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Pernambuco
Fungi
Eum ycota: mushrooms, sac fungi, yeast, molds, rusts, smuts, etc.
Introduction
The organisms of the fungal lineage include mushrooms, rusts, smuts, puffballs, truffles, morels,
molds, and yeasts, as well as many less well-known organisms (Alexopoulos et al., 1996). More
than 70,000 species of fungi have been described; however, some estimates of total numbers
suggest that 1.5 million species may exist (Hawksworth, 1991; Hawksworth et al., 1995).
As the sister group of animals and part of the eukaryotic crown group that radiated about a billion
years ago, the fungi constitute an independent group equal in rank to that of plants and animals.
They share with animals the ability to export hydrolytic enzymes that break down biopolymers,
which can be absorbed for nutrition. Rather than requiring a stomach to accomplish digestion,
fungi live in their own food supply and simply grow into new food as the local environment
becomes nutrient depleted.
Most biologists have seen dense filamentous fungal colonies growing on rich nutrient agar plates,
but in nature the filaments can be much longer and the colonies less dense. When one of the
filaments contacts a food supply, the entire colony mobilizes and reallocates resources to exploit
the new food. Should all food become depleted, sporulation is triggered. Although the fungal
filaments and spores are microscopic, the colony can be very large with individuals of some
species rivaling the mass of the largest animals or plants.
Figure 1: Hyphae of a wood-decaying fungus found growing on the underside of a fallen log. The
metabolically active hyphae have secreted droplets on their surfaces. Copyright M. Blackwell 1996.
Prior to mating in sexual reproduction, individual fungi communicate with other individuals
chemically via pheromones. In every phylum at least one pheromone has been characterized, and
they range from sesquiterpines and derivatives of the carotenoid pathway in chytridiomycetes and
zygomycetes to oligopeptides in ascomycetes and basidiomycetes.
Within their varied natural habitats fungi usually are the primary decomposer organisms present.
Many species are free-living saprobes (users of carbon fixed by other organisms) in woody
substrates, soils, leaf litter, dead animals, and animal exudates. The large cavities eaten out of
living trees by wood-decaying fungi provide nest holes for a variety of animals, and extinction of
the ivory billed woodpecker was due in large part to loss, through human activity, of nesting trees
in bottom land hardwoods. In some low nitrogen environments several independent groups of
fungi have adaptations such as nooses and sticky knobs with which to trap and degrade
nematodes and other small animals. A number of references on fungal ecology are available
(Carroll and Wicklow, 1992; Cooke and Whipps, 1993; Dix and Webster, 1995).
However, many other fungi are biotrophs, and in this role a number of successful groups form
symbiotic associations with plants (including algae), animals (especially arthropods), and
prokaryotes. Examples are lichens, mycorrhizae, and leaf and stem endophytes. Although lichens
may seem infrequent in polluted cities, they can form the dominant vegetation in nordic
environments, and there is a better than 80% chance that any plant you find is mycorrhizal. Leaf
and stem endophytes are a more recent discovery, and some of these fungi can protect the plants
they inhabit from herbivory and even influence flowering and other aspects of plant reproductive
biology. Fungi are our most important plant pathogens, and include rusts, smuts, and many
ascomycetes such as the agents of Dutch elm disease and chestnut blight. Among the other well
known associations are fungal parasites of animals. Humans, for example, may succumb to
diseases caused by Pneumocystis (a type of pneumonia that affects individuals with supressed
immune systems), Coccidioides (valley fever), Ajellomyces (blastomycosis and histoplasmosis),
and Cryptococcus (cryptococcosis) (Kwon-Chung and Bennett, 1992).
Figure 2: The fluffy white hyphae of the mycorrhizal fungus Rhizopogon rubescens has enveloped the
smaller roots of a Virginia pine seedling. Note that some of the mycelium extends out into the
surrounding environment. Copyright J. B. Anderson 1996.
Figure 3: Entomophthora, "destroyer of insects", is the agent of a fungual infection that kills flies.
After their death the fungal growth erupts through the fly cuticle, and dispersal by forcible spore
discharge is a source of inoculum for infection of new flies. Copyright G. L. Barron 1996.
Fungal spores may be actively or passively released for dispersal by several effective methods.
The air we breathe is filled with spores of species that are air dispersed. These usually are species
that produce large numbers of spores, and examples include many species pathogenic on
agricultural crops and trees. Other species are adapted for dispersal within or on the surfaces of
animals (particularly arthropods). Some fungi are rain splash or flowing water dispersed. In a few
cases the forcible release of spores is sufficient to serve as the dispersal method as well. The
function of some spores is not primarily for dispersal, but to allow the organisms to survive as
resistant cells during periods when the conditions of the environment are not conducive to growth.
Fungi are vital for their ecosystem functions, some of which we have reviewed in the previous
paragraphs. In addition a number of fungi are used in the processing and flavoring of foods
(baker's and brewer's yeasts, Penicillia in cheese-making) and in production of antibiotics and
organic acids. Other fungi produce secondary metabolites such as aflatoxins that may be potent
toxins and carcinogens in food of birds, fish, humans, and other mammals.
A few species are studied as model organisms that can be used to gain knowledge of basic
processes such as genetics, physiology, biochemistry, and molecular biology with results that are
applicable to many organisms (Taylor et al., 1993). Some of the fungi that have been intensively
studied in this way include Saccharomyces cereviseae, Neurospora crassa, and Ustilago maydis.
Most phyla appear to be terrestrial in origin, although all major groups have invaded marine and
freshwater habitats. An exception to this generality is the flagellum-bearing phyla
Chytridiomycota, Blastocladiomycota, and Neocallimastigomycota (collectively referred to as
chytrids), which probably had an aquatic origin. Extant chytrid species also occur in terrestrial
environments as plant pathogenic fungi, soil fungi, and even as anaerobic inhabitants of the guts
of herbivores such as cows (all Neocallimastigomycota).
Characteristics
Fungi are characterized by non-motile bodies (thalli) constructed of apically elongating walled
filaments (hyphae), a life cycle with sexual and asexual reproduction, usually from a common
thallus, haploid thalli resulting from zygotic meiosis, and heterotrophic nutrition. Spindle pole
bodies, not centrioles, usually are associated with the nuclear envelope during cell division. The
characteristic wall components are chitin (beta-1,4-linked homopolymers of N-acetylglucosamine
in microcrystalline state) and glucans primarily alpha-glucans (alpha-1,3- and alpha-1,6- linkages)
(Griffin, 1994).
Figure 4: Portion of a hypha of a zygomycete stained with a blue dye to show the many nuclei
present. Many other fungi have septations that devide the hyphae into compartments that usually
contain one to several nuclei per compartment. Copyright M. Blackwell 1996.
Figure 5: Transmission electron micrograph showing duplicated spindle pole body of a prophase I
meiotic nucleus of a basidiomycete Exobasidium. Only chytrids among fungi have centrioles and lack
spindle pole bodies. Copyright Beth Richardson 1996.
Exceptions to this characterization of fungi are well known, and include the following: Most species
of chytrids have cells with a single, smooth, posteriorly inserted flagellum at some stage in the life
cycle, and centrioles are associated with nuclear division. The life cycles of almost all flagellated
fungi are poorly studied. The thalli of Chytridiomycota have been thought to be haploid, but recent
population genetic data support diploidy for one species (Morehouse et al. 2003; Morgan et al.
2007). Most members of Blastocladiomycota appear to have sporic meiosis and, therefore, an
alternation between haploid and diploid generations. Certain members of Mucoromycotina,
Ascomycota, and Basidiomycota may lack hyphal growth during part or all of their life cycles, and,
instead, produce budding yeast cells. Most fungal species with yeast growth forms contain only
minute amounts of chitin in the walls of the yeast cells. A few species of Ascomycota
(Ophiostomataceae) have cellulose in their walls, and certain members of Blastocladiomycota and
Entomophthoromycotina lack walls during part of their life cycle (Alexopoulos et al., 1996).
Fossil Record
Based on the available fossil record, fungi are presumed to have been present in Late Proterozoic
(900-570 mya). Terrestrial forms of purported ascomycetes are reported in associations with
microarthropods in the Silurian Period (438-408 mya) (Sherwood-Pike and Gray, 1985). Fossil
hyphae in association with wood decay and fossil chytrids and Glomales-Endogonales
representatives associated with plants of the Rhynie Chert are reported from the Devonian Period
(408-360 mya) (Hass et al., 1994; Remy et al., 1994a, 1994b; Taylor et al., 1994a, 1995b).
Fungal fossil diversity increased throughout the Paleozoic Era (Taylor et al., 1994b) with all
modern classes reported in the Pennsylvanian Epoch (320-286 mya).
A first attempt to match molecular data on fungal phylogeny to the geological record shows
general agreement, but does point out some conflicts between the two types of data (Berbee and
Taylor 1993).
Biogeography
Wherever adequate moisture, temperature, and organic substrates are available, fungi are
present. Although we normally think of fungi as growing in warm, moist forests, many species
occur in habitats that are cold, periodically arid, or otherwise seemingly inhospitable. It is
important to recognize that optimum conditions for growth and reproduction vary widely with
fungal species. Diversity of most groups of fungi tends to increase in tropical regions, but detailed
studies are only in their infancy (Isaac et al., 1993).
Although many saprobic and plant pathogenic species with low substrate specificity and effective
dispersal systems have broad distributions, gene flow appears to be restricted in many fungi. For
these species large bodies of water such as the Atlantic and Pacific Oceans create barriers to gene
exchange. Some distributions are limited by substrate availability, and dramatic examples come
from parasites of Gondowanan plants; one of these is the Southern Hemisphere distribution of the
ascomycete Cyttaria, corresponding with part of the distribution of its host plant Nothofagus. The
fossil record shows that fungi were present in Antarctica, as is the case for other organisms with
Gondwanan distributions. Arthropod associates also may show distributions throughout part or all
of a host range, and some fungal species (ex. wood wasp associates) occur outside the range of
the associated arthropod.
Notable Fungi
The largest known basidioma (mushroom or fruiting body) was that of a Rigidioporus
ulmarius (Agaricomycetes), hidden-away in a shady corner of the Royal Botanic Gardens,
Kew, Richmond, Surrey, England. This fruiting body was mentioned in the Guinness Book
of World Records (Matthews, 1994). At the beginning of every New Year the Annual
Mensuration Ceremony of the fruiting body took place, and on 19 January 1996 it had
increased to 170 cm maximum length (up from 159 in 1995) and 146 cm maximum width
(up from 140 in 1995). It also grew 4 cm taller from the soil level, measuring 54 cm. The
weight of the fruiting body was estimated to be 284 kg (625 pounds)! Amid rumors of its
destruction, Dr. Brian M. Spooner, Head of Mycology, Royal Botanic Gardens, has brought
us up to date on the fate of the record specimen. Unfortunately, the basidioma began to
rot at the edges a few years ago, likely because the hyphal body of the fungus digested
away its elm root substrate, reminding us that a fungus needs a good dispersal system to
escape the substrate that eventually inevitably is destroyed. In the life of the fruiting body
many trillions of spores must have been produced, and some of these surely fell on an
appropiate substrate to establish a new infection. The final insult to the fruiting body
came from a fox that burrowed under one side and caused it to collapse.
Figure 6: The largest basidioma world record holder Rigidioporus ulmarius at Kew when it
was still intact. The mushroom is shown in its largest dimension (170 cm or over 5 1/2 feet).
Copyright D. Pegler 1996.
Other large basidiomata are those of a Canadian puffball almost 9 feet in circumference
(over 48 pounds) and a basidiocarp of the sulfur mushroom in England (100 pounds). A
previous Guinness Book of World Records record-sized fruiting body of Bridgeoporus
nobilissimus, an endangered species of the Pacific Northwest of the United States, is over
160 kg (300 pounds) and may have regained the title of largest with the demise of the
Kew specimen. This polypore also may do itself in because its great weight is likely to
eventually cause it to fall as the mycelium depletes its food source, often the noble fir
tree.
Reproductive structures clearly can be very large, but what about the body of the fungus,
which often is hidden from view within the substrate? One fungus body constructed of
tubular filaments (hyphae) was brought to our attention when molecular techniques were
used to show that it was extensive (37 acres and an estimated blue whale equivalent size
of 110 tons). The Michigan fungus clone (Armillaria bulbosa, Agaricomycetes) grew in tree
roots and soil. This report drew attention to an even larger fungal clone of Armillaria
ostoyae, reported earlier in the state of Washington, which covered over 1,500 acres.
Each clone began from the germination of a single spore over a thousand years ago.
Although they probably have fragmented and are no longer continuous bodies, such
organisms give us cause to think about what constitutes an individual.
Penicillium chrysogenum (Ascomycota) is known for its production of the antibiotic
penicillin. Although other antibiotics are produced by a variety of organisms, penicillin was
the first to be developed. In the spring of 1996 a long dried out culture of the original
isolate prepared by its discoverer, Sir Alexander Fleming in the late 1920s, was auctioned
by Sotheby's of London and sold to a pharmaceutical company for 23,000 pounds. This
price is insignificant when one considers the worth of this fungus, not only in sales of
penicillin, but in terms of illnesses cured and lives saved. In the past a simple scratch or
blister sometimes could result in a fatal infection such as the blister that resulted in the
death of John Calvin Coolidge (1918-1924), the son of a U. S. president. However, misuse
of penicillin and other antibiotics has resulted in selection of resistant microorganisms,
and the threat of untreatable bacterial infections and diseases (for
example Staphylococcus aureus and Escherichia coli and tuberculosis and syphilis) is still
present in our homes and recreation areas.
Fungal spores fill the air we breathe. On many days in some localities the number of
fungal spores in the air far exceeds the pollen grains. Fungal spores also cause allergies;
however, unlike seasonal pollen production, some fungi can produce spores all year long.
The largest number of fungal spores ever sampled was over 5.5 million per cubic foot in
Wales (Matthews, 1994).
Basidiomycetes have always attracted a lot of attention because some of them have large
basidiocarps, but the realization that all fungi are important in ecosystem function has
drawn more attention to microscopic forms as well. For example a report on the secret
sex life of a yeast-like ascomycete human pathogen, Coccidioides immitis, made a
headline of the New York Times (6 February 1996, p. B7). This fungus causes Valley Fever
and is endemic in parts of the southwestern United States. Although no one has been able
to observe sexual reproduction in this species, molecular studies show genetic diversity
that is best explained by occurrence of sexual reproduction in the life cycle.
Another yeast-like ascomycete reported in the Dallas Morning News (28 August 1995, p.
8D) lives in the gut of cigar beetles and is essential to the beetle's health. Without the gut
fungi to detoxify the plant material of toxins, the beetles would be poisoned. Keep on the
lookout for other reports of fascinating fungal feats.
The kingdom Fungi is a diverse clade of heterotrophic organisms that shares some characters with
animals such as chitinous structures, storage of glycogen, and mitochondrial codon UGA encoding
tryptophan. Both animals and fungi have spores or gametes with a single smooth, posteriorly
inserted flagellum, but only species of the basal chytrid phyla have retained this primitive
character (Barr, 1992; Cavalier-Smith, 1987, 1995). Fungi, animals, and other heterotrophic
protist-like organisms such as choanoflagellates and Mesomycetozoea are now considered part of
the larger group termed opisthokonts (Cavalier-Smith, 1987) in reference to the posterior
flagellum.
The branch uniting the fungi and animals is well-supported based on a number of molecular
phylogenetic datasets, including the nuclear small subunit ribosomal RNA gene (Wainwright et al.,
1993; Bruns et al. 1993), unique and shared sequence insertions in proteins such as elongation
factor 1 (Baldauf and Palmer, 1993), entire mitochondrial genomes (Lang et al., 2002), and
concatenated protein-coding genes (Steenkamp et al., 2006).
Prior classification systems of Fungi based primarily on morphology are in need of updating to
more accurately reflect phylogenetic relationships as determined by molecular systematics.
Molecular characters have been essential for phylogenetic analysis in cases when morphological
characters are convergent, reduced, or missing among the taxa considered. This is especially true
of species that never reproduce sexually, because characters of sexual reproduction traditionally
have been the basis for classification of Fungi. Use of molecular characters allows asexual fungi to
be placed among their closest relatives.
Figure 7: The endoparasitic chytrid Rozella allomycis inside the hyphae of another chytrid Allomyces.
Thick spiny spores of the parasite are seen inside some cells while zoospores are produced in other
cells. Timothy Y. James
Fungi with non-septate or irregularly septate hyphae and thick-walled spores were traditionally
placed in the phylum Zygomycota. However, evidence for a monophyletic Zygomycota is lacking
(Seif et al., 2005), and the deconstruction of the Zygomycota into four unordered subphyla
(Entomophthoromycotina, Kickxellomycotina, Mucoromycotina, Zoopagomycotina) has been
proposed (Hibbett et al., 2007). The separation of the superficially similar arbuscular mycorrhizal
fungi (that lack septa in hyphae but also lack zygospores) into the phylum Glomeromycota has
been previously proposed (Schler et al., 2001). Whether this phylum is more closely related to
the Ascomycota and Basidiomycota lineage or to other zygomycete lineages is controversial
(Redecker et al., 2006).
Evidence from shared morphological characters such as regularly septate hyphae and a dikaryotic
stage (two separate and different nuclei in a single hyphal segment) in the life cycle usually has
been interpreted as support for a close relationship between Basidiomycota and Ascomycota.
Numerous phylogenetic studies such as SSU rDNA (Berbee and Taylor, 1992), RNA polymerase
genes (Liu et al., 2006), and mitochondrial genome sequencing (Seif et al., 2005) provide strong
support for this relationship. A subkingdom termed Dikarya is proposed (Hibbett et al., 2007),
creating a division between a highly speciose subkingdom (Dikarya) and the remaining early
diverging lineages whose relationships are not precisely known.
Fungal classification is far from static, and even which organisms are actually members of Fungi is
changing. For example, the group trichomycetes describes gut inhabitants of arthropods that
share similarities with zygomycetes. Molecular phylogenetic studies have demonstrated that two of
the four orders of trichomycetes are actually members of the Mesomycetozoea protist group
(Benny and ODonnell, 2000; Cafaro, 2005). Other organisms that were previously considered to
be Fungi because of their heterotrophic, mold-like growth forms are now classified as
stramenopiles (Oomycota, Hyphochytriomycota, and Labyrinthulomycota) or slime molds
(Myxomycota, Plasmodiomycota, Dictyosteliomycota, Acrasiomycota) (Bhattacharya et al., 1992;
Leipe et al., 1994; Van der Auwera et al., 1995). More interesting for mycologists are the findings
that some species previously considered protozoa are actually Fungi. For example, the
species Hyaloraphidium curvatum was assumed to be a green alga that had adopted a
heterotrophic lifecycle concomitantly with losing its chloroplast. It is now known to be a chytrid
fungus related to Monoblephariomycetes but lacking a flagellated stage (Ustinova et al., 2000).
Other examples include the parasitic organisms presumed to be protozoa, such as the cockroach
parasite Nepridiophaga (Wylezich et al., 2004) and the Daphnia parasite Polycarum (Johnson et
al., 2006) recently demonstrated to be members of the fungal kingdom based on SSU rDNA
phylogenies.
The most revolutionary addition to the fungal lineage has occurred with phylogenetic evidence
indicating the protist group microsporidia is closely related to Fungipossibly derived from
zygomycetes (Keeling, 2003) or sister to the genus Rozella on the earliest branch in the fungal
kingdom (James et al., 2006a). Microsporidia are highly specialized intracellular parasites
(primarily of animals) that lack mitochondria but have chitin and trehalose in their spores (similar
to Fungi). All molecular studies have shown that microsporidia evolve at an extremely accelerated
rate of evolution, making their placement in the Tree of Life difficult. The relationship with fungi is
supported by many single and multiple gene phylogenies (e.g., Liu et al., 2006), but an exact
placement within the fungi has not received strong support (Keeling and Fast, 2002).
More recently the nuclearid amoebae have been demonstrated to be a sister group to the Fungi
with strong support (Steenkamp et al., 2006). This finding is significant because Nuclearialacks a
cell wall and has phagotrophic nutrition in which the food source (such as a bacterium or algal cell)
is engulfed wholly, unlike fungi and microsporidia which utilize absorptive nutrition. Further
sampling of basal fungal lineages will be needed to determine whether a Nuclearia-like organism
was the cenancestor (most recent common ancestor) of Fungi.
Mycota
Eumycota
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Popular Sites
MykoWeb. WWW pages devoted to the science of mycology and the hobby of
mushrooming.
Mycological Society of San Francisco. North America's largest local amateur mycological
association.
Mushroom Observer. The purpose of this site is to record observations about mushrooms,
help people identify mushrooms they aren't familiar with, and expand the community
around the scientific exploration of mushrooms (mycology).
The Fungal Jungal. To further educate people about fungi, edible and otherwise, To
encourage sustainable and responsible mushroom harvest, and preserve mushroom
habitat.
Tom Volk's Fungi.
Dave Fischer's North American Mushroom Basics.
MushroomExpert.Com.
Forest Fungi.
Pilze, Pilze, Pilze. In deutsch.
Westflischen Pilzbriefe. In deutsch.
Micologi Associati. Nell'italiano.
The WWW Virtual Library: Mycology. A well indexed entrance to almost all mycology and
fungal biology resources on the Internet.
Mycology.Net. An internet site containing information about diversity of fungi.
Mycorrhiza Information Exchange.
Mycology Online. A WWW resource of clinically significant mycological information.
Yahoo Mycology.
Mycologists Online. World-wide directory for mycology and lichenology.
Fungal Databases. Systematic Botany and Mycology Laboratory. Agricultural Research
Service. United States Department of Agriculture. Beltsville, Maryland, USA.
The Fifth Kingdom online. A mycological encyclopedia.
Mycological and Lichenological Collection Catalogs. UCMP Berkeley.
University of Alberta Microfungus Collection & Herbarium (UAMH).
University of Michigan Fungus Collection.
Mycological Herbarium. The Natural History Museums and Botanical Garden, University of
Oslo.
Herbarium Mycologicum. National Botanic Garden of Belgium.
Index Fungorum. Names of fungi.
Centraalbureau voor Schimmelcultures (CBS). Fungal Biodiversity Center - Utrecht, The
Netherlands.
Fungal Genetics Stock Center.
Canadian Collection of Fungal Cultures.
Images
Professional Societies
The International Mycological Association. A group that represents mycologists and fungal
biologists throughout the world.
British Mycological Society.
Mycological Society of America.
Asociacin Latinoamericana de Micologa.
Australasian Mycological Society.
The International Society for Mushroom Science. To further the cultivation of edible
(including medicinal) macrofungi.
Title Illustrations
Scientific Name Chytridium (Chytridiomycota)
Comments Individual growing on a single pine pollen grain. Successive photos show zoospore release from the
sporangium, and the arrow points to a flagellum.
Copyright 1996 H. Whisler, M. Fuller
Many thanks to Soren Rosendahl and Atul Batra for scanning photos and David Maddison and Atul Batra for
page design advice.
Meredith Blackwell
Louisiana State University, Baton Rouge, Louisiana, USA
Rytas Vilgalys
Duke University, Durham, North Carolina, USA
Timothy Y. James
University of Michigan, Ann Arbor, Michigan, USA
John W. Taylor
University of California, Berkeley, California, USA
Page copyright 2012 Meredith Blackwell, Rytas Vilgalys, Timothy Y. James, and John W. Taylor
Blackwell, Meredith, Rytas Vilgalys, Timothy Y. James, and John W. Taylor. 2012. Fungi. Eumycota: mushrooms,
sac fungi, yeast, molds, rusts, smuts, etc.. Version 30 January
2012.http://tolweb.org/Fungi/2377/2012.01.30 in The Tree of Life Web Project, http://tolweb.org/
Predatory Fungi
In species of Arthrobotrys, traps take many forms but the best known are
adhesive (sticky) knobs, adhesive nets (click here and click here), or
rings (click here). The most sophisticated trap is a constricting ring
composed of three cells. When the nematode sticks its head or tail in a
ring, the physical contact triggers the ring and the cells expand rapidly
inwards crushing the victim (click here and click here), which is then
penetrated and digested within hours. Fungal enzymes break down the
body contents of the nematode and the nutrients are translocated
elsewhere within the hyphal system for growth or to produce
conidiophores and conidia (asexual spores,click here and click
here). Spores are carried off by wind, water, mites, insects or whatever
and start the predatory cycle again at a new location. It has been shown
that trapping devices produce a chemical lure that is attractive to
nematodes and this bait attracts the victims to the site of their
destruction. Much of the earlier work on this group of fungi has been
reviewed in detail (Barron 1977).
For many years these dramatic and fascinating methods for capture of
nematodes by predatory fungi distracted observers from the true
significance of this unique relationship. We had occasion however, to do a
detailed study on another genus of the asexual, predatory fungi
called Nematoctonus and this work gave exciting clues to a much more
fundamental relationship.
Carnivorous Mushrooms
We are well aware that in our own nutrition for a healthy diet, we need the
proper balance of protein (N) and carbohydrate (C). This is no less true
for other organisms, and a proper balance of C to N is essential for
successful growth of all living things. This is often expressed as the ratio
of C to N or C:N. Although the ratio varies a lot amongst organisms, most
require a C:N ranging around 30:1 for good growth.
There is very little nitrogen available in wood and, therefore, the C:N in
wood is very high. It can be 300:1 to 1000:1 or even higher. In theory,
because of these extremely low nitrogen levels, it does not seem possible
for saprobic fungi to decay wood! There are many theories as to how
fungi manage to rot wood and I do not have space to evaluate these at this
time. Suffice to repeat that:
The sticky knobs of Hohenbuehelia are nematode specific and these fungi
are, apparently, restricted to this source of biological nitrogen. Pleurotus,
on the other hand, with its non-specific toxin, has a much wider range of
potential victims to attack. The host limits for Pleurotus have never been
fully established. Toxin droplets can be instrumental, not only in
supplying a nitrogen source to Pleurotus, but can also function as
antifeedents that discourage grazing and help protect Pleurotushyphae
from fungus-feeding microfauna such as mites, springtails, water bears, as
well as fungal-feeding nematodes. It is tempting to suggest that the
predatory habit might have originated as a defence response by fungi
against grazing microfauna. It was found that secretory appendages
similar to those of Pleurotus are produced on hyphae of the lawn
mushroom Conocybe lactea (click here). These apparently function to
repel attacks by fungal feeding nematodes and although the nematodes
can be killed by the toxin, Conocybe shows no interest in utilizing their
bodies as a food source (Hutchison et al. 1995).
The results from our studies and those of others with wood decay fungi
(mushrooms, bracket fungi etc) and litter fungi (including the commercial
mushroom, Agaricus brunnescens, showed that a large percentage could
attack and digest bacterial colonies in the same way as Pleurotus (Barron
1988). In the case of Pleurotusthis involves the following sequence of
events:
All of our studies were carried out in vitro on water agar (almost no
nutrient), and I suggest that the results reflect the reality in the natural
habitat. In my opinion, the complex sequence of events required to utilize
nematodes or bacterial colonies is too sophisticated to be explained as
cultural artifacts.
The conclusion, therefore, is that many of the fungi involved in decay that
we have referred to as saprobes (or saprophytes) have a Jekyll and Hyde
existence and, as well as being apparently innocuous saprobes, they also
have the faculty of becoming aggressive predators of a variety of other
microscopic life forms including nematodes, bacteria, pollen grains,
yeasts etc. A fungus that is basically a saprophyte but has a parasitic phase
in its life cycle is referred to as a facultative parasite. In the case of plant
parasites this process is usually sequential, where a saprophytic phase is
followed by a parasitic phase or vice versa. In the case of predatory wood
decay fungi, however, the saprophytic phase and parasitic phase run
concurrently and the two phases are parallel rather than sequential. There
is also evidence to show that wood decay fungi can attack other wood
decay fungi (= mycoparasitism) and this may eventually prove to be an
important alternative for some species. Much of the background literature
in this critical area, by a number of individuals and research groups, has
been reviewed previously (Barron 1992).
So! What about the Carbon Cycle? Ecological studies have found that, in
forest soils, the biomass of fungi is 90% of the total and outweighs the
biomass (= living stuff) of all other microorganisms and mesoorganisms
combined. ( Microorganisms include fungi, bacteria, nematodes, protozoa,
rotifers, algae and all organisms that need a microscope to observe and
identify. Mesoorganisms (= mesofauna) are small to very small but are
still big enough to see with the unaided eye and include springtails, mites,
worms, small insects and insect larvae). This massive fungal component is
based largely on the role of fungi in two major biological systems: 1. As
decay organisms of plant debris. 2. As mycorrhizal partners with trees and
other plants.
The forest tree obtains almost all the water and minerals required through
extensive networks of hyphae of mycorrhizal fungi. These fungi are
particularly adept at obtaining low mobility phosphorus ions. They either
grow to the source ions or alternatively they utilize difficult sources of
phosphorus such as rock phosphate. As with decay fungi, nitrogen is also
in short supply. Attacking living animals could kill two birds with one
stone!
SELECTED REFERENCES
Suillus spraguei
Suillus umbonatus,
Syncephalastrum
racemosum *** .......................................................................................................
...................................UPGRADED 2009
Syncephalastrum racemosum **** merosporangia radiating from apical
vesicle......................................................... UPGRADED 2009
Syncephalastrum racemosum rhizoids
Syncephalis nodosa *** mycoparasite on other
Zygomycota .........................................HOW DO I KILL THEE? # 37 NEW SERIES
Syzygospora mycetophila
Syzygospora mycetophila
Taphrina deformans
Tectella patellaris
Tetracladium sp - aquatic Hyphomycete
Tetracoccosporium paxianum
Tetrapyrgos subdendrophora **** not often illustrated and pretty gill structure
underneath
Thamnidium elegans *** Sporangioles and
sporangiospores......................................................................................... UPGR
ADED 2009
Thaxterogaster porphyreus
Thecothejus pelletieri **** 32-spores ascus with operculum
.......................................................................................UPGRADED 2009
Thelephora caryophyllea
Thelephora caryophyllea *** Carnation Thelephora
Thelephora palmata
Trametes versicolor **** colourful Turkey Tail always looks good
Tremella foliacea *** Leafy Tremella
Tremella mesenterica
Tremella reticulata
Tremellodendropsis semivestitum
Tremiscus helvelloides
Tremiscus helvelloides **** very nice shot by Jim Ginns
Triacutus subcuticularis -
............................................................................. HOW DO I KILL THEE? #
18 NEW SERIES
Trichaptum biforme colourful brackets some with a purple edge
Trichoglossum hirsutum
Tricholoma
alboviolaceum......................................................................................................................
.............................................................. NEW ADDITION 2009
Tricholoma saponaceum *** Soapy
Tricholoma...........................................................................................................................
.................. NEW ADDITION 2009
Tricholoma myomyces
Tricholoma
portentosum ........................................................................................................................
............................................................... NEW ADDITION 2009
Tricholoma virgatum.
Tricholomopsis
rutilans ................................................................................................................................
........................................................ NEW ADDITION 2009
Trichurus
spiralis .....................................................................................................
............................................ NEW ADDITION 2009
Tricladium sp - aquatic Hyphomycete
Tylopilus felleus *** Bitter
Bolete.......................................................................................................................
................................................. NEW ADDITION 2009
Typhula incarnata *** snow mould (photo Yijun Yang and Tom Hsiang)
Uncinula necator **** powdery mildew of
grapes...................................................................................................................................
......... UPGRADED 2009
Varicosporium elodeae - an aquatic Hyphomycete
Volucrispora sp.*** aquatic
hyphomycete........................................................................................................................
................................. NEW ADDITION 2009
Volutella ciliata *** fruiting on water surface from a colonized submerged fir
leaf
Venturia inaequalis *** apple scab fungus - symptoms on fruit
................................................................................................................... NEW
ADDITION 2009
Venturia inaequalis *** micrographs of 'perithecium' and Fusicladium conidia
....................................................................................... NEW ADDITION 2009
Xeromphalina campanella ..
Xerula furfuracea ***
Xerula furfuracea
Xylaria hypoxylon *** Candlesnuff or Antler Fungus
Xylaria hypoxylon *** perithecial stromata
Xylaria longipes ***
Xylaria polymorpha Dead Man's
Fingers................................................................................................................ U
PGRADED 2009
Xylobolus frustulatus *** Ceramic
Fungus ..
UPGRADED 2009
Zoopage sp. *** predator on
amobeae .......................................................... HOW DO I KILL THEE #
38 NEW SERIES
Zoophagus
insidians **** ............................................................................. HOW DO I
KILL THEE? # 11 NEW SERIES
Zygorhynchus **** SEM of
Zygospore................................................................................................................
.................................................NEW ADDITION 09