped. 

Oblan'ceolate. Lance-shaped with the broadest part toward the summit. 

Oblate'. Flattened at the ends or poles. 

Ob'ligate. Necessary, indispensable. 

Oblique'. Taking a position between erect and horizontal as in the case of many 

stems. More developed on one side than on the other as in certain leaf 

blades. 

Ob'long. Longer than broad with nearly parallel sides. 

Obo'vate. Ovate with the attachment at the narrower end. 

Obtuse'. Having a blunt or rounded end. 

O'chrea (o'crea). A sheathing stipule. 

Ontog'eny. The history of the development of an individual. 

O'ospore. The fertilized egg. 

Oper'culum. The transversely dehiscent lid or cover of a moss capsule. 

Orbic'ular. Circular. 

Or'der. A division of a class containing one or more families. 

Orthot'ropous. Applied to ovules or seeds which are erect, with the micropyle 

at the apex and the hilum coinciding with the chalaza.

428 


O'vary. The lower part of a pistil or carpel containing the ovules. 

O'vate. Shaped like a lengthwise section of a hen's egg and having the attachment 

at the broader end. 

O'vule. A transformed bud destined to become a seed after fertilization. 

O'vum. The female sexual cell. 

Pal'ate. A convex projection on the base of the lower lip of a personate corolla. 

Pa'lea (Pal'et). An inner bract of a Grass inflorescence which with the lemma 

incloses the" flower. 

Palea'ceous. Chaffy. 

Pal'lid. Pale. 

PaTmate. Divided or lobed in radiate fashion. 

Palmat'ifid. Palmately-cleft. 

Pandu'riform. Fiddle-shaped. 

Pan'icle. A compound raceme. 

Papiliona'ceous. Having butterfly shaped flowers, as in the sub-family Papilio- 

nacece of the Leguminosae. 

Pap'illose. Bearing small nipple-shaped protuberances. 

Pap'pus. The calyx of a Composite flower. 

Papyra'ceous. Papery. 

Paraph'ysis. A sterile filament found among reproductive organs 

in certain 

plants. 

Parasit'ic. Growing upon or within and deriving sustenance from another 

living organism. 

Paren'chyma. Soft cellular tissue whose units do not have tapering extremities. 

Pari'etal. Situated on or pertaining to the wall of an ovary or pericarp. 

Part'ed. Incised nearly to the mid-rib or base. 

Parthenogenesis. The production of an embryo from an unfertilized egg. 

Pathol'ogy. The study of diseases. 

Pec'tinate. Comb-like. 

Ped'ate. Palmately parted or divided with two lateral lobes or divisions from 

each of which more or less linear divisions arise. 

Ped'icel. A branch of an inflorescence axis supporting a single flower. 

Pedun'cle. The main stalk of an inflorescence. 

Pellu'cid. Transparent, clear. 

Pel'tate. Shield shaped and attached by its lower surface to the support. 

Pen'dulous. Hanging nearly vertically downward as in the case of some ovules 

that hang from the sides of a locule. 

Pentam'erous. Applied to flowers having the number five or a multiple thereof 

running throughout each whorl. 

Pentan'drous. Having five stamens. 

Pe'po. A fruit of a Cucurbit ; a gourd. 

Peren'nial. Living more than two years. 

Per'fect. Applied to flowers that contain both stamens and carpels.

GLOSSARY 

429 

Perfo'liate. Applied to leaves which are united around the stem at their has?. 

Per'ianth. The floral envelopes, calyx and corolla or calyx alone when corolla 

is absent. 

Per'iblem. A region of meristem lying between the dermatogen and plerome 

in the growing end of a root or stem. The meristem which gives rise to 

cortex. 

Pericam'bium. A zone of meristematic tissue lying just within the endodermis. 

Per'icarp. The wall of a ripened ovary or fruit surrounding the seed or seeds. 

Pericla'dium. A sheathing petiole. 

Per'icycle. A zone of formative tissue lying outside of the fibrovascular region 

and inside of the endodermis. 

Per'iderm. The cork tissue of plant axes. 

Perid'ium. The outer covering of certain fungus fructifications as puff-b?lls. 

Per'igone. See perianth. 

Perig'ynous. Applied to stamens and petals when they are adherent to the 

calyx throat, and so borne around the gynoecium. 

Per'isperm. The nourishing tissue of some seeds outside of the embryo sac and 

representing the nucellus of the come laden with nutriment. 

ovule, which, during maturation has be- 

Per'istome. The teeth around the mouth of the capsule in mosses. 

Perithe'cium. The receptacle containing asci in certain Ascomycetes. 

Persist'ant. Applied to parts of the flower which remain until the fruit ripens 

or to leaves which remain on the plant over winter. 

Per'sonate. Applied to a bilabiate corolla which has its throat closed by a con- 

vex projection on the base of the lower lip. 

Pet'al. One of the floral leaves of the corolla. 

Pefaloid. Of some other color than green. Having the color of a petal. 

Pet'iole. A leaf stalk. 

Pet'iolule. The stalk of a leaflet. 

Phel'lodemi. Secondary cortex containing chloroplasts formed by the cork 

cambium on its inner face. 

Phel'logen. The meristem which gives rise to cork and frequently secondary 

cortex; cork cambium. 

Phlo'em. That part of a fibrovascular bundle which contains sieve tissue and 

frequently bast fibers. 

Phloroglu'cin. A white crystalline substance having the formula of C 6H 6O 3 , 

obtained by the decomposition of phloretin and from certain gummy ex- 

tracts and used with hydrochloric "acid as a test for lignin. 

Phycocy'anin. The blue pigment found in the Cyanophyceae (Blue Green Algae). 

Phycoeryth'rin. The red pigment occurring in the Rhodophyceae (Red Algae). 

Phycophae'in. The brown pigment found in the Phycophyceae (Brown Algae). 

Phycoxan'thin. A yellowish pigment occurring in some Algae. 

Phyllocla'de. A flattened branch which resembles a leaf as in Ruscus. 

Phyl'lode. A dilated petiole.

430 

Phyl'lotaxy. 

The arrangement 

Phylloxan'thin. See xanthophyll. 

Phylog'eny. The history of the race. 


of leaves on stems. 

Phys'iology. The science which treats of the functions of living organisms. 

Phy'ton. A term given by Gaudichaud to an internode with a node at its upper 

extremity which bears one or more leaves, in the axils of which buds may 

appear. 

Ptteus. The cap of a toadstool. 

Pilif'erous. Bearing hairs. 

Pilose'. Covered with long, straight and scattered hairs. 

Pin'nate. Applied to compound leaves when the leaflets are arranged along the 

mid-rib. 

Pinnat'ifid. Pinnately-cleft . 

Pinnatipar'tite. Pinnately-parted. 

Pinnat'isect. Pinnately-divided. 

Pin'nule. A secondary pinna. 

Pi'siform. Pea shaped. 

Pis'til. The central female organ of a flower consisting of one or more united 

carpels. 

Pis'tillate. Applied to flowers that possess one or more carpels but no fertile 

stamens. 

Placen'ta. The nourishing tissue which connects the ovules with the wall of 

the ovary. 

Placenta'tion. The arrangement of the placenta within the ovary or the peri- 

carp. 

Plasmo'dium. A multinucleated naked mass of protoplasm having amoeboid 

movement. The vegetative body of a Slime Mold. 

Plasmorysis. A contraction of the protoplasm of a cell due to the extraction 

of contained water under the influence of reagents of greater density than 

the protoplasmic sap. 

Plas'tid. Protoplasmic bodies of various shapes scattered about in the cyto- 

plasm. 

Ple'rome. A meristem found in the apical regions of plant axes which gives 

rise to fibrovascular tissue. 

Pli'cate. Folded like a fan. 

Plumose'. Feathery. 

Plu'mule. The rudimentary bud between the cotyledons. 

Pluriloc'ular. Having more than one chamber or cell. 

Po'lar Body. A portion of a gamete budded off before fertilization. 

Pol'len. The fertilizing dust composed of cells produced 

flowers. 

in the anthers of 

Pollina'tion. The transfer of pollen from anther to stigma and subsequent 

germination thereon.

GLOSSARY 431 

Pollin'ium. A coherent mass of pollen grains in Orchids and Milkweeds, ar- 

ranged as to be carried by insects. 

Poly. A prefix of Greek origin meaning many. 

Polyaderphous. Applied to stamens which are united many 

by their filaments into 

sets. 

Polyan'drous. Having many stamens. 

Polyan'thous. Many flowered. 

Polyarch. Said of a radial fibro vascular bundle having many xylem and phloem 

rays. 

Polycar'pellary. Composed of 3 or more carpels. 

Polycar'pic. Fruiting successively. 

PolycephaTic. Bearing many heads. 

Polycotyledon. A plant such as a Conifer which possesses more than 2 cotyledons 

or seed leaves. 

Polyem'bryony. Producing more than one embryo within a seed. * 

Polyg'amous. Applied to species in which staminate, pistillate and herm- 

aphrodite flowers are borne on the same plant. 

Polyg'onal. Having several or many angles. 

Polymor'phous. Having several to many different forms. 

Polypet'alous. Having distinct, disjoined petals. 

Pclyph'yllous. Many-leaved. 

Polysep'a!ous. Having distinct, disjoined sepals. 

Polys'tachous. Having many spikes. 

Polystem'onous. Possessing many more stamens than petals. 

Pome. A fleshy indehiscent fruit, two or more carpelled, with fibrous cartilaginous, 

or stony endocarp, the chief bulk of which consists of an adherent 

torus. 

Preflora'tion. See Aestivation. 

Prefolia'tion. See Vernation. 

Prick'le. A sharp, rigid outgrowth from the epidermis. 

Primor'dial. First formed. 

Primor'dial U'tricle. The outer plasma membrane. The outer layer of proto- 

plasm adjacent to the cell wall. 

Procam'bium. The first formed fibrovascular tissue of any organ before differ- 

entiation has taken place into xylem and phloem. 

Procumbent. Lying flat on the ground. 

Proem'bryo. The primary stage in the development of Chara consisting 

of a 

single filament and a long rhizoidal cell. The suspensor in flowering plants. 

Promyce'lium. A short hyphal growth from resting spores of smuts or rusts 

upon which basidiospores are borne. 

Prosen'chyma. Tissue composed of elongated, taper-pointed cells. 

Protan'drous. A condition of hermaphrodite flowers in which the stamens 

mature before the carpels.

432 


Protog'ynous. Applied to hermaphrodite flowers in which the carpels are mature 

before the stamens. 

Prothal'lus (Prothal'lium). A thalloid body bearing antheridia and arche- 

gonia, produced by the germination of a spore of a Pteridophyte into a 

protonema which later undergoes differentiation. 

Protone'ma. A simple or branched green filament formed by the germination 

of a spore of a moss or fern. , 

Protophlo'em. The first-formed phloem elements in a fibrovascular bundle. 

Pro'toplasm. 

Pro'toplast. 

Living matter. 

A term applied by Hanstein to the smallest body of protoplasm 

capable of individual action, either with or without a cell-wall, and either 

associated with other like units in a tissue or independent. 

Protoxy'lem. The first formed elements of xylem in a fibrova scular bundle. 

Prox'imal. Applied to the basal extremity. The attached end of an organ as 

opposed' to the free or distal end. 

Pseudo. A prefix of Greek origin indicating spurious or false. 

Pseudo-Bulb. The fleshy bulb-like internode of an epiphytic Orchid. 

Pseu'docarp. A fruit which represents the product of the ripening of a single 

ovary as well as one or more accessory parts. 

Pseudoparen'chyma. A tissue consisting of the interlacing and compact 

hyphae of a fungus. 

Puber'ulent. Covered with a fine, soft hairy coating. 

Pubes'cent. Covered with soft, short hairs. 

Pulvi'nus. An enlargement at the base of the petiole or petiolule of some leaves 

or leaflets, as in numerous Leguminosa. 

Punc'tate. Dotted with small spots or minute pits. 

Pus'tular. Applied -to surfaces having blister- or pimple-like elevations. 

Puta'men. The stony endocarp of a drupe. 

Pyre'noids. Small, rounded, colorless, refractile granules embedded in the 

chroma tophores of numerous Algae and thought to be starch forming centers. 

Pyx'is. A capsule which dehisces transversely ir>to pot and lid portions. 

Quad- or Quadri. A prefix of Latin origin signifying four. 

Quadran'gular. Four-angled. 

Quadrifo'liate. Applied to palmate leaves which have four leaflets arising from 

the summit of the petiole. 

Quinquefol'iate. Applied to any compound leaf that has five leaflets. 

Raceme'. An indeterminate inflorescence having pedicelled flowers arranged 

along a lengthened axis. 

Rac'emose. ArrangecHn racemes 

Ra'chis. The extended portion of a peduncle. 

Rad'ical. Arising from the root or base of the stem. 

Rad'icle. The rudimentary root of an embryo plantlet.

GLOSSARY 433 

Ra'mal. Pertaining to a branch. 

Ra'mus. A branch. 

Ramose'. Branching. 

Rank. A row of leaves or other organs arranged vertically on a stem. 

Ra'phe (Rha'phe). The adherent portion of the ovule stalk in inverted and half 

inverted ovules and seeds. 

Raph'ides. Bundles of needle-shaped crystals. 

Recep'tacle. The shortened stem upon which the whorls of floral leaves are 

inserted. 

Receptac'ular. Pertaining to the receptacle. 

Rec'linate. Bent downward. 

Reclin'ing. See Reclinate. 

Recurved'. Curved outward or backward to a moderate extent. 

Reflexed'. Turned outward to backward more abruptly than Recurved. 

Reg ma. A capsular fruit of 2 or more carpels that first splits into separate 

parts and then each of these dehisces. 

Rejuvenes'cence. Applied to a mode of reproductiou in which the protoplasm 

of the cell becomes rounded out, escapes by ruptnre of the cell wall, forms 

cilia and moves about, in time developing into a new Ipant. 

Ren'ifonn. Kidney-shaped. 

Repand'. Having a slightly undulating margin. 

Re'pent. Creeping. 

Re'plum. A spurious membranous septum seen in Cruciferous fruits that persists 

after the valves have fallen away. 

Retic'ulate. Applied to markings or veins which are in the form of a network. 

Retuse'. Having a broad, shallow sinus at the apex. 

Rev'olute. Said of leaves in the bud when their margins are rolled backward. 

Rha'phe. See Raphe. 

Rhi'zoids. Absorptive organs of certain plants below the Pteridophytes that are 

analagous with roots of higher plants. 

Rhizome'. A creeping underground stem. 

Rhi'zomorphs. Root-like structures composed of united hyphae and seen in 

certain fungi. 

Rib. A prominent vein or ridge. 

Rin'gent. Applied to the corolla of a bilabiate type whose throat is open and 

lips separated. 

Ripa'rious. Growing along the banks of rivers or other water-courses. 

Rosette'. A cluster of leaves or other organs. 

Ros'trate. Beaked. 

Ro'tate. Wheel-shaped. 

Rotund'. Rounded in outline. 

Ru'fous. Brownish-red. 

Rugose'. Wrink-1 ed . 

28

434 


Ru'minate. Applied to the albumen of certain seeds when the perisperm is 

. found 

coursing through the endosperm in irregular fashion. 

Run'cinate. Applied to a pinnately-cleft leaf whose lobes are directed backward 

as in the Dandelion. 

Run'ner. A stem or branch which roots at intervals as it trails along the ground. 

Sac'cate. Pouch-like. 

Sagittate. Arrow-shaped. 

Sama'ra. A winged fruit. 

Sap'rophyte. An organism that lives upon decaying or dead organic matter. 

Sar'cocarp. The fleshy portion of a drupe or other fruit. 

Sca'brous. Said of leaves, etc. that are rough or harsh to the touch. 

Scalar'iform. Applied to tracheae or tracheids whose walls show trausversely 

arranged bars, resembling the rongs of a ladder. 

Scan' dent. Climbing. 

Scape. A naked peduncle arising from a root or underground stem. 

Sca'rious. Dry and membranous. 

Schiz'ocarp. A fruit that separates when mature into 2 or more indehiscent 

mericarps. 

Schizogenous. Said of intercellular-air-spaces or of reservoirs that are formed 

by the breaking down of the middle lamellse of cells where several come to- 

gether and the later separation of the cells at these places. 

Sci'on. A shoot intended for grafting. 

Scleren'chyma. Lignified tissue. 

Sclero'tium. A hardened mass of mycelium. 

Scor'pioid. Applied to certain cymes whose flowers are situated on alternate 

sides of the floral axis. 

Scutel'lum. A shield-shaped expansion of the hypocotyl of Graminea, which 

absorbs nouirshment from the endosperm during germination and bales it 

out to the rest of the embryo. 

Sec'undine. The outer coat of the ovule. 

Seed. A fertilized and matured ovule containing an embryo. 

Se'pal. A leaf of the calyx. 

Sep'tate. Possessing one or more partitions. 

Septici'dal. A mode of dehiscence in which the opening occurs along the line 

of junction of the carpels. 

Septifra'gal. A method of dehiscence in which the valves of a capsular fruit 

break away from the partitions or septa. 

Sep'tum. A partition between cavities in an ovary or fruit or between cells in 

a tissue. 

Seric'eous. Silky. Having a covering of fine, soft, appressed, silky hairs. 

Ser'rate. Toothed with teeth projecting toward the apex. 

Ser'rulate. Finely serrate. 

Ses'sile. Without a stalk.

Se'ta. A bristle-like structure. 

Setig'erous. 

Bristle bearing. 

SiTicle. A short silique. 

GLOSSARY 435 

Sil'ique. The characteristic fruit of the Cntcifcra, consisting of a capsule of 2 

valves which separate from the replum in dehiscence. 

Sin'uate. Wavy margined. 

Soft Bast. The unlignified portion of the phloem. 

Somat'ic Cells. The body cells of an individual, in distinction from reproductive 

cells. 

Sore'dium. A scale-like structure found on many lichens and consisting of a 

group of algae cells surrounded by a network of hyphse. When detached 

from the parent-plant it has the power of mature lichen. 

developing vegetatively into a 

Soro'sis. A multiple fruit, as represented by the Mulberry and Osage Orange, 

consisting of a swollen up, condensed and mature spike. 

So'rus. An aggregation of sporangia. 

Spa'dix. A fleshy spike more or less surrounded by a bract called a spathe. 

Spathe.\ A large bract that encloses or subtends an inflorescence. 

Spat'ulate. Said of flat leaves that are narrow at the base and become gradually 

broader toward the summit, which is rounded. 

Sperma'tophyte. A seed plant. 

Spermatozo'id. A male sexual cell. See Antherozoid. 

Spermatozo'on. Another name for Spermatozoid or Antherozoid. 

Sper'moderm. The covering of the seed. 

Sphace'lia. The conidia stage of Claviceps. 

Spic'ate. Arranged in a spike. 

Spic'ule. A small pointed outgrowth. A needle-shaped crystal. 

Spike. An indeterminate inflorescence consisting of sessile florets arranged 

along a lengthened axis. 

Spike'let. A secondary spike. 

Spine. A sharp, rigid termination of a branch as in the Honey Locust. A thorn. 

Spines'cent. Spiny in structure. 

Sporad'ic. Sea ttered. 

Sporan'giophore. The stalk or support of a sporangium. 

Sporan'gium. A spore case. 

Spore. An asexual or sexual reproductive cell usually with a highly resistant 

cell wall. 

Sporogo'nium. The asexual generation in Bryophytes and Pteridophytes. 

Spo'rophyll. A spore bearing leaf. 

Spur. A tubular or saccate appendage of some part of the flower, usually con- 

taining nectar. 

Squamose'. Scale-like. 

Sta'men. A male organ of the flower producing pollen. 

Stam'inode. An abortive and sterile stamen, or any body without an anther 

occupying the normal place of a stamen.

436 


Stel'late. Star-shaped. 

Stem. The ascending axis of a plant bearing leaves or leaf modifications. 

Ste'reome. The supporting elements of a fibrovascular bundle. 

Ster'ile. i. Unproductive, as a stamen without anther, flower without pistil, 

or pericarp without seeds. 2. Devoid of living organisms. 

Steriliza'tion. The process of ridding an object of all living organisms. 

Stig'ma. That part of a pistil or carpel which receives the pollen. 

Stipe. The stem of a moss; the stalk of a fern frond; the stalk of a toadstool or 

other fungus. 

Stip'ulate. Possessing stipules. 

Stip'ule. A modified leaf, usually blade-like and situated at the base of the 

leaf-stalk. 

Sto'lon. A slender running branch above or below the surface of the soil, either 

capable of taking root or bearing a bulb at its end. 

Stolonif erous.: Bearing stolons. 

Sto'ma. A breathing pore in the epidermis of higher plants. 

Stom'ata. Plural for stoma. 

Stomat'al Cham'bei. The intercellular-air-space directly beneath the stoma. 

Stri'ate. Marked with fine longitudinal lines or grooves. 

Strigose'. Covered with sharp and rigid appressed hairs. 

Strob'ile.- A scaly multiple fruit consisting of a scale-bearing axis, each scale of 

which encloses one or more seeds. A cone. 

Style. That portion of a pistil connecting the ovary with the stigma. 

Stylopo'dium. The fleshy disk directly above the ovarian portion of an Um- 

belliferous fruit, formed by the expansion of the bases of the two styles. 

Sub. A prefix of Latin origin meaning under, below, subordinate, nearly or 

partially. 

Su'ber. Cork tissue. 

Subterta'nean. Beneath the surface of the soil. 

Su'bulate. Narrow and tapering to an acute end. 

Suc'culent. Soft and juicy or fleshy. 

Suc'ker. A shoot from the root or lower part of the stem or underground stem. 

Suffru'ticose. Applied to stems or plants that are woody at their base and 

herbaceous above. 

Sul'cate. Having longitudinal grooves. 

Supe'rior. Said of an ovary that is not adherent to and above the calyx; also 

applied to a calyx which is situated on the upgrown receptacle above the 

ovary or to a tubular calyx whose limb appears to spring from the top of the 

ovary. 

Suspen'sor. A row of cells, representing the first development of the fertilized 

egg of a seed plant, upon the end of which an embryo is formed. 

Su'ture. The line of union of two carpels. The line of dehiscence. 

Swarm Spore. A spore which possesses one or more cilia for movement.

GLOSSARY 437 

Sycon'ium. The characteristic multiple fruit of the Fig, which consists of a 

fleshy, invaginated receptacle bearing numerous akenes. 

Symbio'sis. The living together of two individuals haying a communion of life 

interests. 

Symmetrical. Said of flowers when the parts of each whorl are of the same 

number or multiples of the same number. 

Sympet'alous. See Gamopetalous. 

Sym'physis. A union of parts. 

Syncar'pous. Said of fruits and gyncecia when they are formed of two or more 

united carpels. 

Syner'gids. Two nuclei in the upper region of the embryo sac above the egg 

nucleus. 

Syngene'sious. Said of stamens when their anthers are united. 

Syn'onym. Another name for the same thing. 

Synsep'alous. See Gamosepalous. 

Tab'ular. Flattened from above downward. 

Tape'tum. A layer of cells lining the cavity of an anther sac. 

Tap-Root. The main root coursing directly downward. 

Taxon'omy. The science of classification. 

Teg'men. The inner seed coat. 

Teleu'tospore. A spore produced by the Rusts toward the close of the season 

which forms a promycelium the next year. 

Ten'dril. A modified stem, stipule, leaf, or leaflet which has taken on the 

form of a slender appendage that is capable of coiling spirally around some 

object. 

Teratol'ogy. -{The study of monstrosities. 

Terete'. More or less cylindrical. 

Ter'minal. Pertaining to the end or apex. 

Ter'nate. In threes. 

Terres'trial. Growing on land. 

Tes'sellated. Marked like a checkerboard. ' 

Tes'ta. The outer seed coat. 

Tetra. A prefix of Greek origin signifying four. 

Tetracar'pellary. Having four carpels. 

Tetradyn'amous. Having six stamens, four of which are longer than the other 

two. 

Tetrag'onal. Four-angled. 

Tetram'erous. Said of flowers that have the number four or multiple thereof 

running through their various whorls. 

Tetran'drous. Having four stamens. 

Tetrapet'alous. Having four petals. 

Tetrasep'alous. Having four sepals.


Te'trarch. Said of a radial fibrovascular bundle having 4 xylem and 4 phloem 

arms alternating with one another. 

Tet'raspores. Applied tp the asexually produced spores of the Floridea group 

of Red Algae on account of being formed in groups of four'in the mother cell. 

Tetras'tichous. Said of leaves when they are arranged in four vertical rows 

upon a stem. 

ThaTamus. Another name for receptacle. 

Thal'lus. A plant body showing no differentiation into root, stem, or leaf. 

Thermotropism. Response of living matter to the stimulus of heat or cold. 

Thorn. See Spine. 

Throat. The opening into the tube of a gamosepalous cal>x or gamopetalous 

corolla. 

Thyr'sus. A compact panicle of flowers like the Lilac or Sumac. 

Tis'sue. An aggregation of cells of similar source, structure and function in 

intimate union. 

To'mentose. Covered with dense, matted, wooly hairs. 

Tor'tuous. Bent or twisted irregularly. 

To'rus. Another name for receptacle. 

Tra'chea. An elongated cylindrical or prismatic tube found in the fibrovascular 

system 'and serving for the conduction of crude sap. 

Tra'cheid. An undeveloped trachea usually with bordered pits. 

Transpira'tion. The giving off of watery vapor by the plant. 

Tri. Three. 

Triadel'phous. Having the filaments in 3 sets. 

Trian'drous. Possessing three stamens. 

Tri'arch. Applied to a radial fibrovascular bundle having three xylem and three 

phloem arms alternating with one another. % 

Tricar'pellary. Possessing three carpels. 

Trich'oblast. An internal hair, like those projecting into the intercellular-air- 

spaces of the stems of certain Water Lilies. 

Trich'cgyne. A slender appendage to the carpogonium. 

Trich'ome. A plant hair. 

Trichot'omous. Three-branched or forked. 

Trifo'liate. Said of a compound leaf having three leaflets. 

Trimor'phous. Possessing three kinds of hermaphrodite flowers in the same 

species, differing in the relative length of their stamens and carpels. 

Tristichous. Three ranked. 

Triter'nate. Applied to a compound leaf whose petiole divides into three secondary 

petioles, each of which again divides into three tertiary petioles, each 

division bearing 3 leaflets. 

as if cut off or flattened at the summit. 

Trun'cate. Ending abruptly 

Tu'ber. A short excessively thickened end of an underground stem. 

Tu'bercle. A small wart-like outgrowth upon the rootlets, roots or subter- 

ranean stems of various plants. 

'

Tu'berous. Bearing or resembling tubers. 

GLOSSARY 439 

Tu'nicated. Covered with successively overlapping coats as the bulb of an 

Onion. 

Tur'binate. Top-shaped. 

Turges'cent. Swelling. 

Tylo'sis. A protrusion of the wall of a cell through the pit in the wall of an 

adjacent vessel and appearing in the cavity of the latter. 

Type. An individual possessing the essential characteristics of the group to 

which it belongs. 

Um'bel. The typical inflorescence of the family Umbelliferae. A more or 

less flat topped indeterminate inflorescence in which the pedicels spread 

like the stays of an umbrella. 

Un'ciform. Hook-shaped. 

Un'dershrub. A low shrub-like plant whose base is woody and upper portion 

herbaceous. 

Un'dulate. Having a wavy margin. 

Uni. A prefix of Latin origin meaning one. 

Unilateral. One-sided. 

Uniloc'ular. One-celled. 

Unise'riate. Arranged in a single row, as the cells of some plant hairs. 

Ur'ceolate. Urn-shaped. 

Ure'dospore. A one-celled spore produced during the life history of a Rust. 

U'tricle. An akene with a bladdery pericarp as Chenopodium fruit. 

Vacuole. A cavity within the protoplasm of a cell usually containing cell sap. 

Valv'ate. Applied to the leaves of a flower in the bud stage when their margins 

meet but do not overlap. 

Valve. One of the halves of a diatom. One of the parts of a pericarp that 

splits open when ripe. 

Variety. A sub-species. 

Vas'culum. A collecting case used by botanists. 

Veg'etable. A plant. 

Vein. A strand of fibrovascular tissue in a leaf. 

Vala'men. An absorptive tissue composed of several layers of dead cells 

covering the roots of some tropical epiphytic orchids and aroids. 

Vena'tion. The arrangement of veins in a leaf. 

Ven'ter. The enlarged basal portion of an archegonium. 

Ven'tral Canal' Cell. A cell beneath the entrance of the neck portion of an 

archegonium. 

Vermic'ular. Worm-shaped . 

Verna'tion. The manner in which leaves are disposed in the bud. 

. Ver'rucose. Wart-like. 

Verticillas'ter. A pair of dense cymes in the axils of opposite leaves.

440 

Vertic'illate. Whorled. 

Vessel. See trachea. 


Villose'. Covered with soft, thin, rather straight hairs. 

Virides'cent. Greenish. 

Vis'cid. Sticky. 

Vitta. An oil tube in the fruit of an Umbelliferous plant. 

Vol'va. The swollen base of the stipe in some toadstools. 

Xy'lem. That portion of a fibrovascular bundle which contains wood cells and 

fibers. 

Zob'gloe'a. 

A gelatinous mass of bacteria. 

Zo'ospore. A ciliated spore having the power of movement.] 

Zyg'ospore. A spore resulting from the union of two like gametes.! 

Zy'mogen. A microorganism capable of producing fermentation.

Abies, 294 

Abies balsamea, 295 

excelsa, 295 

Abrus precatorius, 344 

Absinthium N. F., 406 

Acacia, 91, 341, 342 

Catechu, 344 

Senegal, 91, 166, 342 

Acajou Gum, 356 

Acanthaceae, 393 

Acanthus Family, 393 

Acer, 4, 359 

spicatum, 359 

Aceraceae, 358 

Achillea millefolium, 407 

Aconiti Folia, 327 

INDEX 

Aconitine, micro-chemic test for, 84 perispermic, 214 

Aconitum, 325 

galea of, 184 

Aconitum Napellus, 325, 327 

Acorus calamus, 300, 409 

Fig. of, 301 

Acrasiales, 3 

Actaea, 325 

Actinomyces Myricarum, 135 

Adhesion, 183 

Adnation, 183 

Adonis N. F., 327 

vernalis, 327 

Êcidium, 271 

Mcium, 271 

ALgle Marmelos, 351 

Aerogens, 219 

^thallia, 230 

Agar, 246 

Agaricaceae, 274 

Agaricales, 274 

Agaricus, 274 

campestris, 274, 276 

Fig. of, 275 

Agathis loranthifolia, 295 

Agave, 410 

Agave americana, 75 

Agavose, 75 

Aggregatae, 402 

Aggregation-body, 167 

Agropyron repens, 299 

Ailanthus Family, 351 

Akene, 207 

Albugo, 251 

Albumen, defined, 214 

endospermic, 214 

mode of formation, 214, 215 

perispermic and endospermic, 215 

Albumens, 89 

Alburnum, 152 

Alcannin, 98 

Aldrovanda, 335 

Alectoria, 282 

Aletris farinosa, 303 

, 224, 312 Aleurone grains, 90 

441 

Alga-like Fungi, 247 

Algae, 231 

blue green, 227 

brown, 242 

green, 231 

red, 246 

siphon, 241 

Alkaloi(fal reagents, 83 

Alkaloids, definition of, 83 

examples of, 83, 84 

properties of, 83 

Alkanet, 390

442 

Allamanda, 384 

Allium, 303 

sativum, 303 

Allyl-iso-sulphocyanide, 96 

Aloe, 172, 303 

ferox, 303 

Perryi, 303 

vera, 303 

Alpinia officinarum, 304 

Althaea, 91, 365 

Flores, 365 

Folia, 365 

officinalis, 85, 365 

rosea, 365 

Amanita muscaria, 276, 278 

Fig. of, 276 

phalloides, 276, 278 

Fig. of, 277 

Amber, 295 

Ament, 179 

Ammoniacum, 378 

Amygdala, 338 

Am ra, 340 

Dulcis, 338 

Amygdalin, 96 

Amylodextrin, 82 

Amylum, 299 

Anacardiaceae, 356 

Anacardium occidentale, 356 

Anacyclus Pyrethrum, 5, 406 

Anamirta cocculus, 328 - 

Andreaeales, 286 

Androecium, 188 

Andromeda, 379, 380 

Anemone, 207 

Anemone Ludoviciana, 327 

pratense, 327 

Pulsatilla, 327 

Anemonella, 324 

Anethol, 377 

' 

Angelica Archangelica, 377, 378 

atropurpurea, 378 

Angelicas Fructus N. F., 377 

Radix N. F., 378 

INDEX 

Angiosperm, life history of an, 53, 58 

Angiospermous flower, diagram of 

anatomy of, 56 

Angiosperms, 296 

distinctions from gymnosperms, 

58, 59 

resemblances between and gymno- 

sperms, 58 

Angiospermae, 296 

Aniline sulphate, 98 

Anisum, 377 

Annatto, 368 

Annual ring, 147 

thickening, 146, 147 

Annulus, 275, 287, 290 

Anthemis nobilis, 406 

Anther, 188, 190 

adnate, 193 

attachment of, 193 

dehiscence, 191 

development of, 192 

extrorse, 193 

Fig. of, 192 

gross structure of, 190 

histology of, 190 

innate, 193 

introrse, 193 

lobes, 190 

syngenesious, 190 

versatile, 193 

Antheridia, 192, 194, 235 

Anthocarp, 202 

Anthoceros, 283 

Anthocerotales, 283 

Anthocyanins, 94 

Anthophore, 183 

Anthostema, 355 

Anthotaxy, 177 

Antigonum leptopus, 321 

Antipodals, 55 

Antirrhinum, 393 

Apetalae, 310 

Apii Fructus, 378 

graveolens, 378

Apocynaceae, 384 

Apocynum, cannabinum, 386 

latex of, 94 

Apothecia, 256, 281 

Apothecium, 256 

Apple, 210, 338 

Applied Botany, definition, 2 

Apposition, growth by, 97 

Appressorium, 249 

Arabis, 334 

Araceae, 300 

Arales, 300 

Aralia N. F., 376 

hispida, 376 

nudicaulis, 376 

racemosa, 376 

spinosa, 376 

Araliaceae, 376 

Arbutus unedo, 379 

Archegonia, 42 

Archegonium, defined, 282 

Archichlamydeae, 309 

Arctium Lappa, 406 

Arctostaphylos, 380 

Uva Ursi, 379, 380 

Areca catechu, 300 

nut, 300 

Arenga saccharifera, 75 

Aril, 214 

Arillode, 214 

Arisaema triphyllum, 301 

Aristolochia, 320 

reticulata, 321 

Aristolochia, serpentaria, 5, 320 

sipho, 140, 144 

Arnica, 406 

montana, 406 

Artemisia Absinthium, 406 

pauciflora, 406, 407 

Arum Family, 300 

Asafoetlda, 377 

Asagraea ofl&cinalis, 303 

Asarum N. F., 321 

canadensis, 321 

INDEX 443 

Asarum, N. F., flower of, 320 

Asci, 251, 257 

Asclepiadaceae, 386 

Asclepias, latex of, 94 

Asclepias N. F., 386 

Asclepias tuber osa, 386 

Ascocarp, 265 

Ascomycetes, 251 

Ascospores, 251 

of claviceps, 265, 266 

Ascus, 251 

Ash, 381 

mountain, 74 

Asparagine, 85 

Asparagus, 85 

Aspergillus, 256 

fumigatus, 262 

glaucus, 260 

herbariorum, 260^ 261 

niger, 263, 264 

oryzse, 261, 262 

Aspidium. See Dryopteris. 

Aspidosperma Quebracho-bianco, 5, 

386 

Assimilation, defined, 155 

parenchyma, 101 

Asteraceae, 402 

Astragalus gummifer, 91 

Atriplex, 323 

Atropa, 391 

Atropa Belladonna, 85, 391 

Aurantieae, 349 

Aurantii Dulcis Cortex, 350 

Amari Cortex, 350 

Auriculariales, 271 

Austrian Pine, Fig. of cones, 47 

Autobasidiomycetes, 271 

Auxospore, 239, 240 

Awn, 214 

Azalea,' 191, 379 

amena, 379 

apples, 273 

Azolla, 292

444 

Ba,cca, 209 

Bacilli, anthrax, 222 

disease producing, 224 

types of, 220 

Bacillus, 221 

Bacteria, 219 

chemical composition of, 223 

classification according to form, 

220, 222 

disease producing, 224 

forms of cell groups after cleavage, 

223 

morphology due to cleavage, 222 

mounting and staining of, 224, 227 

physical appearance of, 219, 222 

rapidity of multiplication, 223 

reproduction of, 222 

sporulation, 222 

Bacteriaceae, 220 

Bacterioids, 133 

Bacterium, 220 

Balansia claviceps, 266 

Balsam Apple, 401 

Balsamum Peruvianum, 343 

Tolutanum, 343 

Balsams, 92, 93 

Banana, 209 

Baneberry, 325 

Baptisia tinctoria, 344 

Barberry Family, 327 

Bark, 147 

cross-section of stem-bark of 

Cascara Sagrada, 150 

cross-section through root-bark of 

Barosma, 5, 349 

Euonymus atropurpureus, 

358 

betulina, 350 

serratifolia, 351 

Basidiomycetes, 267 

Basidiospore, 267 

Basidium, 267 

Bast, fibers, 105 

hard, 144 

INDEX 

Bast, soft, 144 

Bayberry Family, 312 

Beech, 316 

Beech Family, 316 

Beets, 323 

Begonia, leaf stalk of, 102 

roots of, 1 23 

Beggiatoa, 222 

Beggiatoaceae, 222 

Belae Fructus, 351 

Belladonna, fruit of, 209 

Belladonnas Folia, 391 

Radix, 391 

Benzoin Family, 381 

Benzoinum, 381 

Berberidaceae, 327 

Berberis N. F., 327 

Berry, 208 

Beta, 323 

vulgaris, 324 

var. Rapa, 299 

Betula lenta, 316, 

Betulaceae, 316 

Bicuculla canadensis, 334 

Bifacial leaf blade, 172 

Binomial plan of nomenclature, 4 

Birch Family, 316 

Birthwort Family, 320 

Bistorta, 322 

Bittersweet, 392 

Bixa Orellana, 368 

Bixaceae, 368 

Blackberry, 210, 338 

Black Mold, 248, 249, 250 

Bloodroot, 332 

Blue Flag, 409 

Blue Green Algae, 227 

Bluebell Family, 401 

Bog Mosses, 286 

Boletus, 274 

edulis, 75 

felleus, Fig. of, 273 

Borage Family, 388 

Borraginaceae, 388

Borragineae, 388, 390 

Boswellia carterii, 352 

Botanical nomenclature, 4, 7 

Botany, denned, i 

departments of inquiry, i, 2 

Bract, 178 

Bracteolar leaf, 178 

Branching, deliquescent, 139 

excurrent, 139 

Brassica napus 335 

nigra, 96, 335 

Brauneria pallida, 406 

Brayera, 338 

Bread mold, 248, 249, 250 

Brewer's yeast, 254 

Bridal wreath, 338 

Bromelin, 96 

Brooksilk, 236 

Brown Algae, 242 

Brunfelsia Hopeana, 391 

Brunnichia, 321 

Bryales, 286 

Bryonia N. F., 401 

Bryonia alba, 401 

dioica, 401 

Bryony, 400 

Bryophyllum, roots of, 123 

Bryophyta, 282 

outline of, 3 

Buchnera, 392 

Buchu, 350 

Buckthorn Family, 359 

Budding, 70 

Bud, defined, 135 

Buds, 135 

accessory, 136 

adventitious, 136 

alternate, 136 

axillary, 136 

classifications of, 135, 136 

flower, 135 

leaf, 135 

mixed, 136 

naked. 135 

INDEX 445 

Buds, scaly, 135 

whorled, 136 

Bulb, defined, 139 

tunicated, 139 

scaly, 139 

Bur-reed, 409 

Burseraceae, 351 

Butomus, anthers of, 190 

Buttercup Family, 324 

Caesalpineae, 341 

Cacao Praeparatum N. F., 362 

Cactaceae, 370 

Cactus Family, 370 

grandiflorus, 371 

Caffeina, 366 

Caffeine, 84 

micro-chemic test for, 84 

Calamus, 301 

Calamus Draco, 300 

Calceolaria, 189, 393 

Calcium oxalate, types of crystals, 86 

87 

how formed, 85 

tests for, 85 

Calcium pectate, 61 

Calendula officinalis, 406 

California Poppy, 332 

Callicarpa, 395 

Callistemnon, 172 

Callitris quadrivalvis, 295 

Callus, 98 

Caltrop Family, 348 

Calumba, 328 

Calyptra, of moss, 287 

Calyptrogen, 99, 122 

Calyx, bilabiate, 184 

caducous, 184 

campanulate, 184 

chorisepalous, 183 

deciduous, 184 

defined, 183 

epigynous, 184 

gamosepalous, 183, 184

446 

Calyx, half-superior, 184 

hypocrateriform, 184 

hypogynous, 184 

inferior, 184 

irregular, 184 

petaloid, 183 

perigynous, 184 

persistent, 184 

regular, 184 

rotate, 184 

superior, 184 

Cambogia, 366 

Cammeliaceae, 366 

Campanulaceae, 401 

placenta tion in, 197 

Campanulales, 400 

Camphora, 332 

Candytuft, 334 

Cane sugar, micro-chemic test for, 

. 74 

Canella N. F., 368 

Winterana, 368 

Canellaceae, 368 

Canna, starch of, 81 

style of, 198 

Cannabis, 319 

sativa, 88, 319 

var. indica, 319 

Capitulum, 179 

Caprifoliaceae, 399 

Capsicum, 208 

annuum, 391 

frutescens, 391 

Capsule, 206 

histology of a, 211 

Careya, 374 

Carcerulus, 206 

Cardamomii Semen, 304 

Carica Papaya, 369 

Caricacete, 369 

Carina, 186 

Carnauba wax, 300 

Carnoy fluid, 24 

Carotin, 93 

INDEX 

Carpel, denned, 195 

Carpophore, 183, 206 

Carragheen, 246 

Carrion fungi, 279 

Carthamus tinctorius 407 

Carum, 377 

Carvi, 377 

Caruncle, 214 

Carya, 316 

Caryophyllus, 375 

Caryopsis, 208 

Cascara Sagrada, 359 

Cascarilla, 355 

Cassia acutifolia, 343 

angustifolia, 343 

Fistula, 343 

Cassia buds, 332 

Castanea, 88, 316, 319 

dentata, 319 

Castilloa, 94 

Cat-tails, 409 

Cataria N. F., 397 

Catechu, 344 

Cathartocarpus fistula, 6, 207, 343 

Catkin, 179 

Cattleya, 307 

Caulicle, 121 

Caulophyllum, 327 

thalictroides, 327 

Caustics, 63 

Cedrella, 353 

Celandine, latex of, 94 

Celastraceae. 356, 357 

Celastrus, 356 

Cell, typical plant, 60, 61 

as a fundamental unit, 60 

membranes, 61 

Cell formation and reproduction, 70 

resting stage of, 71, 72 

sap, 6 1 

Cell walls, 96 

behavior of, to micro-chemic re 

agents, 98 

growth in area and thickness, 97

Cell walls, various kinds of and be- 

havior of each to micro-chemic 

reagents, 98 

Celloidin sections, 29 

Cells, epidermal-daughter, 175 

neighboring, 174 

of epidermis of onion-bulb scale, 

61 

subsidiary, 174 

stoma-mother, 175 

Cellulose, 96, 98 

ferment, 95 

fungous, 252 

mucilaginous modification of, 98 

reserve, 98 

Celsia, 189 

Centaurium, 384 

Centrospermae, 322 

Cephaelis acuminata, 398 

Ipecacuanha, 398 

Cephalanthus, 398 

Cereus, 370 

giganteus, 370 

grandiflorus, 371 

Cerevisiae Fermentum Compressum, 

255 

Cetraria islandica, 281, 282 

Fig. of, 281 

Chalaza, 197 

Chamselirium luteum, 303 

Chara, 241, 242 

Characeae, 241 

Charales, 241 

Chaulmoogra oil, 368 

Chelidonium, 206, 332 

majus, 333 

Chemotropism, 63 

Chenopodiaceae, 322 

Chenopodiales, 322 

Chenopodium, 324 - 

anthelminticum, 324 

Fig. of fruit, 208 

Cherry, 209, 338 

Cherry gum, 91 

INDEX 447 

Cherry laurel, emulsion in leaves of, 

96 

Chestnut, 316 

Chicory, 404, 405 

Chimaphila, 379 

umbellata, 380 

Chinese Galls, 356 

Chionanthus, 381 

virginiana, 381 

Chionanthus N. F., 381 

Chirata, 384 

Chlamydobacteriaceae, 221 

Chlamydomonas, 232 

Chlamydospores, 267 

Chlorenchyma, 101 

Chlorophycese, 231 

Chlorophyll, 93 

Chlorophyllin, 93 

Chloroplastids, 70 

Chlor-zinc-iodine, 98 

Choke cherry, 6, 96 

Chondodendron tomentosum, 328 

Chondrus crispus, 246 

Choripetalae, 310 

Chromatin, 71 

Chromatophores, 69 

Chrome-acetic fluids, 24 

Chromic acid, 23 

Chromophyll, 93 


Chromoplastids, 70 

Chromogens, 219 

Chromosomes, 73 

Chrysanthemum, corolla of, 187 

Chrysanthemum cinerariifolium, 406 

Marschalii, 406 

roseum, 406 

Chrysarobinum, 344 

Chytridiales, 251 

Cichorium, 407 . 

Intybus, 407 

Cimicifuga, 325 

racemosa, 325 

Fig. of, 326

448 

Cinchona, 5, 398 

Calisaya, 398 

Ledgeriana, 398 

Rubra, 398 

succirubra, 398 

Cinnamomum Burmanii, 332 

Camphora, 332 

Cassia, 332 

Loureirii, 332 

Saigonicum, 332 

Zeylanicum, 332 

Cinquefoil, 338 

Cistaceae, 369 

Citrullus colocynthis, 401 

vulgaris, 401 

Citrus, 349 

Aurantium, 350 

amara, 350 

Bergamia, 351 

sinensis, 350 

medica Limomum, 350 

acida, 351 

Cladonia, 281 

Clarodendron, 395 

Class, 2 

Classification, principles of, 2, 3 

Clavaria flava, Fig. of, 272 

Clavariales, 273 

Claviceps purpurea, life history of, 

265, 266, 267 

Clematis, 207, 324 

Clitandra, 94 

Close fertilization, prevention of, 199 

Clove, 375 

Clove Bark, 332 

Club Mosses, 288 

Coalescence, 183 

Coca, 348 

leaf epidermis, 174, 175 

Coca Family, 347 

Cocaine, 84, 348 

Coccaceae, 220 

Coccoloba platyclada, 321 

uvifera, 321 

INDEX 

Cocculus, 328 

Cocillana, 354 

Cocoanut oil, 300 

fruit, 210 

Cocos nucifera, 75, 300 

Coenocyte, 241 

Coffea arabica, 398 

libercia, 398 

Coffea Tosta N. F., 308 

Coffee, 398 

Cohesion, 183 

Cola Family, 361 

Cola acuminata, 362 

Colchici Cormus, 303 

Semen, 303 

Colchicum, 302 

Colchicum autumnale, 84, 303 

Colchicine, micro-chemic test for, 84 

Colocynth, 400 

Colocynthis, 401 

Collenchyma, 102, 103 

Collinsonia, 5, 397 

Columbine, 325 

Columella, 249 

Columnar crystals, 86 

Coma, 214 

Comatricha, 230 

Combretaceae, 375 

Combretum, 376 

Combretum sundaicum, 376 

Comfrey, 390 

Commiphora, 352 

Commiphora myrrha, branch of, 353 

Companion cells, 112, 113 

Compositae, 402 

inulin in, 82 

Compressed yeast, 255 

Comptonia, 314 

Comptonia asplenifolia, epidermis of 

leaf, 1 06 

photomicrograph of long-section 

through staminate catkin of, 

178 

Conceptacles of Fucus, 244, 245

Conducting parenchyma, 101 

Conducting tissues, 120 

Condurango, 386 

Cone, defined, 211 

figs, of, 47, 49 

Confervales, 234 

Conidia, 259 

Conidiospores, 259 

Coniferae, 294 

Coniferin, micro-chemic test for, 83 

Coniferales, 294 

Conifer, 293 

Conium N. F., 377 

Conium maculatum, 377 

Conjugates, 236 

Conjugation, 71, 237 

Connation, 183 

Connective, 188 

Contortse, 381 

Convallaria mpjalis, 303 

Convallariae Radix, 303 

Flores, 303 

Convolvulaceae, 386 

Convolvulus Scammonia, 387, 388 

Copaiba, 344 

Copernicia cerifera, 300 

Copper acetate test for tannins, 88 

test for resins, 93 

Coptis, N. F., 327 

trifoliata, 327 

Coral fungi, 273 

Corallin-soda solution, 98 

Coriandrum, 377 

Cork, defined, no 

microchemic tests for, no 

Cork cambium, origin in roots, 126 

origin in stems, 143 

Cork formation in roots, 126 

in stems, 143 

Corm, 139 

Cora, inflorescence of, 298 

smut, 267, 268, 269 

starch, 79 

stem, figs, of sections of, 152, 153 

29 

INDEX 449 

Cornaceae, 378 

Cornus N. F., 378 

Cornus canadensis, 378 

florida, 378 

sanguinea, 378 

Corolla, defined, 185 

apopetalous, 185 

bilabiate, 186 

campanulate, 187, 188 

carophyllaceous, 188 

choripetalous, 185 

crateriform, 186 

cruciform, 186, 187 

forms of, 187 

Corolla, gamopetalous, 186 

hypocrateriform, 186 

labiate, 186, 187 

ligulate, 1 86, 187 

infundibuliform, 187, 188 

papilionaceous, 186, 187 

persoate, 186 

polypetalous, 186 

ringent, 186, 187 

rotate, 186, 187 

urceolate, 187, 188 

Correa grandiflora, 350 

Corrosive sublimate, 24 

Corydalis, 334 

Corymb, 179 

Corynebacterium, 221 

Coto, 332 

Cotyledons, 121, 156 

Cremocarp, 206, 207 

Crenothrix, 222 

Cribiform tissue, 112 

Crocus sativus, 304 

Croton Eluteria, 355 

tiglium, 355 

Cruciferae, 334 

Crude sap, 39 

Crystal fibers, 86, 87 

Crystal sand, 86, 87 

Cubeba, 311 

Cucumber, 400, 401

450 

Cucurbita Pepo, 401 

Cucurbitaceae, 400 

Cudbear, 282 

Culm, 138 

Cuoxam, 98 

Cup Fungi, 256 

Cuphea, 371, 372 

Cupules, 283 

Cupuliferre, 316 

Curcuma N. F., 304 

Curcuma longa, 304 

Zedoaria, 304 

Currant, 208, 209 

Cutin, 97 

Cyanophyceae, 227 

Cycas revoluta, 293 

Cydonia vulgaris, 341 

Cydonium, 341 

Cyme, 179 

Scorpioid, 180 

Cynoglossum officinale, 390 

Cypress, 410 

Cypripediae, 307 

Cypripedium N. F., 308 

hirsutum, 308 

parviflorum, 308 

Cystolith, 87 

hair, 88 

Cytase, 95 

Cytisus scoparius, 344 

Cytology, i, 60 

Cytoplasm, 60, 68 

Cytoplasmic caps, 73 

d-mannose, 74 

Dacromyces deliquescens, 272 

Dacromycetales, 272 

Dahlia, 85, 404 

Daisy, 404 

Daisy Family, 402 

Damiana, 369 

Dammar, 295 

Dandelion, 404, 406 

INDEX 

Daphne Gnidium, 371 

Laureola, 371 

Mezereum, 371 

fig. of, 370 

Date Palm, 299 

Datura Stramonium, 391 

Tatula, 391 

Decadon, 371 

Dehiscence, apical porous, 

in anthers, 191 

diagrams of valvular in fruits, 

203 

in fruits, 202, 204 

longitudinal anther, 191 

transverse anther, 192 

valvular, 191 

Delafield's haematoxylin, 29 

Delphinium, 325 

Ajatis, 327 

consolida, 327 

spur of, 184 

Staphisagria, 325 

Delphinium N. F., 325 

Dermatogen, 100, 122 

Desilicification of woody sections, 30 

Desmidaceae, 236 

Determinate inflorescences, 179 

solitary, 179 

Deutzia, trichomes on leaves of, 107, 

109 

Dextrin, 81, 82 

white, 8 1 

yellow, 8 1 

Dextrose, 74 

Dianthera, 395 

Diastase, 95 

Diatoma vulgare, 238 

Diatomaceae, 238 

Diatomaceous Earth, 239 

Diatomales, 238 

Diatoms, 91, 238, 239, 240, 241 

Dicentra canadensis, 334 

cucullaria, 334 

Dicotyl Plant, morphology of type, 309

Dicotyl Seed, gross structure of, 217 

stems, 140, 147 

Dicotyledoneae, characteristics of, 308, 

309 

Dictyophora duplicata, 279 

Dicypellium carophyllatum, 332 

Diervilla, 400 

Differences between Gymnosperms 

and Angiosperms, 58, 59 

roots and stems, 123 

histological, between leaves of 

Dicotyls and Monocotyls, 

176 

Digitalis, 392 

purpurea, 393 

trichomes of, 107 

var. gloxinaeflora, 394 

Digitoxin, micro-chemic test for, 83 

Dionaea muscipula, 65, 67, 335 

Dioscorea, 5, 303 

villosa, 303 

Dioscoreaceae, 303 

Diosma, 349, 350 

Dittany, 350 

Division, 2 

Dodder, 124, 13? 

Dog Bane Family, 384 

Dogwood Family, 378 

Dog's Tooth Violet, life history of, 53, 

58 

Dorema Ammoniacum, 378 

Dorsoventral leaf blades, 169 

Dracena Draco, 301 

Dragon's Blood, 300 

Drimys Winteri, 324 

Drosera, 335 

effect of mechanical stimulus in, 

67 

longifolia, 336 

intermedia, 336 

rotundifolia, 336 

Droseraceae, 335 

Drupe, 209 

fig. of, 210 

INDEX 451 

Dryopteris Filix-mas, alternation of 

generations in, 44 

comparative physiology of root, 

stem and leaf, 39, 40 

fig. of, 34 

gross structure of stem, 33 

sori and sporangia, 40 

growth of seedling into mature 

sporophyte, 44 

histology of growing apex, 36 

lamina, 38 

mature root, 36 

stem, 33 

root apex, 37 

sori and sporangia, 40 

stipe, 37 

history of gametophyte genera- 

tion, 41, 45 

sporophyte generation, 33, 4 1 

Dryopteris Filix-mas, origin 

of new 

sporophyte or diploid plant 

from fertilized egg, 44 

vascular bundles of, 35, 292 

Dryopteris marginalis (frontispiece}, 

292 

Duboisia myoporoides, 391 

Dulcamara, 391 

Duramen, 152 

Ear fungi, 271 

Earth Stars, 279 

Earth Tongues, 265 

Ebenales, 380 

Ecballium Elaterium, 401 

Echinacea, 406 

Echinocactus, 370 

Echinocereus, 371 

Echium, 389 

Ecology, i, 408 

Economic Botany, 2 

Ectocarpus siliculosus, 242, 243 

Ectoplasm, 39 

Egg Plant, 391

452 

Ehrlich's Anilin Water Gentian Vio- 

let, 226 

Elaeis guineensis, 300 

Elastica, 355 

Elaterinum, 401 

Elettaria Cardamomum, 304 

Elm Family, 319 

Embryo, 121 

Embryo-sac, 49, 55, 197 

maturation of, 55 

Emulsin, 96 

Endocarp, 202 

Endodermis, 109, no 

Endospores, 220, 221 

Endosporium, 41 

Endothecium, 191 

Entada scandens, 207 

Enzyme, 94 

Enzymes, 94, 95 

classification of, 95, 96 

Epicalyx, 185 

Epicarp, 202 

Epidermal, outgrowths, 108-109 

papillae, 108 

scales, 109 

Epidermis, 106 

Epiphegus, 5 

Epithelium, 120 

Equatorial plate, 73 

Equisetineae, 289 

Equisetum, 290 

arvense, 291 

Ergot fungus, 265 

Erica, 379 

Ericaceae, 379 

Ericales, 379 

Erigeron canadensis, 407 

Eriodictyon, 5 

Eriodictyon californicum, 388 


Eriogonum, 321 

Erythraea Centaurium, 384 

Erythronium Americanum, life history 

of, 53-58 

INDEX 

Erythroxylaceae, 347 

Erythroxylon Coca, 348 

Eschscholtzia, 332 

Etaerio, 210 

Etiolin, 93 

Etiology, 2 

Eucalyptol, 375 

Eucalyptus, 374, 375 

globulus, 373, 375 

lamina, 172 

photomicrograph 

leaf-blade, 173 

Kino, 375 

rostrata, 375 

Endorina, 232 

Eugenia aroma tica, 374, 375 

Eugenol, 375 

Euonymus, 356 

Americanus, 356 

atropurpureus, 357 

Europaeus, 356 

Euonymus N. F., 357 

of trans, sect, 

Fig. of flowering and fruiting 

branch, 357 

Fig. of bark in cross-section, 358 

Eupatorium, 406 

perfoliatum, 406 

Euphorbia, 355 

Pilulifera, 355 

Euphorbiaceae, 355 

Euphrasia, 392 

Eurotium, 260 

Exoascus, 251 

Exobasidiales, 273 

Exogonium Purga, 388 

Exosporium, 41 

Exothecium, 191 

Fabaceae, 341 

Fagaceae, 316 

Fagales, 316 

Fagot Cassia, 332 

Fagus grandifolia, 316 

Families, naming of, 7

Family, 2 

Farfara, N. F., 406 

Fats, 91 

Fatsia horrida, 376 

Ferment, 94 

Ferments, 95 

carbohydrate, 95 

fat and oil, 96 

glucoside, 96 

proteinaceous, 96 

Fern, antheridium, 43 

apical cell, 36, 37 

archegonium, 42, 43 

foot of, 44 

prothallium, 42 

sperms of, 43 

Ferns, 290 

true, 292 

water, 292, 409 

Fern Palms, 293 

Fertilization, defined, 71 

in angiosperms, 200, 201 

cross, 199 

self, 199 

Ferula, Asafoetida, 377 

foetida, 377 

galbaniflua, 378 

Sumbul, 377 

Fibro-vascular bundles, definition, 117 

bicollateral, 117, 118 

closed collateral, 117 

concentric, 118 

diagrams of, 117 

open, 117 

radial, 118 

Ficus N. F., 319 

Carica, 319 

elastica, 94, 171 

Figwort Family, 392 

Filament, 188 

gross structure of, 190 

histology of, 190 

Filicales, 292 

Filicineae, 290 

INDEX 453 

Fission, 70 

Fistulina, 274 

Fixation, 23 

Fixing agents, 23-25 

Fixed oils, 91 

tests for, 91 

Flax, 347 

bast fibers of, 105 

Flax Family, 346 

Flemming fluids, 24, 25 

Floral diagrams, 251 

Flower, complete, 181 

diandrous, 188 

dichlamydeous, 183 

double, 182 

essential organs of, 181 

hermaphrodite, 181 

hexandrous, 188 

imperfect, 182 

liliaceous, 186 

monochlamydeous, 183 

neutral, 182 

orchidaceous, 186 

perfect, 181 

pentandrous, 188 

pistillate, 182 

polyandrous, 188 

regular, 182 

stalk, 178 

stamina te, 182 

symmetrical, 182 

Foeniculum, histology of mericarp, 212 

vulgare, 377 

Fcenum graecum, 344 

Foliage leaves, 156 

Follicle, 206. (See pod.) 

Foot, 291 

Fomes, 274 

Forestry, 2 

Forsythia, 381 

Fothergilla, 337 

Fo villa, 55 

Foxglove, 392 

Fragaria, 185

454 

Frangula, 359 

Fraxinus, 381 

Americana, 381 

Ornus, 74, 381 

Free cell formation, 71 

Frond, 33, 290 

Fructose, 74 

Fruit, classification of, 204 

Fruit, definition, 202 

structure, 202-213 

Fruits, achenial, 204, 207, 208 

aggregate, 204, 210 

baccate, 204, 208, 209 

capsular, 204, 205 

drupaceous, 204, 205 

multiple, 204, 210 

schizocarpic, 204, 206, 207 ' 

simple, 204 

Fucus vesiculosus, description 

of, 244, 246 

Figs, of, 243, 244, 245 

Fumariaceas, 334 

Fumitory Family, 334 

Fundamental considerations, 132 

tissue, 101 

Fungi, 247 

Fungi Imperfecti, 280 

Funiculus, 196, 214 

Funtumia africana, 94 

elastica, 94 

Galanga, 304 

Galbalus, denned, 211 

fig. of, 210 

Galbanum, 378 

Galea, 184 

Galega N. F., 344 

officinalis, 344 

Galium, 398 

Galla, 319 

Galls, Chinese, 356 

Japanese, 356 

Galvanotropism, 66 

Gambir, 398 

INDEX 

Gamboge family, 366 

Gamete, 71, 237 

Gametophyte, generation of Male 

Fern, 44 

Gamopetalae, 378 

Garcinia Hanburyi, 366, 367 

Gasteromycetes, 278 

Gaultherase, 96 

Gaultheria, 380 

procumbens, 96, 380 

Gaultherin, 96 

Gaylussacia, 379 

Geaster, 278 

Gelidium, 246 

Gelsemium, 382, 384 

sempervirens, 384 

stem, 145 

Gemmae, 283 

Gemmation, 70 

Genetics, defined, i 

Gentiana, 384 

acauljs, 384 

lutea, 383, 384 

verna, 384 

Gentianaceae, 384 

Gentianales, 381 

Genus, 2 

Geological Botany, denned, 2 

Geotropism, 65 

lateral, 66 

Geraniales, 344 

Geranium, 344 

maculatum, 346 

Geranium Family, 344 

Gerardia, 124, 392 

Germination, 216 

Gesneria, placentation in, 197 

Geum, 183, 185 

Gigartina mamillosa, 246 

Gills of mushrooms, 274, 275 

Ginger, absence of lignin in sclerenchyme 

fibers of, 103 

Family, 304 

Fig. of plant, 306

Ginkgoales, 4 

Ginseng, 376 

Ginseng Family, 376 

Glands, internal, 119 

Glans, 208 

Gleba, 278 

Gloeocapsa, 227, 228 

Gliadins, 89 

Globoid, 90 

Globulins, 89 

Gloiopeltis, 246 

Glomerule, 180 

Gloxinia, 197 

Gluco-alkaloids, 85 

Glucosides, characteristics of, 83 

Glutelins, 89 

Glycerin-Gelatin, 22 

Glycine hispida, 344 

Glycyrrhiza, 343 

glabra, 343 

glandulifera, 343 

Gnaphalium polycephalum, 407 

Gnetales, 4 

Gnetums, 293 

Gonophore, 183 

Gooseberry, 209 

Goosefoo.t Family, 322 

Goosypii Cortex N. F., 365 

Gossypium, 363 

arboreum, 365 

Barbadense, 365 

herbaceum, 364, 365 

hirsutum, 365 

Gossypium Purificatum, 365 

Gourd, 209 

Gourd Family, 400 

Gracilaria lichenoides, 246 

Grain, 208 

Gramma] es, 298 

Gramineae, 298 

silica in, 88 

Gram's method, 225 

Granati Fructus Cortex, 372 

Granatum, 372 

INDEX 455 

Grape, 208 

Grape Family, 359 

Grape Fruit, 209 

Grass Family, 24.7 

Greenbrier, histology of stem, 153, 

i54 

Green felt, 91, 241 

Grindelia camporum, 406 

cuneifolia, 406 

squarrosa, 406 

Gross anatomy, defined, i 

Ground Ivy, 395 

Gruinales, 344 

Guaiaci Lignum, 349 

Guaiacum, 348 

Guaiacum officinale, 349 

sanctum, 349 

Guarana, 358 

Guard cells, 173 

Guarea Rusbyi, 354 

Gum resins, 92 

Gums, 90, 91 

Gutta Percha, 380 

Guttiferae, 366 

Gyncecium, 195 

apocarpous, 196 

syncarpous, 196 

Gynophore, 183 

Gymnosperm, life history of a, 45-52 

Gymnospermse, 292 

Gymnosperms, 292 

differences from angiosperms, 58 

Haas and Hill, 95 

Habitat, defined, i 

Haematoxylon, 343 

campechianum, 343 

Hagenia abyssinica, 338 

Hairs, aggregate, 107 

barbed, 108, 109 

branched, 107 

candelabra shaped, 109 

clavate, 107, 109 

climbing, 108

456 

Hairs, glandular, 107 

hooked, 108, 109 

multicellular, 109 

non-glandular, 109 

root, 121, 122 

stellate, 107, 109 

stinging, 108 

tufted, 109 

unicellular, 107 

uniseriate, 109 

Halophytes, 410 

Haloxylon, 323 

Hamamelidaceae, 337 

Hamamelidis Folia, 338 

Hamamelis virginiana, 338 

Hancornia, 94 

Hanstein, 99 

Harebell, corolla of, 187 

Hard Bast, 144 

Harshberger, 264 

Haustoria, 123, 247 

Hawthorn, 338 

Head, denned, 179 

Heath Family, 379 

Hedeoma, 397 

Hedeoma pulegioides, 397 

Hedera Helix, 376 

Helianthemum, 369, 370 

canadense, 370 

Helianthus tuberosus, 404 

Heliotropeae, 388, 390 

Heliotropism, 64 

Helleborus niger, 327 

Helonias, 303 

Helophytes, 409 

Helvellales, 264 

Henbane, 391 

Henna, 372 

Hepaticae, 283 

Hepburn, 94 

Herb, denned, 139 

Herba Majoranae, 397 

Hertwig, 64 

Hesperidin, micro-chemic test for, 82. 

INDEX 

Hesperidium, 209 

Heterocysts, 229 

Hevea, 94 

Hevea braziliensis, 355 

Hibiscus Syriacus, 363 

Hilum, 

Hip, 207 

of ovule, 19 

of seed, 214 

of starch grain, 76 

Histology, defined, i 

of Aconitum, 132 

of a dicotyl root, 127, 131 

of annual dicotyl stem, 140 

of anther, 190 

of Aristolochia Sipho stem, 140, 

144 

of bark, 149 

of California Privet root, 127, 

131 

of Cascara Sagrada, 149 

Histology of dicotyl tuberous root, 132 

of dicotyl roots, of primary 

growth, 125, 126 

of secondary growth, 126, 127 

of dicotyl stems, 140, 146 

of fern lamina, 38, 39 

root, 36, 37 

sori and sporangia, 40, 41 

stem, 33, 36 

stipe, 37 

of filament, 190 

of fruits, 211, 213 

of Greenbrier stem, 153, 154 

of herbaceous monocotyl stem, 152 

of leaves, 167, 176 

of mericarp, 212 

of monocotyl stems, herbaceous, 

152 

woody, 153, 154 

of perennial dicotyl stem, 143, 144 

of seeds, 215, 218 

of Vanilla capsule, 211 

Holdfast, 244

Honduras Sarsaparilla, histology of 

root, 124 

Honey dew, 265 

Honeysuckle Family, 399 

fruit, 209 

Hordeum sativum, 299 

Horsetails, 289 

Host, 247 

Hounds' tongue, 390 

Humulus, 319 

glandular trichomes of, 107, 109 

Humulus lupulus, 109, 319 

Hura crepitans, 206 

Huxley, 62 

Hyacinth, 302 

Hybrid, 2 

Hydnaceae, 274 

Hydnum, 274 

Hydrangea, 336, 337 

arborescens, 337 

Hydrastis canadensis, 325 

Hydrophytes, 408 

Hydrophyllacese, 388 

Hydropterales, 292 

Hydrotropism, 64 

Hymenium, 272, 275 

Hymenomycetes, 272 

Hyoscyamus, 391 

muticus trichome, 107 

niger, 391 

Hypericaceae, 367 

Hypericum, 367, 368 

perforatum, 368 

Hypha, 247 

Hyphae, aerial, 248 

stoloniferous, 248 

submerged, 248 

Hyssopus officinalis, 397 

Ignatia, 384 

Illicium verum, 324 

Imbedding, celloidin, 28, 29 

paramne, 23, 26 

Impatiens, 5 

INDEX 457 

Indigenous, denned, i 

Indigo, 344 

Indigofera tinctoria, 344 

Indirect nuclear division, 71, 74 

Indusium, 290 

Inflorescence, defined, 177 

ascending, 177 

centrifugal, 177 

centripetal, 177 

cymose, 177 

determinate, 177 

descending, 177 

indeterminate, 177 

mixed, 177 

Inflorescences, of Pine, 294 

of Zea Mays, 298 

Imperatoria Ostruthium, 378 

Indehiscent fruit, 203 

Indeterminate inflorescence, solitary. 

178 

Indian Corn, histology of seed, 215 

Indian Turnip, 301 

Indirect nuclear division, 71 

Integuments of angiospermous ovule, 

196 

Intercellular-air-spaces, lysigenous. 119 

schizogenous, 119 

Internal glands, 119 

phloem, 146 

Internode, defined, 137 

Intussusception, 62 

Inula N. F., 406 

Inula Helenium, 406 

Inulase, 95 

Inulin, 82 

Invertase, 75, 95 

Involucre, 178 

Ipecacuanha, 398 

Ipomcea, 388 

orizabensis, 388 

pandurata, 388 

simulans, 388 

Iridaceae, 304 

Iris, 304, 309

458 

Iris, concentric bundle from rhizome 

of, 116 

florentina, 304 

germanica, 304 

pallida, 304 

versicolor, 304 

Irish Broom, corolla of, 187 

Dulse, 247 

Moss, 246 

Irritability, 62 

Irritable reactions, 62, 68 

Isoetaceae, 288 

Iscetes, 288 

Jalapa, 388 

Male, 388 

Tampico, 388 

Wild, 388 

Janczewski, 99 

Japanese Galls, 356 

Jateorhiza palmata, 328 

Jeffersonia, 5 

Jerusalem Artichoke, 351 

Jungermanniales, 283 

Juglandaceae, 315 

Juglandales, 315 

Juglans N. F., 316 

cinerea, 316 

Juniperus, 294 

communis, 295 

Oxycedrus, 295 

Sabina, 295 

Virginiana, 295 

Juniperus N. F., 295 

Jute, bast fiber of, 105 

Kalmia, 380 

Kamala, 107, 355 

Karyokinesis, 71, 74 

Kava, 311 

Kieselguhr, 239 

Kino, 344 

Kochia, 323 

Kcenigia, 321 

INDEX 

Kola, 362 

Krameria argentea, 344 

Ixina, 344 

triandra, 344 

Kiihne, 66 

Labellum, 186 

Labiatae, 395 

Labrador tea, 410 

Lachnea, 256 

Fig. of, 257 

Lactase, 95 

Lactuca, 94 

virosa, 94, 406 

Lactucarium, 406 

Lamellae, of starch grain, 76 

Lamiaceae, 395 

Lamina, defined, 169 

Lamina, mode of development of, 169 

Landolphia, 94 

Lappa, 406 

Larix Europaea, 295 

Larkspur, 324 

corolla of, 187 

Lateral rootlets, 123, 128 

Latex, 94, 112 

cells, no 


Laticiferous tissue, no 

vessels, in 

Lauraceae, 330 

Laurus, 332 

Lavendula vera, 397 

spica, 397 

Lawsonia inermis, 372 

Leaf, color, 166 

complete, 158 

definition of, 155 

exstipulate, 159 

functions, 155 

petaloid, 166 

petiolate, 159 

sessile, 159 

stipulate, 159

Leaf apex, 163 

' - 

acuminate, 163 

acute, 163 

aristate, 163 

cuspidate, 163 

emarginate, 163 

mucronate, 163 

obcordate, 163 

obtuse, 163 

retuse, 163 

truncate, 163 

arrangement, 156 

alternate, 157 

decussate, 157 

fascicled, 157 

opposite, 157 

verticillate, 157 

base, 163 

auriculate, 163 

cordate, 163 

cuneate, 163 

Leaf base, hastate, 163 

reniform, 163 

sagittate, 163 

duration, 167 

caducous, 167 

deciduous, 167 

evergreen, 167 

persistent, 167 

insertion, 159, 161 

amplexicaul, 161 

cauline, 159 

clasping, 161 

connate-perfoliate, 161 

equitant, 161 

perfoliate, 159 

radical, 159 

ramal, 159 

rank, 157 

margin, 163, 165 

crenate, 163 

cleft, 165 

dentate, 163 

divided, 165 

INDEX 459 

Leaf base, margin, entire, 163 

incised, 165 

lobed, 165 

parted, 165 

repand, 165 

runcinate, 165 

serrate, 163 

sinuate, 165 

outline, 161 

acerose, 163 

acicular, 163 

cuneate, 161 

deltoid, 163 

elliptical, 161 

ensiform, 163 

falcate, 163 

filiform, 161 

inequilateral, 161 

linear, 161 

lanceolate, 161 

oblong, 161 

oblanceolate, 161 

orbicular, 161 

ovate, 161 

peltate, 161 

spatulate, 163 

surface, 166 

glabrous, 166 

glaucous, 

hispid, 166 

1 66 

pubescent, 166 

pellucid-punctate, 166 

rugose, 1 66 

scabrous, 166 

sericious, 166 

spinose, 166 

tomentose, 166 

villose, 1 66 

verrucose, 166 

texture, 166 

coriaceous, 166 

membranous, 166 

succulent, 166 

venation, 159

460 

Leaf venation, anastomosing, 159 

furchate, 159 

palmate, 159 

parallel, 159 

pinni, 159 

reticulate, 159 

Leaves, bifacial, 172, 173 

binate, 166 

bipinnate, 123, 126 

biternate, 166 

bract, 156 

bracteolar, 155 

centric, 172 

compound, 161 

palmately, 165 

pinnately, 165 

convergent, 169, 172 

decompound, 166 

development of lamina of, 169 

dorsoventral, 169 

hydrophy tic, 169, 172 

mesophytic, 169, 171 

umbrophytic, 169, 170 

xerophytic, 169, 171 

foliage, 156 

Leaves, imparipinnate, 165 

interruptedly pinnate, 165 

lyrate, 165 

origin and development of, 

156 

palmate, 165 

paripinnate, 165 

pinnate, 165 

primordial, 156 

scale, 156 

simple, 161 

stomata of, 173 

ternate, 166 

trifoliate, 166^ 

types of Angiospermous, 155 

xerophytic, 169, 172 

Lecanora, 281 

Lechea, 369 

Legume, 206 

INDEX 

Leguminosae, 341 


root tubercles of, 132, 133 

Lemon, fruit of, 209 

Lenticels, 146 

Leptandra, 393 

Leptome, 112 

Leucoplastids, 69, 76 

Levisticum, 378 

officinale, 378 

Levulose, 74 

Lichens, 280, 282 

Licmophora, 239 

Life history, of an angiosperm, 53, 58 

of aspidium, 33, 45 

of black mold, 248, 250 

of claviceps purpurea, 265, 267 

of fern, 33, 45 

of gymnosperm, 45, 53 

of moss, 286, 287 

of mushrooms, 274, 278 

of rust, 269, 271 

of smut, 267 

Lignin, 97 

Lignocellulose, 98 

Ligule, 1 68 

Lilac, 381 

Liliaceae, 301, 303 

Liliales, 301 

Lilium, 301 

Lily Family, 301 

Lima bean, histology of, 217, 218 

Limb, 184 

Limonis Cortex, 350 

Linaceae, 346 

Linaria, 189, 392, 393 

corolla of, 187 

Linden Family, 363 

Linin, 71 

stem, 147 

Linnaeus, 4 

Linum, 347 

usitatissimum, 347 

Lipase, 96

Lippia lanceolata, 395 

Liquidambar orientalis, 338 

Fig. of, 337 

Litmus, 282 

Liverworts, 283 

Lobelia inflata, 402 

Lobeliaceae, 402 

Loganiaceae, 382 

Loment, ,207 

Lomentum, 207 

Lonicera, 399 

Lunaria, 184 

Lunularia, Fig. of, 282 

Lupulinum, 319 

Lychnis, 183 

Lycoperdales, 278 

Lycoperdon, 278 

Lycopodiaceae, 288 

Lycopodium, 5 

Lycopodium clavatum, 288 

Lythraceae, 371 

Mace, 82 

Macis, 330 

Macrocystis, 242 

Macromorphology, i 

Madder family, 398 

Magnolia family, 324 

Magnoliacese, 324 

Mahogany Family, 352 

Making of sections, 15 

Male Fern, 33, 44 

Male Jalap, 388 

Mallotus philippinensis, 355 

Mallow Family, 363 

Maltase, 95 

Maltose, 75 

Maltum, 299 

Malva Folia N. F., 365 

Malva rotundifolia, 363, 365 

sylvestris, 365 

Malvaceae, 363 

anthers of, 191 

Malvales, 361 

INDEX 461 

Mamillaria, 371 

Mariaca, 391 

Mangels, 323 

Mangroves, 410 

Manihot, 94, 355 

utilissima, 81, 96, 355 

Manna, 381 

Maple, 4, 207 

Maple Family, 358 

Maranta arundinacea, 79 

starch, 79 

Marchantia, 283 

Marchantiales, 283 

Marrubium, 5 

Marrubium vulgare, 109, 397 

Marsdenia Condurango, 386 

Marsh, 409 

Mastiche, 304 

Matico, 311 

Matricaria, 406 

> 

Chamomilla, 405, 406 

Matufation of pollen grain in Angiosperms, 

generally, 200 

in Erythronium, 55 

of embryo sac and formation of 

female gametophyte, 200 

Maw seed, 333 

May Apple, calyx of, 184 

Mayer's albumin fixative, 27 

Mechanical tissues, 120 

Medullary rays, 116 

Megaceros, 283 

Megasorus, 213 

Megaspore, 49, 55, 197 

Megasporophyll, defined, 195 

Melaleuca, 5 

Melaeluca Leucadendron, 375 

Melia, 353 

Meliaceae, 352 

Melibiose, 75 

Melilotus, 344 


Melissa officinalis, 397 

Melochia, 361

462 

Melon tree, 369 

Membrane crystals, 87 

Menispermaceae, 327 

Menispermum, 328 

canadense, 328 

Mentha piperita, 397 

Fig. of glandular trichomes, 107 

spicata, 397 

viridis, 397 

Menyanthes trifoliata, 384 

Mericarp, 206 

histology of a, 212 

Meristem, 100 

Merulius, 274 

Mesocarp, 202 

Mesophytes, 411 

Metabolism, 62 

Metachlamydeae, 378 

Method for mounting and staining 

bacteria, 224, 225 

for preparation of Canada balsam 

mount, 21 

for the preparation of a glycerin- 

gelatin mount, 22, 23 

for fixing, dehydrating, hardening 

and imbedding in paraffine, 

23, 24, 25, 26 

for sectioning and mounting ma- 

terial imbedded in paraffine, 

26, 27 

for imbedding in celloidin, 28, 29 

for sectioning celloidin material, 

29 

for staining and mounting cel- 

loidin sections, 29, 30 

of examining reserve starches, 77 

for staining and mounting par- 

affine material, amxed to 

slide, 27, 28 

Gram's, 225 

Methylene Blue, 91, 98 

Loffler's, 226 

Methylis Salicylas, 316, 380 

Mexican Grass, 75, 267 

INDEX 

Mezereum, 371 

Micrandra, 355 

Micrococci, types of, 220 

Micrococcus catarrhalis, 224 

gonorrhoeas, 224 

melitensis, 224 

meningitidis, 224 

Micro-crystals, 86, 87 

Micrometer, ocular, 31 

stage, 31 

Micrometry, 31 

Micromorphology, i 

Micron, 31 

Micropyle, 197 

Microscope, binocular, 12 

care of, 14 

compound, 9, 14 

defined, 7 

dissecting, 8 

Figs, of, 8, 10, n, 12 

lamp, 12 

rules for care of, 14 

simple, 7 

Microscopic measurement, 31, 32 

Microsorus, 188, 190 

Microsporangia, 190 

Microspore, 193 

Microsporophylls, 188 

Microspira, 221 

Microtomes, 16, 20 

hand, 16, 17 

rotary, 18, 19, 20 

sliding, 17, 1 8 

Middle lamella, 61 

Milkweed Family, 386 

Milkwort Family, 354 

Mimosa, 66, 341 

Spegazzini, 66, 67, 68 

thigmotropic reactions in. 68 

Mimoseae, 341 

Mint Family, 395 

Mistletoe, 101, 124 

Mitchella, 398 

Mitella, 336

Mitosis, 71, 74 

Mold, black, 248 

blue, 256 

bread, 248 

green, 256 

Momordica Balsamina, 401 

Monarda, 396 

punctata, 397 

Monkshood, 324 

Monoclinic prisms, 86 

Monocotyledon, morphology of typical 

plants of, 296 

Monocotyledoneae, 296 

Monotropa, 379 

uniflora, 379 

Moonseed Family, 327 

Moraceae, 319 

Morchella esculenta, 265 

Fig. of, 264 

Morels, 265 

Morning Glory Family, 386 

Morphology, denned, i 

Mosses, 284, 287 

Motherwort, 397 

Mountain Ash, 74 

Mountain Mint, 397 

Mounts, Canada balsam, 21,22 

glycerin-gelatin, 22, 23 

temporary, 20 

permanent, 20, 23 

Mucilage, 90, 91 

cell content, 90, 91 

denned, 90 

, membrane, 90, 91 


Mucor mucedo, 250 

stolonifer, 248 

Mucorales, 248 

Mucuna pruriens, 344 

Muhlenbeckia, 321 

Mulberry Family, 319 

Musci, 284 

Mushroom, 274, 275 

Mustard, corolla of, 186 

INDEX 463 

Mustard Family, 334 

stamens of, 189 

Mutisia, 404 

Mycelium, 247 

Mycobacteriaceae, 221 

Mycobacterium, 221 

Mycorrhiza, 264 

Myosotis, 389 

Myrcia acris, 375 

Myrica N. F., 315 

Caroliniensis, 133, 314 

cerifera, 133, 313, 314, 315 

Gale, 315 

Macfarlanei, 133 

Myricaceae, characters of family, 312, 

3i5 

root tubercles of, 132, 133. 134, 

i35, 312 

Myricales, 312 

Myrobalans Family, 375 

Myristica, 215, 329, 330 

fragrans, 329, 330 

Myristicaceae, 329 

Myronase, 96 

Myrosin, 96 

Myrrh Family, 351 

Myrrha, 352 

Myrtaceae, 374 

laminae of, 172 

My r tales, 371 

Myrtiflorse, 371 

Myrtle Family, 374 

Myxogastrales, 3 

Myxomycetes, 230 

Natural system, 2 

Naturalized, defined, i 

Neea, 84 

Negundo, 359 

Nepenthes, 96 

Nepenthin, 96 

Nepeta cataria, 397 

Nephrodium. See Dryopteris. 

Nest fungi, 279

464 

Nettle family, cystoliths in, 88 

Nicotiana tabacum, 391 

Nicotine, micro-chemic test for, 84 

Nidulariales, 279 

Nightshade, 391 

Family, 390 

Node, 137 

Nomenclature, 4-7 

Non-protoplasmic cell contents, 74-96 

Nostoc, 229 

Nucellus, 54, 196 

Nuclear membrane, 61, 69 

Nuclei, assisting, 200 

polar, 

200 I 

Nucleins, 89 

Nucleolus, 60, 6 1, 68 

Nucleus, 60, 68 

division, 71, 72 

endosperm, 57 

Nut, 208 

Nutgall, Fig. of, 317 

Nutlet, 206, 207 

Nutmeg Family, 329 

Nux Vomica, 215, 329, 384 

Enzyme in, 95 

Nyssa, 378 

Oaks, 316 

black, 317 

white, 317 

Ochrea, 169 

Ocimum Basilicum, 397 

Ocrea, 321 

Octant cells, 44 

Ocular micrometer, 31 

(Edogonium, 71 

Oil, benne, 393 

bitter almond, 92 

cassia, 92 

cedarwood, 295 

chaulmoogra, 368 

cocoanut, 300 

linseed, 347 

mustard, 92 

INDEX 

Oil, palm, 300 

rose geranium, 346 

spike, 397 

turpentine, 92 

Oils, fixed, 91 

nitrogenated, 92 

oxygenated, 92 

sulphurated, 92 

volatile, 92 

Olea Europaea, 381 

Oleaceae, 381 

Oleander, latex of, 94 

Oleoresins, 92 

Oleum anisi, 324 

aurantii florum, 351 

betulae, 316 

bergamottae, 351 

cadinum, 295 

cajeputi, 375 

chenopodii, 324 

cinnamomi, 332 

erigerontis, 407 

gossypii seminis, 365 

juniperi, 295 

lavendulae, 397 

lini, 347 

myristicae, 330 

olivae, 381 

pini pumilionis, 295 

rosmarini, 397 

ricini, 355 

santali, 320 

sesami, 393 

terebinthinae, 295 

theobromatis, 362 

thymi. 397 

tiglii, 355 

Olibanum, 352 

Olive Family, 381 

Oogonia, 241 

Oogonium, 245 

Oomycetes, 251 

Oospore, 234, 242 

Operculina Turpethum, 388

Operculum, 287 

Ophioglossales, 3 

Opium, 94, 333 

Optical combination, 32 

Opuntia, 370 

Opuntiales, 370 

Orcein, 282 

Orchid family, 305 

floral organs of, 307 

Orchidaceae, 305 

Orchideae, 307 

Orange, fruit of, 210 

Order, 2 

Orders, naming of, 7 

Ordinary Parenchyma, 101 

Oregon balsam, 295 

Organs, essential, 181 

plant, 121 

reproductive, 121 

vegetative, 121 

Origanum, 397 

majorana, 397 

vulgare, 397 

Oscillatoria, 228, 229 

Osmic acid, 23 

Ostwald, 94 

Ourouparia Gambir, 398 

Outline of plant groups, 3, 4 

Ovary, 195 

Ovule, amphitropous, 197 

anatropous, 197 

campylotropous, 197 

immature angiospermous, Fig. 

of, 201 

matured embryo sac in angio- 

spermous ovule, Fig. of, 

201 

orthotropous, 197 

shape of, 197 

Ovules, 195 

denned, 196 

in angiosperms, 196 

in gymnosperms, 196 

position of, 196 

30 

INDEX 465 

Ovum, 43 

Oxytropism, 64 

Pachistima, 356, 357 

Palaquium, 380 

Palm cocoanut, 299 

Date, 299 

Family, 299 

oil, 300 

Palmeae, 299 

Panax quinquefolium, 376 

repens, 376 

Pandorina, 232 

Panicle, 179 

Pansy, papillae on epidermis. 108 

Papaver, 332 

latex of, 94 


somniferum, 333 

Papaveraceae, 332 

Papaverales, 332 

Papain, 96, 369 

Papaw, 96, 369 

Papaw Family, 369 

Papilionaceae, 341, 342 

Pappus, 202 

Paprika, 391 

Papyrus, 409 

Paraphyses, 245, 256, 286 

Parasite, 247 

Pareira, 328 

Parenchyma, 101 

assimilation, 101 

conducting, 101 

ordinary, 101 

phloem, IDT 

reserve, 103 

wood, 101 

Parietales, 366 

Parmelia, 282 

Parsley Family, 377 

Passiflora, 183, 369 

Passiflora N. F., 369 

incarnata, 369

466 

Passifloraceae, 369 

Passion Flower Family, 369 

Pathogens, 219 

Paullinia, 5 

Paullinia Cupana, 358 

Paulownia imperialis, 392 

Pea Family, 341 

Peach, 209, 338 

Pear, 338 

fruit of, 2 10 

stone cells of, 104 

Peat mosses, 286 

Pedaliaceae, 393 

Pedicel, 178 

Pedicularis, 392 

Peduncle, 178 

Peireskia, 370, 371 

Pelargonium, 345 

capitatum, 346 

odoratissimum, 346 

Radula, 346 

Penicillium, 256 

brevicaule, 260, 261 

camemberti, 259, 260 

expansum, 260 

glaucum, 257, 258, 259 

roqueforti, 259 

Pentstemon, 189, 393 

Peony, 325 

Pepo, 209 

Pepper Family, 310 

Perianth, 185 

Periblem, 100, 122 

Pericambium, 37, 124, 126, 127, 128,140 

Pericarp, 202 

Pericladium, 167, 377 

Pericycle, 37, 140 

Periderm, 147 

Peridiolum, 279 

Peridium, 278 

Perigone, 183 

Perithecia, 265, 266 

Permanent mounts, preparation of , 20- 

30 

INDEX 

Peronosporales, 251 

Pertusaria, 281 

Petals, 185 

Petiole, 167, 168 

Petroselini Radix, 377 

Petroselinum, 377 

sativum, 377 

Petunia, 391 

Peziza, 256 

repanda, Fig. of, 256 

Pezizales, 256 

Pfeffer, 63 

Phaeophyceae, 242 

Phallales, 279 

Phanerogamia, 292 

Pharmaceutical Botany, denned, 2 

Phaseolus lunatus, structure of seed 

of, 217, 218 

Phaseolus multiflorus, 

cross-sections of root of. 126 

Phelloderm, 147 

Philadelphus, 336 

Phloem, defined, 119 

internal, 146 

interxylary, 146 

intraxylary, 146 


Phloroglucin, 98 

Phoenix, 295 


Photogens, 219 

Photosynthesis, 40 

denned, 155 

Phragmidiothrix, 221 

Phragmites, 409 

Phycocyanin, 94 

Phycoerythrin, 94 

Phycophaein, 94, 242 

Phycomycetes, 247 

Phycoxanthin, 242 

Phyllotaxy, 156 

Physcia stellaris, Fig. of, 280 

Physiology, defined, i 

Physostigma, 198, 344

Physostigma, Fig. of, 345 

venenosum, 344 

Phytogeography, i 

Phytoglobulins, 89, 90 

Phytolacca, 132, 324 

decandra, 324 

Fig. of, 323 

Phytolaccaceae, 324 

Phytopalaeontology, 2 

Phytopathology, i 

Phytophthora, 251 

Picea, 294 


mariana, 295 

rubra, 295 

Pickerel weed, 101 

Picrasma excelsa, 351 

Picric acid, 23 

Pigments, 93 

Pileus, 274 

Pilobolus, 248 

Pilocarpus, 349, 350 

Jaborandi, 350 

microphyllus, 350 

neighboring-cells of leaf epidermis, 

174, 175 

Pimenta, 374, 375 


Pimiento, 391 

Pimpinella, 378 

Anisum, 377 

magna, 378 

Saxifraga, 378 

Pinaceae, 294 

Pine, inflorescence of, 294 

Family, 294 

Stem, 45 

Pineapple, 96 

Pinenes, 92 

Pinites succinifer, 295 

Pinna, 290 

Pinnule, 290 

Pinus, 294 

alba, 295 

INDEX 

Pinus, austriaca, 47 

maritima, 295 

montana, 295 

palustris, 295 


strobus, 45, 295 

Piperacese, 310, 311 

Piper angustifolium, 311 

cubeba, 311 

fruit of, 209 

methysticum, 311 

nigrum, 311 

placentation of, 197 

Piscidia Erythrina, 344 

Pistacia vera, 356 

lentiscus, 356 

Pistachio, 356 

Pistil, 196 

dicarpellary, 196 

monocarpellary, 196 

polycarpellary, 196 

tricarpellary, 196 

system, 195 

Pix Burgundica, 295 

Canadensis, 295 

Liquida, 295 

Placenta, defined, 197 

Placentation^ 197 

Plankton, 409 

Planococcus, 220 

Planosarcina, 220 

Plant, acaulescent, 136 

anemophilous, 199 

annual, 124 

biennial, 124 

caulescent, 136 

cell, 60 

dioecious, 183 

entomophilous, 199 

geography, i 

hairs, 108, 109 

hydrophilous, 200 

indigenous, i 

monoecious, 182 

467

468 

Plant, naturalized, i 

organs, 121-218 

perennial, 124 

tissues, 90-120 

zoophilous, 200 

Plantaginaceae, placenta tion in, 197 

Plasma membranes, 61, 73 

Plasmodiocarp, 231 

Plasmodium, 230 , 

Plasmolysis, figure illustrating, 69 

Plastids, 68, 69 

Platystemon, 332 

Plectascales, 256 

Plerome, 100, 122 

Pleurococcaceae, 232 

Pleurococcus, 232 

Pleuston, 409 

Plum, 209, 338 

Plumule, 121, 135 

Pod, 206 

Fig. of, 205 

Podophyllum, 327 

peltatum, 327, 328 

Polemoniales, 386 

Pollen, 54, 55, 193, 195 

maturation of, 55 

Pollen grains, Fig. of, 194 

germinating, 56 

pine, 83 

Pollination, 50, 56, 199 

Pollinia, 195 

Polyembryony, 59 

Polygala, 96 

lutea, 355 

senega, 354, 355 

Polygalaceae, 354 

Polygonaceae, 197, 321 

Polygonales, 321 

Polygonatum, 302 

Polygonum, 321 

Bistorta, 322 

Polypodiaceae, 292 

Polyporaceae, 274 

Polyporus, enzyme in, 95 

INDEX 

Polyporus, officinalis, 274 

Polytrichum commune, Fig. of, 285 

life history of, 286, 287 

Pome, 209 

of apple, Fig. of, 210 

Pomegranate Family, 371 

Ponteria, 380 

Poplar, 312 

Poppy , calyx of, 184 

Family, 332 

Populi Gemmae, 312 

Populus balsamifera, 312 

nigra, 312 

Pore Fungi, 274 

Porella, 283 

Potato, 391 

Potentilla, 185 

silvestris, 341 

Prefloration, 181 

Prefoliation, 157, 158 

Prickles, 138 

Primula, style of, 199 

Primulacese, placenta tion in, 197 

Principes, 299 

Principles of classification, 2, 3 

Privet, 381 

California, 127 

Proembryo, 242 

Promycelium, 267 

Proportions of stamens, 189 

Protective tissues, 120 

Proteins, 40, 89 

tests for, 89 

Prothallial cushion, 43 

Prothallus, 41', 290 

Protoascales, 251 

Protobasidiomycetes, 267 

Protococcales, 232 

Protonema, 41, 283 

Protophloem, 127, 129, 143 

Protophyta, 219 

Protoplasm, 62 

properties of, 62, 68 

Protoplasmic cell contents, 68-70

Protoxylem, 127, 129, 143 

Prune, 209 

Prunum N. F., 340 

Primus, 4 

amygdalus var. amara, 340 

cerasus, 91 

domestica, 340, 341 

serotina, 338 

virginiana, 6 

Pseudocarp, 202 

Pseudobulbs, 137 

Pseudomonas radicicola, 132 

Fig. of, 134 

Pseudopodia, 230 

Pseudotsuga mucronata, 295 

Ptelea, 350 

trifoliata, 3^1 

Pteridophyta, 287-292 

Pterocarpus marsupium, 344 

santalinus, 343 

Puccinia Graminis, 270 

Puff Balls, 278 

Pulsatilla, 327 

Pulvinus, 167 

Pumpkin, 400 

Punica, 371 

Granatum, 372 

Punicaceae, 371 

Purging Cassia, 6 

Purified Siliceous Earth, 239 

Putamen, 202 

Pycnanthemum Montanum, 397 

Pyrenoid, 235, 236 

Pyrenomycetales, 265 

Pyrethri Flores, 406 

Pyrethrum, 406 

Pyrola, 380 

Pyrus malus, 340 

Pyxis, 206 

Quassia, 351 

amara, 351 

Quercus N. F., 319 

alba, 316, 319 

INDEX 469 

Quercus, N. F., chrysolepis, 316 

garryana, 316 

infectoria, 317, 319 

michauxii, 316 

occidentalis, 316 

platanoides, 316 

prinus, 316 

Suber, 316 

velutina, 316 

virginica, 316 

Quillaja N. F., 338 

Saponaria, 338 

Quince, fruit of, 210 

Raceme, 179 

Rachis, 178 

Radicle, 121 

Ranales, 324 

Ranunculaceae, 324 

Ranunculus, 324 

Raphe, 197. 214 

Raphides, 86, 87 

Raspberry, 210 

fruit of, 210 

Receptacle, 183 

Receptacles of Fucus, 243, 244 

Regma, 205, 206 

Rejuvenescence, 71 

Reproduction, asexual, 70 

defined, 68, 70 

sexual, 70 

vegetative, 42 

Resin, micro-chemic test for, 93 

Resina, 295 

Resins, 92 

Resinogenous layer, 92 

Reserve cellul6se, 98 


starches, 77-81 

Respiration, 155 

Rhamnales, 359 

Rhamnase, 96 

Rhamnose, 96

470 

Rhamnus cathartica, 359 

Frangula, 96, 359 

Purshiana, 359 

Rheum, 322 


palmatum, 321, 322 

tanguticum, 322 

Rhinanthus, 392 

Rhizoids, 283, 286, 290 

Rhizome, defined, 139 

Rhizopora, 410 

Rhizopus, 249 

nigricans, 248 

Rhododendron, 191 

Rhodophyceae, 246 

Rhodymenia palmata, 247 

Rhoeadales, 332 

Rhubarb, 322 

Rhus, cotinus, 356 

fruit of, 209 

Glabra N. F., 356 

japonica, 356 

semialata, 356 

toxicodendron. 356 

typhina, 356 

venenata, 356 

Ribes, 336 

Riccia, 283, 409 

Ricciocarpus, 409 

Ricinus communis, 355 

Rocella, 281 

Rock Rose Family, 369 

Root, defined, 121 

adventitious, 123 

anomalous, 123 

California Privet, 127-131 

cap, 121, 122 

classification as to form, 123 

conical, 123 

distinction from stem, 123 

duration of, 1 24 

epiphytic, 123 


fusiform, 123 

INDEX 

Root, generative tissues of, 122 

hairs, 121, 122 

histology, 124-132 

lateral, 123 

primary, 123 

secondary, 123 

system, 133 

tubercles, 132-135 

Roots, Dicotyledon, 125 

abnormal structure in. 132 

of parasitic plants, 1 23 

of primary growth, 125, 126 

of secondary growth, 126, 127 

Monocotyledon, 124 

multiple, 123 

napiform, 123 

nodule producing, 132 

Rootlet, cross section of in region of 

root hairs, 122 

Roridula, 335 

Rosa canina, 341 

centifolia, 341 

gallica, 338 

Rosa Gallica, 338 

Rosaceae, 338 

Resales, 336 

Rose, 338 

Family, 338 

Rosette aggregates, 86, 87 

Rosmarinus officinalis, 397 

Rubber, 94 

Rubi Fructus, 340 

Rubi Idaei Fructus, 340 

Rubiaceae, 398 

Rubiales, 398 

Rubus N. F., 338 

cuneifolius, 338 

Idaeus, 340 

nigrobaccus, 338 

strigosus, 340 

villosus, 338 

Rue Family, 349 

Ruellia ciliosa, 395 

Rumex N. F., 270

Rumex, 321 

acetosella, 321 

crispus, 322 

obtusifolius, 322 

Rust, wheat, 269, 270, 271 

Rusts, 268 

Ruta, 349 

graveolens, 351 

Rutaceae, 349 

hesperidin in, 82 

Ruteae, 349 

Sabal, 209, 300 

Sabbatia angularis, 384 

Sabina, 295 

Saccharomyces anomalus, 252 

apiculatus, 256 

cerevisiae, 251, 252, 253, 254 

ellipsoideus, 251, 252, 256 

exiguus, 251 

farinosus, 252 

hyalosporus, 252 

Ludwigii, 251 

mali Duclauxii, 251 

marxianus, 251 

membranifaciens, 252 

octosporus, 95 

pastorianus, 251 

Saccharomycetaceae, 251 

Saccharum, 299, 324 

officinarum, 299 

Saddle Fungi, 264 

Salep, 91 

Salicaceae, 311 

Salicales, 311 

Salicin, 82, 311, 312 


Salicornia, 410 

Salix, 312 

alba, 312 

Salvia, 396, 397 


Salvina, 292 

Samara, 207 

INDEX 47 1 

Sambucus, 399, 400 


nigra, 400 

Samphire, 410 

Sandalwood Family, 319 

Sandaraca, 295 

Sandbox, 206, 355 

Sanguinaria canadensis, 333 

Sanseviera cylindrica, leaf of, 172 

Santalaceae, 319 

Santalales, 319 

Santalum album, 320 

Rubrum, 343 

Santonica, 407 

Santoninum, 406 

Sap, cell, 258 

crude, 39 

elaborated, 116 

nuclear, 71 

vacuoles, 61 

Sapindaceae, 358 

Sapindales, 356 

Saponin, micro-chemic test for, 83 

Sapotaceae Family, 380 

Saprogens, 219 

Saprolegniales, 251 

Saprophyte, 247 

Sarcina, 220, 222 

Sarcocarp, 203 

Sarcosphaera, 257 

Sarracenia, style of, 198 

Sarraceniales, 335 

Sarsaparilla, 303 

endodermis of Mexican root, no 

of Honduras root, no 

Fig. of Honduras root in T. S., 125 

Sassafras, 331, 332 

Medulla, 332 

variifolium, 332 

Satureia hortensis, 397 

Saw Palmetto Palm, Fig. of, 300 

Saxifragaceae, 336 

Saxifrage, 336 

Family, 336

472 

Scale leaves, 156 

Scales, 109 

Scammoniae Radix, 388 

Scape, 138 

Schizomycetes, 219 

Schizophyta, 219 

Schleiden, 60 

Schulze's process of maceration, 30, 31 

Scilla, 303 

Scitaminales, 304 

Sclerenchyma, 103 

fibers, 105 

Sclerotium, 265 

Scoparius, 344 

Scopola Carniolica, 391 

Scorpioid cyme, 180 

Scotch Pine, 48 

Scouring Rushes, 289 

Scrophularia, 393 

Scrophulariaceae, 392 

Scullcap, 396 

Scutellaria lateriflora, 397 

Secale cereale, 265 

Secretion canals, 119, 120 

reservoirs, 119, 120 

sacs, 119 

Secretory cells, 113 

Sections, making of, 15 

celloidin, 29 

paraffine, 26, 27 

radial-longitudinal, 16 

tangential-longitudinal, 16 

transverse, 16 

Seed, 213 

appendages, 213, 214 

albuminous, 215 

coats, 213, 214 

defined, 213 

Dicotyl, 217 

exalbuminous, 215 

germination, 216 

histology, 215-218 

Monocotyl, 215, 216 

Seed-grass, 409 

INDEX 

Seedling, 216, 217 

of White Pine, 52 

Selaginella, 288 

Martensii, 289 

Selaginellaceae, 288 

Semen Rapae, 335 

Senecio N. F., 406 

aureus, 406 

Senega, 355 

Senna, 343 

neighboring cells of, 174 

stomatal apparatus of, 175 


Sepaline, position, 185 

spurs, 184 

stipules, 184 

Sepals, 183 



perigynous, 184 

Serenoa serrulata, 300 

Serpentaria, 321 

Serphyllum, 397 

Sesame Family, 393 

Sesamum indicum, 393 

Sieve, 112 

Silica, 88 

callus of plates, 98 

Siliceous Earths, 239 

Silicule, 206 

Siliqua, 206 

Simaruba amara, 351 


Simarubaceae, 351 

Sinapis alba, 335 

nigra, 335 

Sinigrin, 96 

Siphonales, 241 

Sitotropism, 64 

Skunk Cabbage, 301 

Slime Molds, 230 

Smilax, 302 

medica, 303 

oflficinale, 303

Smilax, ornata, 303 

Smuts, 267 

Soapwort Family, 358 

Soft bast, 144 

Solanaceae, 390 

Solaneae, 191 

Solanine, micro-chemic test for, 85 

Solanum N. F., 391 

Solanum, 391 

carolinense, 391 

Dulcamara, 391 

tuberosum, 85 

Sorbinose, 74 

Soredia, 28"i 

Sorghum, 75, 299 

Sorosis, 2ii 

Sorus, 40, 290 

Sour Gum, 378 

Soy Bean, 344 

Spadix, 179 

Sparganium, 409 

Spathe, 178 

Species, 2 

Spermacia, 271 

Spermagonia, 271 

Spermatocyte, 43 

Spermatophyta, 292 

outline of, 4 

Spermatozoid of fern, 43 

Spermoderm, 213 

Sperms, 245 

Sphaerella, 232 

Sphagnales, 286 

Sphagnum, 286 

acutifolium, 284 

squarrosum, 284 

Spicebush, 331 

Family, 330 

Spiderwort, 66 

Spigelia, 382, 383 

marilandica, 384 

Spike, 179 

Spinach, 323 

Spine, 138 

INDEX 473 

Spirilla, types of, 221 

Spirillum, 221 

cholerae asiaticae, 224 

obermeieri, 224 

Spirochaeta, 221 

Spirogyra, 236 

life history of, 236, 237, 238 

Spirosoma, 221 

Sporangiophores, 248 

Spore, 41 

brand, 267 

Sporogonium, 283, 286 

Sporophore, 274 

Sporophyte, 33, 44 

Spruce Gum, 295 

Spurge Family, 355 

Spur shoot, 47 

Spurs, sepaline, 184 

Staff Tree Family, 356, 357 

Stage micrometer, 31 

Stain, for "acid proof" bacteria, 226 

Broca's differential, 227 

Delafield's haematoxylin, 29 

Ehrlich'sx gentian violet, 226^ 

Gram's, 225 

Loffler's methylene-blue, 226 

Van Ermengem's flagella, 226, 227 

Ziehl's carbol-fuchsin, 226 

Staining, 21, 22, 28, 29 

of bacteria, 224-227 

Stamen System, 188 

Stamens, color of, 190 

connation of, 189 

definite, 188 

diadelphous, 189 

didynamous, 189 


gynandrous, 189 

histology of parts, 190-195 


indefinite, 188 

insertion of, 189 

monadelphous, 189 

perigynous, 189

474 

Stamens, polyadelphous, 190 

proportions of, 189 

tetradynamous, 189 

triadelphous, 189 

Standardization of ocular micrometer, 

3i 

Staphisagria, 325 

Staphylococcus, 220, 222 

pyogenes aureus, 224 

Star Apple Family, 380 

Starch, 75 

assimilation, 75 

Barley, 78, 79 

Bean, 80, 81 

Buckwheat, 79 

Canna, 81 

Cassava, 80, 81 

characteristics of commercial 

kinds, 77-81 

compound grains, 77 

Cora, 79, 80 

fill, 77 

Maranta, 78, 79 

Pea, 80, 8 1 

Potato, 78, 79 

Rice, 79, 80 

Rye, 78, 79 

Sago, 78, 81 

reserve, 76 

structure and composition of, 76 

Wheat, 78, 79 

Steckbeck, 68 

Stem, defined, 136 

direction of growth, 137 

distinction from root, 123 


generative tissues of, 137 

modification of, 137, 138 

Stemonitis fusca, 230 

Stems, above-ground, 138 

aerial-tuberous, 137 

annual, 137, 138 

ascending, 137 

biennial, 137 

INDEX 

Stems, cactoid, 138 

climbing, 138 

decumbent, 137 

duration of, 137 

elongation of, 137 

endogenous, 139, 140 

exceptional types of dicotyl, 145, 

146 

exogenous, 139 

fruticose, 138 

herbaceous, 138 

histology of annual dicotyl, 140- 

142 

of perennial dicotyl, 143-145 

of herbaceous monocotyl, 152 

of woody monocotyl, 153, 154 

perennial, 138 

phylloid, 138 

procumbent, 137 

reclining, 137 

repent, 137 

scandent, 138 

size of, 136 

spiny, 137 

subterranean tuberous, 138 

sunruticose, 138 

tendriliform, 137 

twining, 137 

underground, 139 

Sterculiaceae, 361 

Sterigmata, 258, 259, 266 

Sterigmatocystis niger, 264^ 

Stevens, 85 

Stigma, 196, 199 

defined, 199 

of animal pollinated flowers, 199 

of wind pollinated flowers, 199 

papillae of, 199 

Stillingia, 355 

sylvatica, 355 

Stimuli, extrinsic, 62 

intrinsic, 62 

Stink horn, 279 

Stipe, 265, 274, 290

Stipule, defined, 168 

Stipules, 1 68, 169 

axillary, 169 

caducous scaly, 169 

lateral, 168 

modified, 169 

sepaline, 184, 185 

Stolon, 138 

Stoma mother-cell, 174 

Stomata, 106 

structure and development of, 

173-176 

transpiration, 106 

water, 106 

Stone cells, 103 

Stone cells, defined, 103 

Figs, of, 102, 104 

Stoneworts, 241 

Stramonium, 391 

Strawberry, 338 

Streptococcus, 220, 222 

erysipelatis, 224 

pneumoniae, 224 

Strep tothrix, 221 

bo vis, 224 

Strobile, 210, 211 

Stromata, 214 

Strophanthin, micro-chenaic test for, 

82 

Strophanthus, 386 

hispidus, 385, 386 

Kombe, 386 

Strychnine, micro-chemic test for, 83 

Strychnos Ignatii, 384 

Nux Vomica, 382, 384 

Style, 195, 198 

arms, 198 

collecting hairs of, 198 

column, 198 

Styloids", Fig. of, 86 

Styraceae, 381 

Styrax, 338 

Benzoin, 381 

Suberin, 97 

INDEX 475 

Suberized walls, 98 

Sub-hymenium, 275 

Succinum, 295 

Succus Citri, 351 

Pomorum, 340 

Sucrose, 74 

Sudan III, 98 

Sugar cane, 75 

Sugars, 74 

Pliickiger's micro-chemic test for 

determination of various 

kinds of, 74 

Sumac Family, 356 

Sumbul, 377 

Summer Savory, 397 

Sundew, effect of stimulus applied to 

tentacles of, 67 

Surirella, 240 

Sutures, 202 

Swarm spores, 231 

Sweet Basil, 397 

Sweet Potato, 387 

Swertia Chirayita, 384 

Syconium, 210, 211 

Sympetalae, 378 

Symphytum, 389, 390 

Symplocarpus fcetidus, 301 

Synchytrium, 251 

Synedra, 240 

Synergids, 55 

Syringa, 381 

Systematic Botany, 219-407 

defined, i 

Tabacum, 391 

Tamarindus, 344 

indica, 344 

Tanacetum, 407 

vulgare, 407 

Tannins, definition, 88 

kinds of, 88 

properties of, 88 

tests for, 88 

Tapetum, 191, 193

476 

Tapioca, 355 

Taraxacum officinale, 406 

Taxodium, 410 

Taxonomy, defined, i 

Taxus, 294 

Tea Family, 366 

Teak-wood, 395 

Tectona, 395 

Tegmen, 213 

Tela contexta, 275 

Teleutospores, 270 

Teliospores, 270 

Tendril, 138 

Terebinthina, 295 

Canadensis, 295 

Laricis, 295 

Ternstrcemiaceae, 366 

Terpenes, 92 

Terra Silicea Purificata, 239 

Testa, 213 

Tetrads, 193 

Tetraspores, 246 

Thalamus, 183 

Thallophyta, 219 

outline of, 3 

Thallus, 219 

Thamnidium, 248, 250 

Thea sinensis, 366 

Theaceae, 366 

Thelepho rales, 273 

Theobroma, 361 

Theobroma Cacao, 362 

Thermotropism, 63 

Thigmotropism, 66-68 

Thiothrix, 222 

Thistle, 404, 406 

Thorn, 138 

Thornapple, 391 

Throat, 184 

Thuja, 294, 295 

occidentalis, 295 

Thymeleaceae, 371 

Thymus Serphyllum, 397 

vulgaris, 397 

INDEX 

Thyrsus, 180 

Tilia, 363 

Tiliaceae, 361, 363 

Tissue, 99 

assimilation parenchyma, 101 

chlorenchyma, 101 

collenchyma, 102, 103 

conducting parenchyma, 101 

cork, no 

cribiform, 112 

embryonic, 100 

endodermal, no 

epidermal, 106-109 

fundamental, 101 

generative, 99, 100 

ground, 101 

laticiferous, no 

medullary ray, 116 

meristematic, 100 

ordinary parenchyma, 101 

reserve parenchyma, 103 

sclerenchyma, 103-105 

sieve, 112 

stony, 103 

suberous, no 

tracheary, 112, 116 

Tissues, 99-120 

classified accd. to function, 120 

Tobacco, 391 

Toluifera Balsamum, 343 

Pereirae, 343 

Tomato, 208, 391 

Tooth Fungi, 274 

Tormentilla, 341 

Torus, 183 

Touranose, 75 

Toxins, 63 

Tracheas, 112, 113, 115 

Tracheary tissue, 112 

Tracheids, 116, 151 

Tradescantia zebrina, 69 

Tragacanth, 91 

Tragacantha, 342 

Trailing Arbutus, 380

Trama, 275 

Transpiration, 155 

Stomata, 106 

Tree, 138 

Trees, branching in, 139 

Trehalase, 95 

Trehalose, 75 

Tremellaceae, 272 

Tremellales, 271 

Trichomes, 107, 108, 109 

Trifolium N. F., 344 

pratense, 344 

repens, 341 

Trigonella Fcenum-graecum, 344 

Trillium, 303 

Tristicha, 88 

Triticum, 299 

Tropophytes, 411 

True Mosses, 286 

Truffles, 264 

Trunk, 138 

Trypsin, 96 

Tschirch, 92 

Tsuga canadensis, 295 

Tuber, 139 

Tuberales, 264 

Tubiflorae, 386 

Turnera aphrodisiaca, 369 

diffusa, 369 

Turneraceae, 368 

Turpentine, 295 

Bordeaux, 295 

Turpeth Root, 388 

Tussilago Farfara, 406 

Twin crystals, 86 

Typha, 193, 194, 409 

Ulmaceae, 319 

Ulmus, 91, 319 

fulva, 319 

Ulothricaceae, 234 

Ulothrix zonata, 234, 235 

Umbel, 179 

Umbellales, 376 

INDEX 477 

Umbelliferae, 377 

Umbelliflorae, 376 

Uredinales, 268 

Uredinium, 270 

Uredo linearis, 270 

Uredospore, 270 

Urginea maritima, 303 

Urticales, 319 

Usnea, 281, 282 

Ustilaginales, 267 

Ustilago Maydis, 267 

Zeae, 267 

Fig. of, 268 

Utricle, 208 

Utricularia, 172 

Uva, 209 

Uva Ursi, 380 

neighboring cells of leaf epider- 

Vaccineae, 380 

Vaccinium, 380 

mis, 174, 175 

Vacuole, glycogen, 252 

Vacuoles, sap, 61 

Valerian Family, 402 

Valeriana, 402 


Valerianacese, 402 

Valves of fruits, 202 

Vanilla, 308 

planifolia, 308 

histology of fruit, 211, 212 

Varieties, naming of, 6 

Variety, 2 

Vaucheria, 91, 241 

Vaucheria terrestris, 235 

Vavaea, 353 

Veil, partial, 275 

universal, 277 

Venation, leaf, 159 

Venus fly-trap, 65, 67 

Veratrina, 303 

Veratrine, micro-chemic test for, 

84

478 

Veratrum, 303 

album, 84 

viride, 303 

Verbasci Flores, 393 

Folia, 393 

Verbascum, 189, 393 

Blattaria, 392, 393 

phlomoides, 393 

Thapsiforme, 393 

Thapsus, 392, 393 

Verbena N. F., 395 

hastata, 395 

Verbenaceae, 395 

Vernation, 157, 158 

Veronica, 189, 392. 393 

virginica, 393 

Verticillaster, 180 

Viburnum, 399 

Lentago, 400 

Opulus, 400 

Prunifolium, 400 

Vinca, style of, 198 

Vinum Xericum, 361 

Viola tricolor, 368 

style of, 198 

Violaceae, 368 

Viscum, 59 

Vitaceae, 359 

Vitis, 361 

Volatile oils, 92 

Volvocales, 232 

Volvox globator, life history of, 232- 

234 

Vouacapoua Araroba, 344 

Walnut Family, 315 

Waltheria, 361 

Water Ferns, 292 

Water Leaf Family, 388 

Watermelon, 400 

seed, 401 

Water stomata, 106 

Weigelia, 399, 400 

Welwitchia, 58 

c. 

~ 

INDEX 

Weymouth Pine, 45 

Wheat, Fig. of plant, 297 

rust, 269 

Wheat, Fig. of fruit, 208 

starch, 78, 79 

White Pine, life history of, 45-52 

Wild Black Cherry, 6, 96 

Cherry, 4 

Willow, 311, 312 

Winter's Bark, 324 

Witchhazel Family, 337 

Wood, denned, 150 

fibers, 105 

heart-, 152 

microscopic characters of angio- 

spermous and gymnospermous, 

152 


pine, 151 

sap-, 152 

Wood Rush, parenchyma of, 101 

Xanthophyll, 93 

Xanthoxyli Fructus N. F., 35 r 

Xanthoxylum, 350, 351 

americanum, 351 

Clava-Herculis, 351 

Xerophytes, 410 

Xylem, 119 

stages in development of elements 

of, 114 

Yam Family, 393 

Yeasts, bottom, 255 

defined, 251 

Hansen's classification of, 251, 252 

top, 255 

Yucca, 301 

Zanthoxyleae, 349 

Zea N. F., 299 

Mays, 299 

bundle of stem of, 115 

inflorescences of, 298

Zedoaria, 255 

Ziehl's Carbol-fuchsin, 226 

Zingiber, 304 


Fig. of, 306 

Zingiberaceae, 304 

Zoogloea, 133 

Zoo spore, 71 

INDEX 

Zoospores, 231, 235 

Zygnemaceas, 236 

Zygomycetes, 248 

Zygophyllaceae, 348 

Zygospore, 231, 235, 250 

Zygote, 71 

Zymase, 74, 95 

Zymogens, 219 

479

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