Subphyla incertae sedis
A FUNGUS (plural : FUNGI or FUNGUSES ) is any member of the group of
eukaryotic organisms that includes microorganisms such as yeasts and
molds , as well as the more familiar mushrooms . These organisms are
classified as a kingdom , FUNGI, which is separate from the other
eukaryotic life kingdoms of plants and animals .
A characteristic that places fungi in a different kingdom from
plants, bacteria, and some protists is chitin in their cell walls .
Similar to animals, fungi are heterotrophs ; they acquire their food
by absorbing dissolved molecules, typically by secreting digestive
enzymes into their environment. Fungi do not photosynthesise . Growth
is their means of mobility, except for spores (a few of which are
flagellated), which may travel through the air or water. Fungi are the
principal decomposers in ecological systems. These and other
differences place fungi in a single group of related organisms, named
Eumycota (true fungi or Eumycetes), which share a common ancestor
(form a monophyletic group), an interpretation that is also strongly
supported by molecular phylogenetics . This fungal group is distinct
from the structurally similar myxomycetes (slime molds) and oomycetes
(water molds). The discipline of biology devoted to the study of fungi
is known as mycology (from the Greek μύκης mykes, mushroom). In
the past, mycology was regarded as a branch of botany , although it is
now known fungi are genetically more closely related to animals than
Abundant worldwide, most fungi are inconspicuous because of the small
size of their structures, and their cryptic lifestyles in soil or on
dead matter. Fungi include symbionts of plants, animals, or other
fungi and also parasites . They may become noticeable when fruiting ,
either as mushrooms or as molds. Fungi perform an essential role in
the decomposition of organic matter and have fundamental roles in
nutrient cycling and exchange in the environment. They have long been
used as a direct source of human food, in the form of mushrooms and
truffles ; as a leavening agent for bread; and in the fermentation of
various food products, such as wine , beer , and soy sauce . Since the
1940s, fungi have been used for the production of antibiotics , and,
more recently, various enzymes produced by fungi are used industrially
and in detergents . Fungi are also used as biological pesticides to
control weeds, plant diseases and insect pests. Many species produce
bioactive compounds called mycotoxins , such as alkaloids and
polyketides , that are toxic to animals including humans. The fruiting
structures of a few species contain psychotropic compounds and are
consumed recreationally or in traditional spiritual ceremonies . Fungi
can break down manufactured materials and buildings, and become
significant pathogens of humans and other animals. Losses of crops due
to fungal diseases (e.g., rice blast disease ) or food spoilage can
have a large impact on human food supplies and local economies.
The fungus kingdom encompasses an enormous diversity of taxa with
varied ecologies, life cycle strategies, and morphologies ranging from
unicellular aquatic chytrids to large mushrooms. However, little is
known of the true biodiversity of Kingdom Fungi, which has been
estimated at 2.2 million to 3.8 million species, of which only 120,000
have been described. 8000 of them are detrimental to plants and 300
can be pathogenic to humans . Ever since the pioneering 18th and 19th
century taxonomical works of
Carl Linnaeus , Christian Hendrik Persoon
Elias Magnus Fries , fungi have been classified according to
their morphology (e.g., characteristics such as spore color or
microscopic features) or physiology . Advances in molecular genetics
have opened the way for DNA analysis to be incorporated into taxonomy,
which has sometimes challenged the historical groupings based on
morphology and other traits.
Phylogenetic studies published in the
last decade have helped reshape the classification within Kingdom
Fungi, which is divided into one subkingdom , seven phyla , and ten
* 1 Etymology
* 2 Characteristics
* 3 Diversity
* 4.1 History
* 5 Morphology
* 5.1 Microscopic structures
* 6 Growth and physiology
* 7.4 Other sexual processes
* 9 Taxonomy
* 9.1 Taxonomic groups
* 9.2 Fungus-like organisms
* 10.1.1 With plants
* 10.1.2 With algae and cyanobacteria
* 10.1.3 With insects
* 10.1.4 As pathogens and parasites
* 10.1.5 As targets of mycoparasites
* 12 Pathogenic mechanisms
* 13 Human use
* 13.1 Therapeutic uses
* 13.1.1 Modern chemotherapeutics
* 13.1.2 Traditional and folk medicine
* 13.2 Cultured foods
* 13.3 In food
* 13.4 Poisonous fungi
* 13.5 Pest control
* 13.8 Others
* 14 See also
* 15 References
* 15.1 Cited literature
* 16 External links
The English word fungus is directly adopted from the
(mushroom), used in the writings of
Horace and Pliny . This in turn
is derived from the Greek word sphongos (σφογγος "sponge"),
which refers to the macroscopic structures and morphology of mushrooms
and molds; the root is also used in other languages, such as the
German Schwamm ("sponge") and Schimmel ("mold"). The use of the word
mycology, which is derived from the Greek mykes (μύκης
"mushroom") and logos (λόγος "discourse"), to denote the
scientific study of fungi is thought to have originated in 1836 with
Miles Joseph Berkeley 's publication The English
Flora of Sir James Edward Smith, Vol. 5. A group of all the fungi
present in a particular area or geographic region is known as
mycobiota (plural noun, no singular), e.g., "the mycobiota of
FUNGAL HYPHAE CELLS
* Hyphal wall
Before the introduction of molecular methods for phylogenetic
analysis, taxonomists considered fungi to be members of the plant
kingdom because of similarities in lifestyle: both fungi and plants
are mainly immobile , and have similarities in general morphology and
growth habitat. Like plants, fungi often grow in soil and, in the case
of mushrooms , form conspicuous fruit bodies , which sometimes
resemble plants such as mosses . The fungi are now considered a
separate kingdom, distinct from both plants and animals, from which
they appear to have diverged around one billion years ago. Some
morphological, biochemical, and genetic features are shared with other
organisms, while others are unique to the fungi, clearly separating
them from the other kingdoms:
* With other eukaryotes : Fungal cells contain membrane-bound nuclei
with chromosomes that contain DNA with noncoding regions called
introns and coding regions called exons . Fungi have membrane-bound
cytoplasmic organelles such as mitochondria , sterol -containing
membranes, and ribosomes of the
80S type. They have a characteristic
range of soluble carbohydrates and storage compounds, including sugar
alcohols (e.g., mannitol ), disaccharides , (e.g., trehalose ), and
polysaccharides (e.g., glycogen , which is also found in animals ).
* With animals: Fungi lack chloroplasts and are heterotrophic
organisms and so require preformed organic compounds as energy
* With plants: Fungi have a cell wall and vacuoles . They
reproduce by both sexual and asexual means, and like basal plant
groups (such as ferns and mosses ) produce spores . Similar to mosses
and algae, fungi typically have haploid nuclei.
* With euglenoids and bacteria: Higher fungi, euglenoids, and some
bacteria produce the amino acid L-lysine in specific biosynthesis
steps, called the α-aminoadipate pathway .
* The cells of most fungi grow as tubular, elongated, and
thread-like (filamentous) structures called hyphae , which may contain
multiple nuclei and extend by growing at their tips. Each tip contains
a set of aggregated vesicles —cellular structures consisting of
proteins , lipids , and other organic molecules—called the
Spitzenkörper . Both fungi and oomycetes grow as filamentous hyphal
cells. In contrast, similar-looking organisms, such as filamentous
green algae , grow by repeated cell division within a chain of cells.
There are also single-celled fungi (yeasts ) that do not form hyphae,
and some fungi have both hyphal and yeast forms.
* In common with some plant and animal species, more than 70 fungal
species display bioluminescence .
* Some species grow as unicellular yeasts that reproduce by budding
or binary fission .
Dimorphic fungi can switch between a yeast phase
and a hyphal phase in response to environmental conditions.
* The fungal cell wall is composed of glucans and chitin ; while
glucans are also found in plants and chitin in the exoskeleton of
arthropods , fungi are the only organisms that combine these two
structural molecules in their cell wall. Unlike those of plants and
oomycetes, fungal cell walls do not contain cellulose.
Omphalotus nidiformis , a bioluminescent mushroom
Most fungi lack an efficient system for the long-distance transport
of water and nutrients, such as the xylem and phloem in many plants.
To overcome this limitation, some fungi, such as
Armillaria , form
rhizomorphs , which resemble and perform functions similar to the
roots of plants. As eukaryotes, fungi possess a biosynthetic pathway
for producing terpenes that uses mevalonic acid and pyrophosphate as
chemical building blocks .
Plants and some other organisms have an
additional terpene biosynthesis pathway in their chloroplasts, a
structure fungi and animals do not have. Fungi produce several
secondary metabolites that are similar or identical in structure to
those made by plants. Many of the plant and fungal enzymes that make
these compounds differ from each other in sequence and other
characteristics, which indicates separate origins and convergent
evolution of these enzymes in the fungi and plants.
Bracket fungi on a tree stump
Fungi have a worldwide distribution, and grow in a wide range of
habitats, including extreme environments such as deserts or areas with
high salt concentrations or ionizing radiation , as well as in deep
sea sediments. Some can survive the intense UV and cosmic radiation
encountered during space travel. Most grow in terrestrial
environments, though several species live partly or solely in aquatic
habitats, such as the chytrid fungus
Batrachochytrium dendrobatidis ,
a parasite that has been responsible for a worldwide decline in
amphibian populations. This organism spends part of its life cycle as
a motile zoospore , enabling it to propel itself through water and
enter its amphibian host. Other examples of aquatic fungi include
those living in hydrothermal areas of the ocean.
Around 120,000 species of fungi have been described by taxonomists ,
but the global biodiversity of the fungus kingdom is not fully
understood. A 2017 estimate suggests there may be between 2.2 and 3.8
million species. In mycology, species have historically been
distinguished by a variety of methods and concepts. Classification
based on morphological characteristics, such as the size and shape of
spores or fruiting structures, has traditionally dominated fungal
taxonomy. Species may also be distinguished by their biochemical and
physiological characteristics, such as their ability to metabolize
certain biochemicals, or their reaction to chemical tests . The
biological species concept discriminates species based on their
ability to mate . The application of molecular tools, such as DNA
sequencing and phylogenetic analysis, to study diversity has greatly
enhanced the resolution and added robustness to estimates of genetic
diversity within various taxonomic groups.
Mycology is the branch of biology concerned with the systematic study
of fungi, including their genetic and biochemical properties, their
taxonomy, and their use to humans as a source of medicine, food, and
psychotropic substances consumed for religious purposes, as well as
their dangers, such as poisoning or infection. The field of
phytopathology , the study of plant diseases, is closely related
because many plant pathogens are fungi. In 1729, Pier Antonio
Micheli first published descriptions of fungi.
The use of fungi by humans dates back to prehistory; Ötzi the Iceman
, a well-preserved mummy of a 5,300-year-old
Neolithic man found
frozen in the Austrian Alps, carried two species of polypore mushrooms
that may have been used as tinder (
Fomes fomentarius ), or for
medicinal purposes (
Piptoporus betulinus ). Ancient peoples have used
fungi as food sources–often unknowingly–for millennia, in the
preparation of leavened bread and fermented juices. Some of the oldest
written records contain references to the destruction of crops that
were probably caused by pathogenic fungi.
Mycology is a relatively new science that became systematic after the
development of the microscope in the 17th century. Although fungal
spores were first observed by
Giambattista della Porta in 1588, the
seminal work in the development of mycology is considered to be the
Pier Antonio Micheli
Pier Antonio Micheli 's 1729 work Nova plantarum
genera. Micheli not only observed spores but also showed that, under
the proper conditions, they could be induced into growing into the
same species of fungi from which they originated. Extending the use
of the binomial system of nomenclature introduced by
Carl Linnaeus in
Species plantarum (1753), the Dutch Christian Hendrik Persoon
(1761–1836) established the first classification of mushrooms with
such skill as to be considered a founder of modern mycology. Later,
Elias Magnus Fries (1794–1878) further elaborated the classification
of fungi, using spore color and microscopic characteristics, methods
still used by taxonomists today. Other notable early contributors to
mycology in the 17th–19th and early 20th centuries include Miles
Joseph Berkeley ,
August Carl Joseph Corda ,
Anton de Bary , the
brothers Louis René and
Charles Tulasne , Arthur H. R. Buller ,
Curtis G. Lloyd , and
Pier Andrea Saccardo . The 20th century has seen
a modernization of mycology that has come from advances in
biochemistry , genetics , molecular biology , and biotechnology . The
DNA sequencing technologies and phylogenetic analysis has
provided new insights into fungal relationships and biodiversity , and
has challenged traditional morphology-based groupings in fungal
An environmental isolate of
Most fungi grow as hyphae , which are cylindrical, thread-like
structures 2–10 µm in diameter and up to several centimeters in
Hyphae grow at their tips (apices); new hyphae are typically
formed by emergence of new tips along existing hyphae by a process
called branching, or occasionally growing hyphal tips fork, giving
rise to two parallel-growing hyphae.
Hyphae also sometimes fuse when
they come into contact, a process called hyphal fusion (or anastamosis
). These growth processes lead to the development of a mycelium , an
interconnected network of hyphae.
Hyphae can be either septate or
coenocytic . Septate hyphae are divided into compartments separated by
cross walls (internal cell walls, called septa, that are formed at
right angles to the cell wall giving the hypha its shape), with each
compartment containing one or more nuclei; coenocytic hyphae are not
compartmentalized. Septa have pores that allow cytoplasm , organelles
, and sometimes nuclei to pass through; an example is the dolipore
septum in fungi of the phylum Basidiomycota.
Coenocytic hyphae are in
essence multinucleate supercells.
Many species have developed specialized hyphal structures for
nutrient uptake from living hosts; examples include haustoria in
plant-parasitic species of most fungal phyla, and arbuscules of
several mycorrhizal fungi, which penetrate into the host cells to
Although fungi are opisthokonts —a grouping of evolutionarily
related organisms broadly characterized by a single posterior
flagellum —all phyla except for the chytrids have lost their
posterior flagella. Fungi are unusual among the eukaryotes in having
a cell wall that, in addition to glucans (e.g., β-1,3-glucan) and
other typical components, also contains the biopolymer chitin.
Fungal mycelia can become visible to the naked eye, for example, on
various surfaces and substrates , such as damp walls and spoiled food,
where they are commonly called molds . Mycelia grown on solid agar
media in laboratory petri dishes are usually referred to as colonies .
These colonies can exhibit growth shapes and colors (due to spores or
pigmentation ) that can be used as diagnostic features in the
identification of species or groups. Some individual fungal colonies
can reach extraordinary dimensions and ages as in the case of a clonal
Armillaria solidipes , which extends over an area of more
than 900 ha (3.5 square miles), with an estimated age of nearly 9,000
The apothecium —a specialized structure important in sexual
reproduction in the ascomycetes—is a cup-shaped fruit body that is
often macroscopic and holds the hymenium , a layer of tissue
containing the spore-bearing cells. The fruit bodies of the
basidiomycetes (basidiocarps ) and some ascomycetes can sometimes grow
very large, and many are well known as mushrooms .
GROWTH AND PHYSIOLOGY
Mold growth covering a decaying peach . The frames were taken
approximately 12 hours apart over a period of six days.
The growth of fungi as hyphae on or in solid substrates or as single
cells in aquatic environments is adapted for the efficient extraction
of nutrients, because these growth forms have high surface area to
volume ratios .
Hyphae are specifically adapted for growth on solid
surfaces, and to invade substrates and tissues. They can exert large
penetrative mechanical forces; for example, many plant pathogens ,
Magnaporthe grisea , form a structure called an appressorium
that evolved to puncture plant tissues. The pressure generated by the
appressorium, directed against the plant epidermis , can exceed 8
megapascals (1,200 psi). The filamentous fungus Paecilomyces
lilacinus uses a similar structure to penetrate the eggs of nematodes
The mechanical pressure exerted by the appressorium is generated from
physiological processes that increase intracellular turgor by
producing osmolytes such as glycerol . Adaptations such as these are
complemented by hydrolytic enzymes secreted into the environment to
digest large organic molecules—such as polysaccharides , proteins ,
and lipids —into smaller molecules that may then be absorbed as
nutrients. The vast majority of filamentous fungi grow in a polar
fashion (extending in one direction) by elongation at the tip (apex)
of the hypha. Other forms of fungal growth include intercalary
extension (longitudinal expansion of hyphal compartments that are
below the apex) as in the case of some endophytic fungi, or growth by
volume expansion during the development of mushroom stipes and other
large organs. Growth of fungi as multicellular structures consisting
of somatic and reproductive cells—a feature independently evolved in
animals and plants —has several functions, including the development
of fruit bodies for dissemination of sexual spores (see above) and
biofilms for substrate colonization and intercellular communication .
The fungi are traditionally considered heterotrophs , organisms that
rely solely on carbon fixed by other organisms for metabolism . Fungi
have evolved a high degree of metabolic versatility that allows them
to use a diverse range of organic substrates for growth, including
simple compounds such as nitrate , ammonia , acetate , or ethanol .
In some species the pigment melanin may play a role in extracting
energy from ionizing radiation , such as gamma radiation . This form
of "radiotrophic" growth has been described for only a few species,
the effects on growth rates are small, and the underlying biophysical
and biochemical processes are not well known. This process might bear
similarity to CO2 fixation via visible light , but instead uses
ionizing radiation as a source of energy.
Fungal reproduction is complex, reflecting the differences in
lifestyles and genetic makeup within this diverse kingdom of
organisms. It is estimated that a third of all fungi reproduce using
more than one method of propagation; for example, reproduction may
occur in two well-differentiated stages within the life cycle of a
species, the teleomorph and the anamorph . Environmental conditions
trigger genetically determined developmental states that lead to the
creation of specialized structures for sexual or asexual reproduction.
These structures aid reproduction by efficiently dispersing spores or
spore-containing propagules .
Asexual reproduction occurs via vegetative spores (conidia ) or
through mycelial fragmentation . Mycelial fragmentation occurs when a
fungal mycelium separates into pieces, and each component grows into a
separate mycelium. Mycelial fragmentation and vegetative spores
maintain clonal populations adapted to a specific niche , and allow
more rapid dispersal than sexual reproduction. The "Fungi imperfecti"
(fungi lacking the perfect or sexual stage) or
all the species that lack an observable sexual cycle. Deuteromycota
is not an accepted taxonomic clade, and is now taken to mean simply
fungi that lack a known sexual stage.
Mating in fungi and
Sexual selection in fungi
Sexual reproduction with meiosis has been directly observed in all
fungal phyla except
Glomeromycota (genetic analysis suggests meiosis
Glomeromycota as well). It differs in many aspects from sexual
reproduction in animals or plants. Differences also exist between
fungal groups and can be used to discriminate species by morphological
differences in sexual structures and reproductive strategies. Mating
experiments between fungal isolates may identify species on the basis
of biological species concepts. The major fungal groupings have
initially been delineated based on the morphology of their sexual
structures and spores; for example, the spore-containing structures,
asci and basidia , can be used in the identification of ascomycetes
and basidiomycetes, respectively. Some species may allow mating only
between individuals of opposite mating type , whereas others can mate
and sexually reproduce with any other individual or itself. Species of
the former mating system are called heterothallic , and of the latter
Most fungi have both a haploid and a diploid stage in their life
cycles. In sexually reproducing fungi, compatible individuals may
combine by fusing their hyphae together into an interconnected
network; this process, anastomosis , is required for the initiation of
the sexual cycle. Many ascomycetes and basidiomycetes go through a
dikaryotic stage, in which the nuclei inherited from the two parents
do not combine immediately after cell fusion, but remain separate in
the hyphal cells (see heterokaryosis ). The 8-spore asci of
Morchella elata , viewed with phase contrast microscopy
In ascomycetes, dikaryotic hyphae of the hymenium (the spore-bearing
tissue layer) form a characteristic hook at the hyphal septum. During
cell division , formation of the hook ensures proper distribution of
the newly divided nuclei into the apical and basal hyphal
compartments. An ascus (plural asci) is then formed, in which
karyogamy (nuclear fusion) occurs. Asci are embedded in an ascocarp ,
or fruiting body.
Karyogamy in the asci is followed immediately by
meiosis and the production of ascospores . After dispersal, the
ascospores may germinate and form a new haploid mycelium.
Sexual reproduction in basidiomycetes is similar to that of the
ascomycetes. Compatible haploid hyphae fuse to produce a dikaryotic
mycelium. However, the dikaryotic phase is more extensive in the
basidiomycetes, often also present in the vegetatively growing
mycelium. A specialized anatomical structure, called a clamp
connection , is formed at each hyphal septum. As with the structurally
similar hook in the ascomycetes, the clamp connection in the
basidiomycetes is required for controlled transfer of nuclei during
cell division, to maintain the dikaryotic stage with two genetically
different nuclei in each hyphal compartment. A basidiocarp is formed
in which club-like structures known as basidia generate haploid
basidiospores after karyogamy and meiosis. The most commonly known
basidiocarps are mushrooms, but they may also take other forms (see
In glomeromycetes (formerly zygomycetes), haploid hyphae of two
individuals fuse, forming a gametangium , a specialized cell structure
that becomes a fertile gamete -producing cell. The gametangium
develops into a zygospore , a thick-walled spore formed by the union
of gametes. When the zygospore germinates, it undergoes meiosis ,
generating new haploid hyphae, which may then form asexual
sporangiospores . These sporangiospores allow the fungus to rapidly
disperse and germinate into new genetically identical haploid fungal
Both asexual and sexual spores or sporangiospores are often actively
dispersed by forcible ejection from their reproductive structures.
This ejection ensures exit of the spores from the reproductive
structures as well as traveling through the air over long distances.
The bird's nest fungus
Specialized mechanical and physiological mechanisms, as well as spore
surface structures (such as hydrophobins ), enable efficient spore
ejection. For example, the structure of the spore-bearing cells in
some ascomycete species is such that the buildup of substances
affecting cell volume and fluid balance enables the explosive
discharge of spores into the air. The forcible discharge of single
spores termed ballistospores involves formation of a small drop of
water (Buller's drop), which upon contact with the spore leads to its
projectile release with an initial acceleration of more than 10,000 g
; the net result is that the spore is ejected 0.01–0.02 cm,
sufficient distance for it to fall through the gills or pores into the
air below. Other fungi, like the puffballs , rely on alternative
mechanisms for spore release, such as external mechanical forces. The
bird\'s nest fungi use the force of falling water drops to liberate
the spores from cup-shaped fruiting bodies. Another strategy is seen
in the stinkhorns , a group of fungi with lively colors and putrid
odor that attract insects to disperse their spores.
OTHER SEXUAL PROCESSES
Besides regular sexual reproduction with meiosis, certain fungi, such
as those in the genera
Aspergillus , may exchange
genetic material via parasexual processes, initiated by anastomosis
between hyphae and plasmogamy of fungal cells. The frequency and
relative importance of parasexual events is unclear and may be lower
than other sexual processes. It is known to play a role in
intraspecific hybridization and is likely required for hybridization
between species, which has been associated with major events in fungal
Evolution of fungi
In contrast to plants and animals , the early fossil record of the
fungi is meager. Factors that likely contribute to the
under-representation of fungal species among fossils include the
nature of fungal fruiting bodies , which are soft, fleshy, and easily
degradable tissues and the microscopic dimensions of most fungal
structures, which therefore are not readily evident. Fungal fossils
are difficult to distinguish from those of other microbes, and are
most easily identified when they resemble extant fungi. Often
recovered from a permineralized plant or animal host, these samples
are typically studied by making thin-section preparations that can be
examined with light microscopy or transmission electron microscopy .
Researchers study compression fossils by dissolving the surrounding
matrix with acid and then using light or scanning electron microscopy
to examine surface details.
The earliest fossils possessing features typical of fungi date to the
Paleoproterozoic era, some 2,400 million years ago (Ma ); these
multicellular benthic organisms had filamentous structures capable of
anastomosis . Other studies (2009) estimate the arrival of fungal
organisms at about 760–1060 Ma on the basis of comparisons of the
rate of evolution in closely related groups. For much of the
Paleozoic Era (542–251 Ma), the fungi appear to have been aquatic
and consisted of organisms similar to the extant chytrids in having
flagellum-bearing spores. The evolutionary adaptation from an aquatic
to a terrestrial lifestyle necessitated a diversification of
ecological strategies for obtaining nutrients, including parasitism ,
saprobism , and the development of mutualistic relationships such as
mycorrhiza and lichenization. Recent (2009) studies suggest that the
ancestral ecological state of the
Ascomycota was saprobism, and that
independent lichenization events have occurred multiple times.
It is presumed that the fungi colonized the land during the Cambrian
(542–488.3 Ma), long before land plants. Fossilized hyphae and
spores recovered from the
Ordovician of Wisconsin (460 Ma) resemble
Glomerales , and existed at a time when the land flora
likely consisted of only non-vascular bryophyte -like plants.
Prototaxites , which was probably a fungus or lichen, would have been
the tallest organism of the late
Silurian . Fungal fossils do not
become common and uncontroversial until the early Devonian
(416–359.2 Ma), when they occur abundantly in the
Rhynie chert ,
Chytridiomycota . At about this same time,
approximately 400 Ma, the
Basidiomycota diverged, and
all modern classes of fungi were present by the Late Carboniferous
(Pennsylvanian , 318.1–299 Ma).
Lichen -like fossils have been found in the
Doushantuo Formation in
southern China dating back to 635–551 Ma.
Lichens formed a
component of the early terrestrial ecosystems, and the estimated age
of the oldest terrestrial lichen fossil is 400 Ma; this date
corresponds to the age of the oldest known sporocarp fossil, a
Paleopyrenomycites species found in the Rhynie Chert. The oldest
fossil with microscopic features resembling modern-day basidiomycetes
is Palaeoancistrus, found permineralized with a fern from the
Pennsylvanian. Rare in the fossil record are the Homobasidiomycetes
(a taxon roughly equivalent to the mushroom-producing species of the
Agaricomycetes ). Two amber -preserved specimens provide evidence that
the earliest known mushroom-forming fungi (the extinct species
Archaeomarasmius leggetti ) appeared during the late
Cretaceous , 90
Some time after the Permian–
Triassic extinction event (251.4 Ma), a
fungal spike (originally thought to be an extraordinary abundance of
fungal spores in sediments ) formed, suggesting that fungi were the
dominant life form at this time, representing nearly 100% of the
available fossil record for this period. However, the relative
proportion of fungal spores relative to spores formed by algal species
is difficult to assess, the spike did not appear worldwide, and in
many places it did not fall on the Permian–
Although commonly included in botany curricula and textbooks, fungi
are more closely related to animals than to plants and are placed with
the animals in the monophyletic group of opisthokonts . Analyses
using molecular phylogenetics support a monophyletic origin of the
Fungi. The taxonomy of the Fungi is in a state of constant flux,
especially due to recent research based on DNA comparisons. These
current phylogenetic analyses often overturn classifications based on
older and sometimes less discriminative methods based on morphological
features and biological species concepts obtained from experimental
There is no unique generally accepted system at the higher taxonomic
levels and there are frequent name changes at every level, from
species upwards. Efforts among researchers are now underway to
establish and encourage usage of a unified and more consistent
nomenclature . Fungal species can also have multiple scientific
names depending on their life cycle and mode (sexual or asexual) of
reproduction. Web sites such as
Index Fungorum and ITIS list current
names of fungal species (with cross-references to older synonyms).
The 2007 classification of Kingdom Fungi is the result of a
large-scale collaborative research effort involving dozens of
mycologists and other scientists working on fungal taxonomy. It
recognizes seven phyla , two of which—the
Ascomycota and the
Basidiomycota—are contained within a branch representing subkingdom
Dikarya, the most species rich and familiar group, including all the
mushrooms, most food-spoilage molds, most plant pathogenic fungi, and
the beer, wine, and bread yeasts. The accompanying cladogram depicts
the major fungal taxa and their relationship to opisthokont and
unikont organisms, based on the work of Philippe Silar and "The
Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for
Basic and Applied Research".
List of fungal orders Main groups of fungi.
The major phyla (sometimes called divisions) of fungi have been
classified mainly on the basis of characteristics of their sexual
reproductive structures. Currently, seven phyla are proposed:
Microsporidia, Chytridiomycota, Blastocladiomycota,
Neocallimastigomycota, Glomeromycota, Ascomycota, and Basidiomycota.
Phylogenetic analysis has demonstrated that the
unicellular parasites of animals and protists, are fairly recent and
highly derived endobiotic fungi (living within the tissue of another
species). One 2006 study concludes that the
Microsporidia are a
sister group to the true fungi; that is, they are each other's closest
evolutionary relative. Hibbett and colleagues suggest that this
analysis does not clash with their classification of the Fungi, and
Microsporidia are elevated to phylum status, it is
acknowledged that further analysis is required to clarify evolutionary
relationships within this group.
Chytridiomycota are commonly known as chytrids. These fungi are
Chytrids and their close relatives
Blastocladiomycota (below) are the only
fungi with active motility, producing zoospores that are capable of
active movement through aqueous phases with a single flagellum ,
leading early taxonomists to classify them as protists . Molecular
phylogenies , inferred from rRNA sequences in ribosomes , suggest that
Chytrids are a basal group divergent from the other fungal phyla,
consisting of four major clades with suggestive evidence for paraphyly
or possibly polyphyly .
Blastocladiomycota were previously considered a taxonomic clade
within the Chytridiomycota. Recent molecular data and ultrastructural
characteristics, however, place the
Blastocladiomycota as a sister
clade to the Zygomycota, Glomeromycota, and
Basidiomycota). The blastocladiomycetes are saprotrophs , feeding on
decomposing organic matter, and they are parasites of all eukaryotic
groups. Unlike their close relatives, the chytrids, most of which
exhibit zygotic meiosis , the blastocladiomycetes undergo sporic
Neocallimastigomycota were earlier placed in the phylum
Chytridomycota. Members of this small phylum are anaerobic organisms ,
living in the digestive system of larger herbivorous mammals and in
other terrestrial and aquatic environments enriched in cellulose
(e.g., domestic waste landfill sites). They lack mitochondria but
contain hydrogenosomes of mitochondrial origin. As in the related
chrytrids, neocallimastigomycetes form zoospores that are posteriorly
uniflagellate or polyflagellate.
Arbuscular mycorrhiza seen
Flax root cortical cells containing paired
arbuscules. Diagram of an apothecium (the typical cup-like
reproductive structure of Ascomycetes) showing sterile tissues as well
as developing and mature asci.
Members of the
Glomeromycota form arbuscular mycorrhizae , a form of
mutualist symbiosis wherein fungal hyphae invade plant root cells and
both species benefit from the resulting increased supply of nutrients.
Glomeromycota species reproduce asexually. The symbiotic
association between the
Glomeromycota and plants is ancient, with
evidence dating to 400 million years ago. Formerly part of the
Zygomycota (commonly known as 'sugar' and 'pin' molds), the
Glomeromycota were elevated to phylum status in 2001 and now replace
the older phylum Zygomycota. Fungi that were placed in the Zygomycota
are now being reassigned to the Glomeromycota, or the subphyla
Kickxellomycotina , the
Zoopagomycotina and the
Entomophthoromycotina . Some well-known
examples of fungi formerly in the
Zygomycota include black bread mold
Rhizopus stolonifer ), and
Pilobolus species, capable of ejecting
spores several meters through the air. Medically relevant genera
Rhizomucor , and
Ascomycota , commonly known as sac fungi or ascomycetes,
constitute the largest taxonomic group within the Eumycota. These
fungi form meiotic spores called ascospores , which are enclosed in a
special sac-like structure called an ascus . This phylum includes
morels , a few mushrooms and truffles , unicellular yeasts (e.g., of
Pichia , and Candida ), and
many filamentous fungi living as saprotrophs, parasites, and
mutualistic symbionts (e.g. lichens). Prominent and important genera
of filamentous ascomycetes include
Fusarium , and
Claviceps . Many ascomycete species have only been
observed undergoing asexual reproduction (called anamorphic species),
but analysis of molecular data has often been able to identify their
closest teleomorphs in the Ascomycota. Because the products of
meiosis are retained within the sac-like ascus, ascomycetes have been
used for elucidating principles of genetics and heredity (e.g.,
Neurospora crassa ).
Members of the
Basidiomycota , commonly known as the club fungi or
basidiomycetes, produce meiospores called basidiospores on club-like
stalks called basidia . Most common mushrooms belong to this group, as
well as rust and smut fungi , which are major pathogens of grains.
Other important basidiomycetes include the maize pathogen Ustilago
maydis , human commensal species of the genus
Malassezia , and the
opportunistic human pathogen,
Cryptococcus neoformans .
Because of similarities in morphology and lifestyle, the slime molds
(mycetozoans , plasmodiophorids , acrasids ,
labyrinthulids , now in
Excavata , Opisthokonta
Stramenopiles , respectively), water molds (oomycetes ) and
Stramenopiles ) were formerly classified in the
kingdom Fungi, in groups like
Phycomycetes . The slime molds were studied also as protozoans ,
leading to a ambiregnal , duplicated taxonomy.
Unlike true fungi, the cell walls of oomycetes contain cellulose and
lack chitin . Hyphochytrids have both chitin and cellulose. Slime
molds lack a cell wall during the assimilative phase (except
labyrinthulids, which have a wall of scales), and ingest nutrients by
ingestion (phagocytosis , except labyrinthulids) rather than
absorption (osmotrophy , as fungi, labyrinthulids, oomycetes and
hyphochytrids). Neither water molds nor slime molds are closely
related to the true fungi, and, therefore, taxonomists no longer group
them in the kingdom Fungi. Nonetheless, studies of the oomycetes and
myxomycetes are still often included in mycology textbooks and primary
Amoebidiales are opisthokont protists , previously
thought to be zygomycete fungi. Other groups now in Opisthokonta
Ichthyosporea ) were also at given time
classified as fungi. The genus
Blastocystis , now in
was originally classified as a yeast.
Ellobiopsis , now in
was considered a chytrid. The bacteria were also included in fungi in
some classifications, as the group Schizomycetes.
Rozellida clade, including the "ex-chytrid"
Rozella , is a
genetically disparate group known mostly from environmental DNA
sequences that is a sister group to fungi. Members of the group that
have been isolated lack the chitinous cell wall that is characteristic
The nucleariids , protists currently grouped in the Choanozoa
Opisthokonta ), may be the next sister group to the eumycete clade,
and as such could be included in an expanded fungal kingdom.
Actinobacteria ), a group with many filamentous
bacteria, were also long believed to be fungi.
A pin mold decomposing a peach
Although often inconspicuous, fungi occur in every environment on
Earth and play very important roles in most ecosystems . Along with
bacteria, fungi are the major decomposers in most terrestrial (and
some aquatic) ecosystems, and therefore play a critical role in
biogeochemical cycles and in many food webs . As decomposers, they
play an essential role in nutrient cycling , especially as saprotrophs
and symbionts , degrading organic matter to inorganic molecules, which
can then re-enter anabolic metabolic pathways in plants or other
Many fungi have important symbiotic relationships with organisms from
most if not all Kingdoms . These interactions can be mutualistic or
antagonistic in nature, or in the case of commensal fungi are of no
apparent benefit or detriment to the host.
Mycorrhizal symbiosis between plants and fungi is one of the most
well-known plant–fungus associations and is of significant
importance for plant growth and persistence in many ecosystems; over
90% of all plant species engage in mycorrhizal relationships with
fungi and are dependent upon this relationship for survival. The
dark filaments are hyphae of the endophytic fungus Neotyphodium
coenophialum in the intercellular spaces of tall fescue leaf sheath
The mycorrhizal symbiosis is ancient, dating to at least 400 million
years ago. It often increases the plant's uptake of inorganic
compounds, such as nitrate and phosphate from soils having low
concentrations of these key plant nutrients. The fungal partners may
also mediate plant-to-plant transfer of carbohydrates and other
nutrients. Such mycorrhizal communities are called "common mycorrhizal
networks". A special case of mycorrhiza is myco-heterotrophy ,
whereby the plant parasitizes the fungus, obtaining all of its
nutrients from its fungal symbiont. Some fungal species inhabit the
tissues inside roots, stems, and leaves, in which case they are called
endophytes. Similar to mycorrhiza, endophytic colonization by fungi
may benefit both symbionts; for example, endophytes of grasses impart
to their host increased resistance to herbivores and other
environmental stresses and receive food and shelter from the plant in
Algae And Cyanobacteria
Lobaria pulmonaria , a symbiosis of fungal, algal ,
and cyanobacterial species
Lichens are a symbiotic relationship between fungi and photosynthetic
algae or cyanobacteria . The photosynthetic partner in the
relationship is referred to in lichen terminology as a "photobiont".
The fungal part of the relationship is composed mostly of various
species of ascomycetes and a few basidiomycetes .
Lichens occur in
every ecosystem on all continents, play a key role in soil formation
and the initiation of biological succession , and are prominent in
some extreme environments, including polar , alpine , and semiarid
desert regions. They are able to grow on inhospitable surfaces,
including bare soil, rocks, tree bark , wood, shells, barnacles and
leaves. As in mycorrhizas , the photobiont provides sugars and other
carbohydrates via photosynthesis to the fungus, while the fungus
provides minerals and water to the photobiont. The functions of both
symbiotic organisms are so closely intertwined that they function
almost as a single organism; in most cases the resulting organism
differs greatly from the individual components. Lichenization is a
common mode of nutrition for fungi; around 20% of fungi—between
17,500 and 20,000 described species—are lichenized. Characteristics
common to most lichens include obtaining organic carbon by
photosynthesis, slow growth, small size, long life, long-lasting
(seasonal) vegetative reproductive structures, mineral nutrition
obtained largely from airborne sources, and greater tolerance of
desiccation than most other photosynthetic organisms in the same
Many insects also engage in mutualistic relationships with fungi.
Several groups of ants cultivate fungi in the order
their primary food source, while ambrosia beetles cultivate various
species of fungi in the bark of trees that they infest. Likewise,
females of several wood wasp species (genus
Sirex ) inject their eggs
together with spores of the wood-rotting fungus Amylostereum areolatum
into the sapwood of pine trees; the growth of the fungus provides
ideal nutritional conditions for the development of the wasp larvae.
At least one species of stingless bee has a relationship with a fungus
in the genus
Monascus , where the larvae consume and depend on fungus
transferred from old to new nests.
Termites on the African savannah
are also known to cultivate fungi, and yeasts of the genera Candida
and Lachancea inhabit the gut of a wide range of insects, including
neuropterans , beetles , and cockroaches ; it is not known whether
these fungi benefit their hosts. Fungi ingrowing dead wood are
essential for xylophagous insects (e.g. woodboring beetles ). They
deliver nutrients needed by xylophages to nutritionally scarce dead
wood . Thanks to this nutritional enrichment the larvae of woodboring
insect is able to grow and develop to adulthood. The larvae of many
families of fungicolous flies, particularly those within the
Sciaroidea such as the
Mycetophilidae and some
Keroplatidae feed on fungal fruiting bodies and sterile mycorrhizae .
As Pathogens And Parasites
The plant pathogen Aecidium magellanicum causes calafate rust ,
seen here on a
Berberis shrub in Chile.
Many fungi are parasites on plants, animals (including humans), and
other fungi. Serious pathogens of many cultivated plants causing
extensive damage and losses to agriculture and forestry include the
rice blast fungus
Magnaporthe oryzae , tree pathogens such as
Ophiostoma ulmi and
Ophiostoma novo-ulmi causing
Dutch elm disease
Cryphonectria parasitica responsible for chestnut blight , and
plant pathogens in the genera
Alternaria , and
Cochliobolus . Some carnivorous fungi , like
Paecilomyces lilacinus ,
are predators of nematodes , which they capture using an array of
specialized structures such as constricting rings or adhesive nets.
Some fungi can cause serious diseases in humans, several of which may
be fatal if untreated. These include aspergillosis , candidiasis ,
coccidioidomycosis , cryptococcosis , histoplasmosis , mycetomas , and
paracoccidioidomycosis . Furthermore, persons with immuno-deficiencies
are particularly susceptible to disease by genera such as Aspergillus
, Candida , Cryptoccocus ,
Histoplasma , and
Pneumocystis . Other
fungi can attack eyes, nails, hair, and especially skin, the so-called
dermatophytic and keratinophilic fungi, and cause local infections
such as ringworm and athlete\'s foot . Fungal spores are also a cause
of allergies , and fungi from different taxonomic groups can evoke
As Targets Of Mycoparasites
The organisms which parasitize fungi are known as mycoparasitic
organisms. Certain species of the
Pythium genus, which are oomycetes ,
have potential as biocontrol agents against certain fungi. Fungi can
also act as mycoparasites or antagonists of other fungi, such as
Hypomyces chrysospermus , which grows on bolete mushrooms.
Fungi can become the target of infection by mycoviruses .
Ergotamine , a major mycotoxin produced by
which if ingested can cause gangrene , convulsions, and hallucinations
Many fungi produce biologically active compounds, several of which
are toxic to animals or plants and are therefore called mycotoxins .
Of particular relevance to humans are mycotoxins produced by molds
causing food spoilage, and poisonous mushrooms (see above).
Particularly infamous are the lethal amatoxins in some Amanita
mushrooms, and ergot alkaloids , which have a long history of causing
serious epidemics of ergotism (St Anthony's Fire) in people consuming
rye or related cereals contaminated with sclerotia of the ergot
Claviceps purpurea . Other notable mycotoxins include the
aflatoxins , which are insidious liver toxins and highly carcinogenic
metabolites produced by certain
Aspergillus species often growing in
or on grains and nuts consumed by humans, ochratoxins , patulin , and
T-2 mycotoxin ) and fumonisins , which have
significant impact on human food supplies or animal livestock .
Mycotoxins are secondary metabolites (or natural products ), and
research has established the existence of biochemical pathways solely
for the purpose of producing mycotoxins and other natural products in
Mycotoxins may provide fitness benefits in terms of
physiological adaptation, competition with other microbes and fungi,
and protection from consumption (fungivory ). Many fungal secondary
metabolites (or derivatives) are used medically, as described under
Human Use below.
Ustilago maydis is a pathogenic plant fungus that causes smut disease
in maize and teosinte .
Plants have evolved efficient defense systems
against pathogenic microbes such as U. maydis. A rapid defense
reaction after pathogen attack is the oxidative burst where the plant
produces reactive oxygen species at the site of the attempted
invasion. U. maydis can respond to the oxidative burst with an
oxidative stress response, regulated by the gene
YAP1 . The response
protects U. maydis from the host defense, and is necessary for the
pathogen’s virulence. Furthermore, U. maydis has a well-established
DNA repair system which acts during mitosis and
meiosis. The system may assist the pathogen in surviving DNA damage
arising from the host plant’s oxidative defensive response to
Cryptococcus neoformans is an encapsulated yeast that can live in
both plants and animals. C. neoformans usually infects the lungs,
where it is phagocytosed by alveolar macrophages . Some C. neoformans
can survive inside macrophages, which appears to be the basis for
latency , disseminated disease, and resistance to antifungal agents.
One mechanism by which C. neoformans survives the hostile macrophage
environment is by up-regulating the expression of genes involved in
the oxidative stress response. Another mechanism involves meiosis .
The majority of C. neoformans are mating "type a". Filaments of mating
"type a" ordinarily have haploid nuclei, but they can become diploid
(perhaps by endoduplication or by stimulated nuclear fusion) to form
blastospores . The diploid nuclei of blastospores can undergo meiosis,
including recombination, to form haploid basidiospores that can be
dispersed. This process is referred to as monokaryotic fruiting. this
process requires a gene called
DMC1 , which is a conserved homologue
of genes recA in bacteria and
RAD51 in eukaryotes, that mediates
homologous chromosome pairing during meiosis and repair of DNA
double-strand breaks. Thus, C. neoformans can undergo a meiosis,
monokaryotic fruiting, that promotes recombinational repair in the
oxidative, DNA damaging environment of the host macrophage, and the
repair capability may contribute to its virulence.
Saccharomyces cerevisiae cells shown with DIC microscopy
The human use of fungi for food preparation or preservation and other
purposes is extensive and has a long history.
Mushroom farming and
mushroom gathering are large industries in many countries. The study
of the historical uses and sociological impact of fungi is known as
ethnomycology . Because of the capacity of this group to produce an
enormous range of natural products with antimicrobial or other
biological activities, many species have long been used or are being
developed for industrial production of antibiotics , vitamins, and
anti-cancer and cholesterol-lowering drugs. More recently, methods
have been developed for genetic engineering of fungi, enabling
metabolic engineering of fungal species. For example, genetic
modification of yeast species —which are easy to grow at fast rates
in large fermentation vessels—has opened up ways of pharmaceutical
production that are potentially more efficient than production by the
original source organisms.
Medicinal fungi The mould
Penicillium chrysogenum was
the source of penicillin G .
Many species produce metabolites that are major sources of
pharmacologically active drugs. Particularly important are the
antibiotics, including the penicillins , a structurally related group
of β-lactam antibiotics that are synthesized from small peptides .
Although naturally occurring penicillins such as penicillin G
Penicillium chrysogenum ) have a relatively narrow
spectrum of biological activity, a wide range of other penicillins can
be produced by chemical modification of the natural penicillins.
Modern penicillins are semisynthetic compounds, obtained initially
from fermentation cultures, but then structurally altered for specific
desirable properties. Other antibiotics produced by fungi include:
ciclosporin , commonly used as an immunosuppressant during transplant
surgery ; and fusidic acid , used to help control infection from
methicillin-resistant Staphylococcus aureus bacteria. Widespread use
of antibiotics for the treatment of bacterial diseases, such as
tuberculosis , syphilis , leprosy , and others began in the early 20th
century and continues to date. In nature, antibiotics of fungal or
bacterial origin appear to play a dual role: at high concentrations
they act as chemical defense against competition with other
microorganisms in species-rich environments, such as the rhizosphere ,
and at low concentrations as quorum-sensing molecules for intra- or
interspecies signaling. Other drugs produced by fungi include
griseofulvin isolated from
Penicillium griseofulvum , used to treat
fungal infections, and statins (
HMG-CoA reductase inhibitors), used
to inhibit cholesterol synthesis . Examples of statins found in fungi
include mevastatin from
Penicillium citrinum and lovastatin from
Aspergillus terreus and the oyster mushroom . Fungi produce compounds
that inhibit viruses and cancer cells . Specific metabolites,
such as polysaccharide-K , ergotamine , and β-lactam antibiotics ,
are routinely used in clinical medicine. The shiitake mushroom is a
source of lentinan , a clinical drug approved for use in cancer
treatments in several countries, including
Japan . In
Japan , polysaccharide-K (brand name Krestin), a chemical derived from
Trametes versicolor , is an approved adjuvant for cancer therapy.
Traditional And Folk Medicine
The medicinal fungi
Ganoderma lucidum (left) and
Ophiocordyceps sinensis (right)
Certain mushrooms enjoy usage as therapeutics in folk medicines ,
Traditional Chinese medicine . Notable medicinal mushrooms
with a well-documented history of use include
Agaricus subrufescens ,
Ganoderma lucidum , and
Ophiocordyceps sinensis .
Baker\'s yeast or
Saccharomyces cerevisiae , a unicellular fungus, is
used to make bread and other wheat-based products, such as pizza dough
and dumplings .
Yeast species of the genus
Saccharomyces are also
used to produce alcoholic beverages through fermentation.
Aspergillus oryzae ) is an essential ingredient in brewing Shoyu
(soy sauce ) and sake , and the preparation of miso , while Rhizopus
species are used for making tempeh . Several of these fungi are
domesticated species that were bred or selected according to their
capacity to ferment food without producing harmful mycotoxins (see
below), which are produced by very closely related Aspergilli . Quorn
, a meat substitute , is made from
Fusarium venenatum .
A selection of edible mushrooms eaten in Asia
Edible mushrooms include commercially raised and wild-harvested
Agaricus bisporus , sold as button mushrooms when small or
Portobello mushrooms when larger, is the most widely cultivated
species in the West, used in salads, soups, and many other dishes.
Many Asian fungi are commercially grown and have increased in
popularity in the West. They are often available fresh in grocery
stores and markets, including straw mushrooms (
Volvariella volvacea ),
oyster mushrooms (
Pleurotus ostreatus ), shiitakes (Lentinula edodes
), and enokitake (
Stilton cheese veined with
Many other mushroom species are harvested from the wild for personal
consumption or commercial sale. Milk mushrooms , morels , chanterelles
, truffles , black trumpets , and porcini mushrooms (
Boletus edulis )
(also known as king boletes) demand a high price on the market. They
are often used in gourmet dishes.
Certain types of cheeses require inoculation of milk curds with
fungal species that impart a unique flavor and texture to the cheese.
Examples include the blue color in cheeses such as Stilton or
Roquefort , which are made by inoculation with
. Molds used in cheese production are non-toxic and are thus safe for
human consumption; however, mycotoxins (e.g., aflatoxins, roquefortine
C , patulin, or others) may accumulate because of growth of other
fungi during cheese ripening or storage.
Amanita phalloides accounts for the majority of fatal mushroom
poisonings worldwide. It sometimes lacks the greenish color seen here.
Many mushroom species are poisonous to humans, with toxicities
ranging from slight digestive problems or allergic reactions as well
as hallucinations to severe organ failures and death. Genera with
mushrooms containing deadly toxins include
Lepiota , and, the most infamous,
Amanita . The latter genus includes
the destroying angel (A. virosa ) and the death cap (A. phalloides ),
the most common cause of deadly mushroom poisoning. The false morel
Gyromitra esculenta ) is occasionally considered a delicacy when
cooked, yet can be highly toxic when eaten raw. Tricholoma equestre
was considered edible until it was implicated in serious poisonings
causing rhabdomyolysis . Fly agaric mushrooms (
Amanita muscaria) also
cause occasional non-fatal poisonings, mostly as a result of ingestion
for its hallucinogenic properties. Historically, fly agaric was used
by different peoples in
Europe and Asia and its present usage for
religious or shamanic purposes is reported from some ethnic groups
such as the Koryak people of north-eastern
As it is difficult to accurately identify a safe mushroom without
proper training and knowledge, it is often advised to assume that a
wild mushroom is poisonous and not to consume it.
Grasshoppers killed by
In agriculture, fungi may be useful if they actively compete for
nutrients and space with pathogenic microorganisms such as bacteria or
other fungi via the competitive exclusion principle , or if they are
parasites of these pathogens. For example, certain species may be used
to eliminate or suppress the growth of harmful plant pathogens, such
as insects, mites , weeds , nematodes , and other fungi that cause
diseases of important crop plants. This has generated strong interest
in practical applications that use these fungi in the biological
control of these agricultural pests.
Entomopathogenic fungi can be
used as biopesticides , as they actively kill insects. Examples that
have been used as biological insecticides are
Beauveria bassiana ,
Paecilomyces (Isaria) spp, and
Lecanicillium lecanii .
Endophytic fungi of grasses of the genus
Neotyphodium , such as N. coenophialum , produce alkaloids that are
toxic to a range of invertebrate and vertebrate herbivores . These
alkaloids protect grass plants from herbivory , but several endophyte
alkaloids can poison grazing animals, such as cattle and sheep.
Infecting cultivars of pasture or forage grasses with Neotyphodium
endophytes is one approach being used in grass breeding programs; the
fungal strains are selected for producing only alkaloids that increase
resistance to herbivores such as insects, while being non-toxic to
Certain fungi, in particular "white rot" fungi, can degrade
insecticides , herbicides , pentachlorophenol , creosote , coal tars ,
and heavy fuels and turn them into carbon dioxide , water, and basic
elements. Fungi have been shown to biomineralize uranium oxides ,
suggesting they may have application in the bioremediation of
radioactively polluted sites.
Several pivotal discoveries in biology were made by researchers using
fungi as model organisms , that is, fungi that grow and sexually
reproduce rapidly in the laboratory. For example, the one gene-one
enzyme hypothesis was formulated by scientists using the bread mold
Neurospora crassa to test their biochemical theories. Other important
model fungi are
Aspergillus nidulans and the yeasts Saccharomyces
Schizosaccharomyces pombe , each of which with a long
history of use to investigate issues in eukaryotic cell biology and
genetics , such as cell cycle regulation, chromatin structure, and
gene regulation . Other fungal models have more recently emerged that
address specific biological questions relevant to medicine , plant
pathology , and industrial uses; examples include
Candida albicans , a
dimorphic, opportunistic human pathogen,
Magnaporthe grisea , a plant
Pichia pastoris , a yeast widely used for eukaryotic
protein production .
Fungi are used extensively to produce industrial chemicals like
citric , gluconic , lactic , and malic acids, and industrial enzymes,
such as lipases used in biological detergents , cellulases used in
making cellulosic ethanol and stonewashed jeans , and amylases ,
invertases , proteases and xylanases . Several species, most notably
Psilocybin mushrooms (colloquially known as magic mushrooms), are
ingested for their psychedelic properties, both recreationally and
* Fungi portal
Outline of fungi
Conservation of fungi
* ^ Moore RT (1980). "Taxonomic proposals for the classification of
marine yeasts and other yeast-like fungi including the smuts".
Botanica Marina. 23: 361–373.
* ^ The classification system presented here is based on the 2007
phylogenetic study by Hibbett et al.
* ^ /ˈfʌndʒaɪ/ ( listen ) or /ˈfʌŋɡaɪ/ ( listen )
* ^ "Fungus". Oxford Dictionaries. Retrieved 26 February 2011.
* ^ https://www.nature.com/articles/nmicrobiol2017120
* ^ Simpson DP (1979). Cassell's
Latin Dictionary (5 ed.). London,
UK: Cassell Ltd. p. 883. ISBN 0-304-52257-0 .
* ^ A B Ainsworth, p. 2.
* ^ Mitzka W, ed. (1960). Etymologisches Wörterbuch der deutschen
Sprache. Berlin: Walter de Gruyter.
* ^ Alexopoulos et al., p. 1.
* ^ "LIAS Glossary". Retrieved 14 August 2013.
* ^ Bruns T (October 2006). "
Evolutionary biology: a kingdom
revised". Nature. 443 (7113): 758–61.
doi :10.1038/443758a . PMID 17051197 .
* ^ Baldauf SL, Palmer JD (December 1993). "Animals and fungi are
each other\'s closest relatives: congruent evidence from multiple
proteins" . Proceedings of the National Academy of Sciences of the
United States of America. 90 (24): 11558–62. Bibcode
:1993PNAS...9011558B. doi :10.1073/pnas.90.24.11558 . PMC 48023 .
PMID 8265589 .
* ^ Deacon, p. 4.
* ^ A B Deacon, pp. 128–129.
* ^ Alexopoulos et al., pp. 28–33.
* ^ Alexopoulos et al., pp. 31–32.
* ^ Shoji JY, Arioka M, Kitamoto K (2006). "Possible involvement of
pleiomorphic vacuolar networks in nutrient recycling in filamentous
fungi". Autophagy. 2 (3): 226–7. doi :10.4161/auto.2695 . PMID
* ^ Deacon, p. 58.
* ^ Zabriskie TM, Jackson MD (February 2000). "Lysine biosynthesis
and metabolism in fungi". Natural Product Reports. 17 (1): 85–97.
doi :10.1039/a801345d . PMID 10714900 .
* ^ Xu H, Andi B, Qian J, West AH, Cook PF (2006). "The
alpha-aminoadipate pathway for lysine biosynthesis in fungi". Cell
Biochemistry and Biophysics. 46 (1): 43–64. doi :10.1385/CBB:46:1:43
. PMID 16943623 .
* ^ Alexopoulos et al., pp. 27–28.
* ^ Alexopoulos et al., p. 685.
* ^ A B C Alexopoulos et al., p. 30.
* ^ Desjardin DE, Perry BA, Lodge DJ, Stevani CV, Nagasawa E
(2010). "Luminescent Mycena: new and noteworthy species". Mycologia.
102 (2): 459–77. doi :10.3852/09-197 . PMID 20361513 .
* ^ Alexopoulos et al., pp. 32–33.
* ^ Bowman SM, Free SJ (August 2006). "The structure and synthesis
of the fungal cell wall". BioEssays. 28 (8): 799–808. doi
:10.1002/bies.20441 . PMID 16927300 .
* ^ Alexopoulos et al., p. 33.
* ^ Mihail JD, Bruhn JN (November 2005). "Foraging behaviour of
Armillaria rhizomorph systems". Mycological Research. 109 (Pt 11):
1195–207. doi :10.1017/S0953756205003606 . PMID 16279413 .
* ^ A B C D Keller NP, Turner G, Bennett JW (December 2005).
"Fungal secondary metabolism - from biochemistry to genomics". Nature
Reviews. Microbiology. 3 (12): 937–47. doi :10.1038/nrmicro1286 .
PMID 16322742 .
* ^ Wu S, Schalk M, Clark A, Miles RB, Coates R, Chappell J
(November 2006). "Redirection of cytosolic or plastidic isoprenoid
precursors elevates terpene production in plants". Nature
Biotechnology. 24 (11): 1441–7. doi :10.1038/nbt1251 . PMID 17057703
* ^ Tudzynski B (March 2005). "Gibberellin biosynthesis in fungi:
genes, enzymes, evolution, and impact on biotechnology". Applied
Microbiology and Biotechnology. 66 (6): 597–611. doi
:10.1007/s00253-004-1805-1 . PMID 15578178 .
* ^ Vaupotic T, Veranic P, Jenoe P, Plemenitas A (June 2008).
"Mitochondrial mediation of environmental osmolytes discrimination
during osmoadaptation in the extremely halotolerant black yeast
Hortaea werneckii". Fungal
Genetics and Biology. 45 (6): 994–1007.
doi :10.1016/j.fgb.2008.01.006 . PMID 18343697 .
* ^ A B Dadachova E, Bryan RA, Huang X, Moadel T, Schweitzer AD,
Aisen P, Nosanchuk JD, Casadevall A (2007). "Ionizing radiation
changes the electronic properties of melanin and enhances the growth
of melanized fungi" . PLoS One. 2 (5): e457. Bibcode
:2007PLoSO...2..457D. doi :10.1371/journal.pone.0000457 . PMC 1866175
. PMID 17520016 .
* ^ Raghukumar C, Raghukumar S (1998). "Barotolerance of fungi
isolated from deep-sea sediments of the Indian Ocean". Aquatic
Microbial Ecology. 15 (2): 153–163. doi :10.3354/ame015153 .
* ^ Sancho LG, de la Torre R, Horneck G, Ascaso C, de Los Rios A,
Pintado A, Wierzchos J, Schuster M (June 2007). "
Lichens survive in
space: results from the 2005 LICHENS experiment". Astrobiology. 7 (3):
Bibcode :2007AsBio...7..443S. doi :10.1089/ast.2006.0046 .
PMID 17630840 .
* ^ Brem FM, Lips KR (September 2008). "Batrachochytrium
dendrobatidis infection patterns among Panamanian amphibian species,
habitats and elevations during epizootic and enzootic stages".
Diseases of Aquatic Organisms. 81 (3): 189–202. doi
:10.3354/dao01960 . PMID 18998584 .
* ^ Le Calvez T, Burgaud G, Mahé S, Barbier G, Vandenkoornhuyse P
(October 2009). "Fungal diversity in deep-sea hydrothermal ecosystems"
. Applied and Environmental Microbiology. 75 (20): 6415–21. doi
:10.1128/AEM.00653-09 . PMC 2765129 . PMID 19633124 .
* ^ A B Mueller GM, Schmit JP (2006). "Fungal biodiversity: what do
we know? What can we predict?".
Biodiversity and Conservation. 16:
1–5. doi :10.1007/s10531-006-9117-7 .
* ^ Hawksworth DL, Lücking R (July 2017). "Fungal Diversity
Revisited: 2.2 to 3.8 Million Species".
Microbiology Spectrum. 5 (4):
79. doi :10.1128/microbiolspec.FUNK-0052-2016 . ISBN 9781555819576 .
PMID 28752818 .
* ^ A B Kirk et al., p. 489.
* ^ A B C D E F G H Hibbett DS, Binder M, Bischoff JF, Blackwell M,
Cannon PF, Eriksson OE, et al. (May 2007). "A higher-level
phylogenetic classification of the Fungi" (PDF). Mycological Research.
111 (Pt 5): 509–47. doi :10.1016/j.mycres.2007.03.004 . PMID
17572334 . Archived from the original (PDF) on 26 March 2009.
* ^ According to one 2001 estimate, some 10,000 fungal diseases are
known. Struck C (2006). "Infection strategies of plant parasitic
fungi". In Cooke BM, Jones DG, Kaye B. The Epidemiology of Plant
Diseases. Berlin, Germany: Springer. p. 117. ISBN 1-4020-4580-8 .
* ^ Peintner U, Pöder R, Pümpel T (1998). "The Iceman's fungi".
Mycological Research. 102 (10): 1153–1162. doi
* ^ Ainsworth, p. 1.
* ^ Alexopoulos et al., pp. 1–2.
* ^ Ainsworth, p. 18.
* ^ Hawksworth DL (September 2006). "Pandora's mycological box:
molecular sequences vs. morphology in understanding fungal
relationships and biodiversity". Revista Iberoamericana de MicologíA.
23 (3): 127–33. doi :10.1016/S1130-1406(06)70031-6 . PMID 17196017 .
* ^ Harris SD (2008). "Branching of fungal hyphae: regulation,
mechanisms and comparison with other branching systems". Mycologia.
100 (6): 823–32. doi :10.3852/08-177 . PMID 19202837 .
* ^ Deacon, p. 51.
* ^ Deacon, p. 57.
* ^ Chang S-T, Miles PG (2004). Mushrooms: Cultivation, Nutritional
Value, Medicinal Effect and Environmental Impact. Boca Raton, Florida:
CRC Press. ISBN 0-8493-1043-1 .
* ^ Parniske M (October 2008). "Arbuscular mycorrhiza: the mother
of plant root endosymbioses".
Nature Reviews. Microbiology. 6 (10):
763–75. doi :10.1038/nrmicro1987 . PMID 18794914 .
* ^ Steenkamp ET, Wright J, Baldauf SL (January 2006). "The
protistan origins of animals and fungi". Molecular
Evolution. 23 (1): 93–106. doi :10.1093/molbev/msj011 . PMID
* ^ Stevens DA, Ichinomiya M, Koshi Y, Horiuchi H (September 2006).
"Escape of Candida from caspofungin inhibition at concentrations above
the MIC (paradoxical effect) accomplished by increased cell wall
chitin; evidence for beta-1,6-glucan synthesis inhibition by
Antimicrobial Agents and Chemotherapy. 50 (9):
3160–1. doi :10.1128/AAC.00563-06 . PMC 1563524 . PMID 16940118 .
* ^ Hanson, pp. 127–141.
* ^ Ferguson BA, Dreisbach TA, Parks CG, Filip GM, Schmitt CL
(2003). "Coarse-scale population structure of pathogenic Armillaria
species in a mixed-conifer forest in the Blue Mountains of northeast
Canadian Journal of Forest Research . 33 (4): 612–623. doi
* ^ Alexopoulos et al., pp. 204–205.
Moss ST (1986). The
Biology of Marine Fungi. Cambridge, UK:
Cambridge University Press
Cambridge University Press . p. 76. ISBN 0-521-30899-2 .
* ^ Peñalva MA, Arst HN (September 2002). "Regulation of gene
expression by ambient pH in filamentous fungi and yeasts" .
Microbiology and Molecular
Biology Reviews. 66 (3): 426–46, table of
contents. doi :10.1128/MMBR.66.3.426-446.2002 . PMC 120796 . PMID
* ^ A B Howard RJ, Ferrari MA, Roach DH, Money NP (December 1991).
"Penetration of hard substrates by a fungus employing enormous turgor
pressures" . Proceedings of the National Academy of Sciences of the
United States of America. 88 (24): 11281–4. Bibcode
:1991PNAS...8811281H. doi :10.1073/pnas.88.24.11281 . PMC 53118 .
PMID 1837147 .
* ^ Money NP (1998). "Mechanics of invasive fungal growth and the
significance of turgor in plant infection". Molecular
Host-Specific Toxins in
Plant Disease: Proceedings of the 3rd Tottori
International Symposium on Host-Specific Toxins, Daisen, Tottori,
Japan, August 24–29, 1997. Netherlands: Kluwer Academic Publishers.
pp. 261–271. ISBN 0-7923-4981-4 .
* ^ Wang ZY, Jenkinson JM, Holcombe LJ, Soanes DM, Veneault-Fourrey
C, Bhambra GK, Talbot NJ (April 2005). "The molecular biology of
appressorium turgor generation by the rice blast fungus Magnaporthe
grisea". Biochemical Society Transactions. 33 (Pt 2): 384–8. doi
:10.1042/BST0330384 . PMID 15787612 .
* ^ Pereira JL, Noronha EF, Miller RN, Franco OL (June 2007).
"Novel insights in the use of hydrolytic enzymes secreted by fungi
with biotechnological potential". Letters in Applied Microbiology. 44
(6): 573–81. doi :10.1111/j.1472-765X.2007.02151.x . PMID 17576216 .
* ^ Schaller M, Borelli C, Korting HC, Hube B (November 2005).
"Hydrolytic enzymes as virulence factors of Candida albicans".
Mycoses. 48 (6): 365–77. doi :10.1111/j.1439-0507.2005.01165.x .
PMID 16262871 .
* ^ Farrar JF (October 1985). "Carbohydrate metabolism in
biotrophic plant pathogens". Microbiological Sciences. 2 (10):
314–7. PMID 3939987 .
* ^ Fischer R, Zekert N, Takeshita N (May 2008). "Polarized growth
in fungi--interplay between the cytoskeleton, positional markers and
membrane domains". Molecular Microbiology. 68 (4): 813–26. doi
:10.1111/j.1365-2958.2008.06193.x . PMID 18399939 .
* ^ Christensen MJ, Bennett RJ, Ansari HA, Koga H, Johnson RD,
Bryan GT, Simpson WR, Koolaard JP, Nickless EM, Voisey CR (February
2008). "Epichloë endophytes grow by intercalary hyphal extension in
elongating grass leaves". Fungal
Genetics and Biology. 45 (2):
84–93. doi :10.1016/j.fgb.2007.07.013 . PMID 17919950 .
* ^ Money NP (October 2002). "
Mushroom stem cells". BioEssays. 24
(10): 949–52. doi :10.1002/bies.10160 . PMID 12325127 .
* ^ Willensdorfer M (February 2009). "On the evolution of
differentiated multicellularity". Evolution; International Journal of
Organic Evolution. 63 (2): 306–23. doi
:10.1111/j.1558-5646.2008.00541.x . PMID 19154376 .
* ^ Daniels KJ, Srikantha T, Lockhart SR, Pujol C, Soll DR (May
2006). "Opaque cells signal white cells to form biofilms in Candida
albicans" . The EMBO Journal. 25 (10): 2240–52. doi
:10.1038/sj.emboj.7601099 . PMC 1462973 . PMID 16628217 .
* ^ Marzluf GA (September 1981). "Regulation of nitrogen metabolism
and gene expression in fungi" . Microbiological Reviews. 45 (3):
437–61. PMC 281519 . PMID 6117784 .
* ^ Hynes MJ (1994). "Regulatory circuits of the amdS gene of
Aspergillus nidulans". Antonie Van Leeuwenhoek. 65 (3): 179–82. doi
:10.1007/BF00871944 . PMID 7847883 .
* ^ Dadachova E, Casadevall A (December 2008). "Ionizing radiation:
how fungi cope, adapt, and exploit with the help of melanin" . Current
Opinion in Microbiology. 11 (6): 525–31. doi
:10.1016/j.mib.2008.09.013 . PMC 2677413 . PMID 18848901 .
* ^ Alexopoulos et al., pp. 48–56.
* ^ Kirk et al., p. 633.
* ^ Heitman J (September 2006). "
Sexual reproduction and the
evolution of microbial pathogens". Current Biology. 16 (17):
R711–25. doi :10.1016/j.cub.2006.07.064 . PMID 16950098 .
* ^ Alcamo IE, Pommerville J (2004). Alcamo's Fundamentals of
Microbiology. Boston, Massachusetts: Jones and Bartlett. p. 590. ISBN
* ^ A B Redecker D, Raab P (2006). "Phylogeny of the glomeromycota
(arbuscular mycorrhizal fungi): recent developments and new gene
markers". Mycologia. 98 (6): 885–95. doi :10.3852/mycologia.98.6.885
. PMID 17486965 .
* ^ Guarro J, Stchigel AM (July 1999). "Developments in fungal
Microbiology Reviews. 12 (3): 454–500. PMC
100249 . PMID 10398676 .
* ^ A B Taylor JW, Jacobson DJ, Kroken S, Kasuga T, Geiser DM,
Hibbett DS, Fisher MC (October 2000). "
recognition and species concepts in fungi". Fungal
Biology. 31 (1): 21–32. doi :10.1006/fgbi.2000.1228 . PMID 11118132
* ^ Metzenberg RL, Glass NL (February 1990). "
Mating type and
mating strategies in Neurospora". BioEssays. 12 (2): 53–9. doi
:10.1002/bies.950120202 . PMID 2140508 .
* ^ Jennings and Lysek, pp. 107–114.
* ^ Deacon, p. 31.
* ^ Alexopoulos et al., pp. 492–493.
* ^ Jennings and Lysek, p. 142.
* ^ Deacon, pp. 21–24.
* ^ Linder MB, Szilvay GR, Nakari-Setälä T, Penttilä ME
(November 2005). "Hydrophobins: the protein-amphiphiles of filamentous
Microbiology Reviews. 29 (5): 877–96. doi
:10.1016/j.femsre.2005.01.004 . PMID 16219510 .
* ^ Trail F (November 2007). "Fungal cannons: explosive spore
discharge in the Ascomycota". FEMS
Microbiology Letters. 276 (1):
12–8. doi :10.1111/j.1574-6968.2007.00900.x . PMID 17784861 .
* ^ Pringle A, Patek SN, Fischer M, Stolze J, Money NP (2005). "The
captured launch of a ballistospore". Mycologia. 97 (4): 866–71. doi
:10.3852/mycologia.97.4.866 . PMID 16457355 .
* ^ Kirk et al., p. 495.
* ^ Brodie HJ (1975). The Bird's Nest Fungi. Toronto, Ontario:
University of Toronto Press. p. 80. ISBN 0-8020-5307-6 .
* ^ Alexopoulos et al., p. 545.
* ^ Jennings and Lysek, pp. 114–115.
* ^ Furlaneto MC, Pizzirani-Kleiner AA (January 1992).
"Intraspecific hybridisation of Trichoderma pseudokoningii by
anastomosis and by protoplast fusion". FEMS
Microbiology Letters. 69
(2): 191–5. doi :10.1111/j.1574-6968.1992.tb05150.x . PMID 1537549 .
* ^ Schardl CL, Craven KD (November 2003). "Interspecific
hybridization in plant-associated fungi and oomycetes: a review".
Molecular Ecology. 12 (11): 2861–73. doi
:10.1046/j.1365-294X.2003.01965.x . PMID 14629368 .
* ^ Donoghue MJ, Cracraft J (2004). Assembling the Tree of Life.
Oxford (Oxfordshire), UK: Oxford University Press. p. 187. ISBN
* ^ Taylor and Taylor, p. 19.
* ^ Taylor and Taylor, pp. 7–12.
* ^ Bengtson, Stefan; Rasmussen, Birger; Ivarsson, Magnus; Muhling,
Janet; Broman, Curt; Marone, Federica; Stampanoni, Marco; Bekker,
Andrey (24 April 2017). "Fungus-like mycelial fossils in
2.4-billion-year-old vesicular basalt".
Ecology & Evolution. 1
(6): 0141. doi :10.1038/s41559-017-0141 . ISSN 2397-334X .
* ^ Lücking R, Huhndorf S, Pfister DH, Plata ER, Lumbsch HT
(2009). "Fungi evolved right on track". Mycologia. 101 (6): 810–22.
doi :10.3852/09-016 . PMID 19927746 .
* ^ A B C James TY, Kauff F, Schoch CL, Matheny PB, Hofstetter V,
Cox CJ, et al. (October 2006). "Reconstructing the early evolution of
Fungi using a six-gene phylogeny". Nature. 443 (7113): 818–22.
Bibcode :2006Natur.443..818J. doi :10.1038/nature05110 . PMID 17051209
* ^ Taylor and Taylor, pp. 84–94 and 106–107.
* ^ Schoch CL, Sung GH, López-Giráldez F, Townsend JP,
Miadlikowska J, Hofstetter V, et al. (April 2009). "The Ascomycota
tree of life: a phylum-wide phylogeny clarifies the origin and
evolution of fundamental reproductive and ecological traits".
Systematic Biology. 58 (2): 224–39. doi :10.1093/sysbio/syp020 .
PMID 20525580 .
* ^ A B Brundrett MC (2002). "Coevolution of roots and mycorrhizas
of land plants". New Phytologist. 154 (2): 275–304. doi
* ^ Redecker D, Kodner R, Graham LE (September 2000). "Glomalean
fungi from the Ordovician". Science. 289 (5486): 1920–1. Bibcode
:2000Sci...289.1920R. doi :10.1126/science.289.5486.1920 . PMID
* ^ Taylor TN, Taylor EL (1996). "The distribution and interactions
Paleozoic fungi". Review of Palaeobotany and Palynology. 95
(1–4): 83–94. doi :10.1016/S0034-6667(96)00029-2 .
* ^ Dotzler N, Walker C, Krings M, Hass H, Kerp H, Taylor TN,
Agerer R (2009). "Acaulosporoid glomeromycotan spores with a
germination shield from the 400-million-year-old Rhynie chert".
Mycological Progress. 8 (1): 9–18. doi :10.1007/s11557-008-0573-1 .
* ^ Taylor JW, Berbee ML (2006). "Dating divergences in the Fungal
Tree of Life: review and new analyses". Mycologia. 98 (6): 838–49.
doi :10.3852/mycologia.98.6.838 . PMID 17486961 .
* ^ Blackwell M, Vilgalys R, James TY, Taylor JW (2009). "Fungi.
Eumycota: mushrooms, sac fungi, yeast, molds, rusts, smuts, etc". Tree
Life Web Project. Retrieved 25 April 2009.
* ^ Yuan X, Xiao S, Taylor TN (May 2005). "Lichen-like symbiosis
600 million years ago". Science. 308 (5724): 1017–20. Bibcode
:2005Sci...308.1017Y. doi :10.1126/science.1111347 . PMID 15890881 .
* ^ Karatygin IV, Snigirevskaya NS, Vikulin SV (2009). "The most
ancient terrestrial lichen Winfrenatia reticulata: A new find and new
interpretation". Paleontological Journal. 43 (1): 107–114. doi
* ^ Taylor TN, Hass H, Kerp H, Krings M, Hanlin RT (2005).
"Perithecial ascomycetes from the 400 million year old Rhynie chert:
an example of ancestral polymorphism". Mycologia. 97 (1): 269–85.
doi :10.3852/mycologia.97.1.269 . PMID 16389979 .
* ^ Dennis RL (1970). "A Middle Pennsylvanian basidiomycete
mycelium with clamp connections". Mycologia. 62 (3): 578–584. doi
JSTOR 3757529 .
* ^ Hibbett DS, Grimaldi D, Donoghue MJ (1995). "Cretaceous
mushrooms in amber". Nature. 377 (6549): 487. Bibcode
:1995Natur.377..487H. doi :10.1038/377487a0 .
* ^ Hibbett DS, Grimaldi D, Donoghue MJ (1997). "Fossil mushrooms
from Miocene and
Cretaceous ambers and the evolution of
homobasidiomycetes". American Journal of Botany. 84 (7): 981–991.
doi :10.2307/2446289 .
JSTOR 2446289 .
* ^ Eshet Y, Rampino MR, Visscher H (1995). "Fungal event and
palynological record of ecological crisis and recovery across the
Triassic boundary". Geology. 23 (1): 967–970. Bibcode
:1995Geo....23..967E. doi :10.1130/0091-7613(1995)0232.3.CO;2 .
* ^ Foster CB, Stephenson MH, Marshall C, Logan GA, Greenwood PF
(2002). "A revision of Reduviasporonites Wilson 1962: description,
illustration, comparison and biological affinities". Palynology. 26
(1): 35–58. doi :10.2113/0260035 .
* ^ López-Gómez J, Taylor EL (2005). "Permian-
in Spain: a multidisciplinary approach". Palaeogeography,
Palaeoclimatology, Palaeoecology. 229 (1–2): 1–2. doi
* ^ Looy CV, Twitchett RJ, Dilcher DL, Van Konijnenburg-Van Cittert
JH, Visscher H (July 2001). "
Life in the end-
Permian dead zone" .
Proceedings of the National Academy of Sciences of the United States
of America. 98 (14): 7879–83.
Bibcode :2001PNAS...98.7879L. doi
:10.1073/pnas.131218098 . PMC 35436 . PMID 11427710 . See image 2
* ^ Ward PD, Botha J, Buick R, De Kock MO, Erwin DH, Garrison GH,
Kirschvink JL, Smith R (February 2005). "Abrupt and gradual extinction
Permian land vertebrates in the Karoo basin, South Africa".
Science. 307 (5710): 709–14.
Bibcode :2005Sci...307..709W. doi
:10.1126/science.1107068 . PMID 15661973 .
* ^ A B Shalchian-Tabrizi K, Minge MA, Espelund M, Orr R, Ruden T,
Jakobsen KS, Cavalier-Smith T (2008). "Multigene phylogeny of
choanozoa and the origin of animals" . PLoS One. 3 (5): e2098. Bibcode
:2008PLoSO...3.2098S. doi :10.1371/journal.pone.0002098 . PMC 2346548
. PMID 18461162 .
* ^ See Palaeos Fungi: Fungi for an introduction to fungal
taxonomy, including recent controversies. archive
* ^ Celio GJ, Padamsee M, Dentinger BT, Bauer R, McLaughlin DJ
(2006). "Assembling the Fungal Tree of Life: constructing the
structural and biochemical database". Mycologia. 98 (6): 850–9. doi
:10.3852/mycologia.98.6.850 . PMID 17486962 .
* ^ Silar P (2016). "Protistes Eucaryotes: Origine,
Biologie des Microbes Eucaryotes". HAL: 462. ISBN 978-2-9555841-0-1 .
* ^ Esser K (2014). The Mycota VII A: Systematics and Evolution
(2nd ed.). Springer. p. 461. ISBN 978-3-642-55317-2 .
* ^ Gill EE, Fast NM (June 2006). "Assessing the
microsporidia-fungi relationship: Combined phylogenetic analysis of
eight genes". Gene. 375: 103–9. doi :10.1016/j.gene.2006.02.023 .
PMID 16626896 .
* ^ Liu YJ, Hodson MC, Hall BD (2006). "Loss of the flagellum
happened only once in the fungal lineage: phylogenetic structure of
kingdom Fungi inferred from RNA polymerase II subunit genes" . BMC
Evolutionary Biology. 6: 74. doi :10.1186/1471-2148-6-74 . PMC 1599754
. PMID 17010206 .
* ^ James TY, Letcher PM, Longcore JE, Mozley-Standridge SE, Porter
D, Powell MJ, Griffith GW, Vilgalys R (2006). "A molecular phylogeny
of the flagellated fungi (Chytridiomycota) and description of a new
phylum (Blastocladiomycota)". Mycologia. 98 (6): 860–71. doi
:10.3852/mycologia.98.6.860 . PMID 17486963 .
* ^ Lockhart RJ, Van Dyke MI, Beadle IR, Humphreys P, McCarthy AJ
(August 2006). "Molecular biological detection of anaerobic gut fungi
(Neocallimastigales) from landfill sites" . Applied and Environmental
Microbiology. 72 (8): 5659–61. doi :10.1128/AEM.01057-06 . PMC
1538735 . PMID 16885325 .
* ^ A B Remy W, Taylor TN, Hass H, Kerp H (December 1994). "Four
hundred-million-year-old vesicular arbuscular mycorrhizae" .
Proceedings of the National Academy of Sciences of the United States
of America. 91 (25): 11841–3.
Bibcode :1994PNAS...9111841R. doi
:10.1073/pnas.91.25.11841 . PMC 45331 . PMID 11607500 .
* ^ Schüssler A, Schwarzott D, Walker C (2001). "A new fungal
phylum, the Glomeromycota: phylogeny and evolution". Mycological
Research. 105 (12): 1413–1421. doi :10.1017/S0953756201005196 .
* ^ Alexopoulos et al., p. 145.
* ^ For an example, see Samuels GJ (February 2006). "Trichoderma:
systematics, the sexual state, and ecology". Phytopathology. 96 (2):
195–206. doi :10.1094/PHYTO-96-0195 . PMID 18943925 .
* ^ Radford A, Parish JH (June 1997). "The genome and genes of
Neurospora crassa". Fungal
Genetics and Biology. 21 (3): 258–66. doi
:10.1006/fgbi.1997.0979 . PMID 9290240 .
* ^ Valverde ME, Paredes-López O, Pataky JK, Guevara-Lara F
(January 1995). "Huitlacoche (
Ustilago maydis) as a food
source--biology, composition, and production". Critical Reviews in
Food Science and Nutrition. 35 (3): 191–229. doi
:10.1080/10408399509527699 . PMID 7632354 .
* ^ Zisova LG (2009). "
Malassezia species and seborrheic
dermatitis". Folia Medica. 51 (1): 23–33. PMID 19437895 .
* ^ Perfect JR (June 2006). "Cryptococcus neoformans: the yeast
that likes it hot". FEMS
Yeast Research. 6 (4): 463–8. doi
:10.1111/j.1567-1364.2006.00051.x . PMID 16696642 .
* ^ Blackwell M, Spatafora JW (2004). "Fungi and their allies". In
Bills GF, Mueller GM, Foster MS.
Biodiversity of Fungi: Inventory and
Monitoring Methods. Amsterdam: Elsevier Academic Press. pp. 18–20.
ISBN 0-12-509551-1 .
* ^ Gadd GM (January 2007). "Geomycology: biogeochemical
transformations of rocks, minerals, metals and radionuclides by fungi,
bioweathering and bioremediation". Mycological Research. 111 (Pt 1):
3–49. doi :10.1016/j.mycres.2006.12.001 . PMID 17307120 .
* ^ A B Lindahl BD, Ihrmark K, Boberg J, Trumbore SE, Högberg P,
Stenlid J, Finlay RD (2007). "Spatial separation of litter
decomposition and mycorrhizal nitrogen uptake in a boreal forest". The
New Phytologist. 173 (3): 611–20. doi
:10.1111/j.1469-8137.2006.01936.x . PMID 17244056 .
* ^ Barea JM, Pozo MJ, Azcón R, Azcón-Aguilar C (July 2005).
"Microbial co-operation in the rhizosphere". Journal of Experimental
Botany. 56 (417): 1761–78. doi :10.1093/jxb/eri197 . PMID 15911555 .
* ^ A B Aanen DK (June 2006). "As you reap, so shall you sow:
coupling of harvesting and inoculating stabilizes the mutualism
between termites and fungi" .
Biology Letters. 2 (2): 209–12. doi
:10.1098/rsbl.2005.0424 . PMC 1618886 . PMID 17148364 .
* ^ Nikoh N, Fukatsu T (April 2000). "Interkingdom host jumping
underground: phylogenetic analysis of entomoparasitic fungi of the
genus cordyceps". Molecular
Biology and Evolution. 17 (4): 629–38.
doi :10.1093/oxfordjournals.molbev.a026341 . PMID 10742053 .
* ^ Perotto S, Bonfante P (December 1997). "Bacterial associations
with mycorrhizal fungi: close and distant friends in the rhizosphere".
Trends in Microbiology. 5 (12): 496–501. doi
:10.1016/S0966-842X(97)01154-2 . PMID 9447662 .
* ^ Arnold AE, Mejía LC, Kyllo D, Rojas EI, Maynard Z, Robbins N,
Herre EA (December 2003). "Fungal endophytes limit pathogen damage in
a tropical tree" . Proceedings of the National Academy of Sciences of
the United States of America. 100 (26): 15649–54. Bibcode
:2003PNAS..10015649A. doi :10.1073/pnas.2533483100 . PMC 307622 .
PMID 14671327 .
* ^ A B Paszkowski U (August 2006). "Mutualism and parasitism: the
yin and yang of plant symbioses". Current Opinion in
Plant Biology. 9
(4): 364–70. doi :10.1016/j.pbi.2006.05.008 . PMID 16713732 .
* ^ A B Hube B (August 2004). "From commensal to pathogen: stage-
and tissue-specific gene expression of Candida albicans". Current
Opinion in Microbiology. 7 (4): 336–41. doi
:10.1016/j.mib.2004.06.003 . PMID 15288621 .
* ^ Bonfante P (April 2003). "Plants, mycorrhizal fungi and
endobacteria: a dialog among cells and genomes". The Biological
Bulletin. 204 (2): 215–20. doi :10.2307/1543562 .
JSTOR 1543562 .
PMID 12700157 .
* ^ van der Heijden MG, Streitwolf-Engel R, Riedl R, Siegrist S,
Neudecker A, Ineichen K, Boller T, Wiemken A, Sanders IR (2006). "The
mycorrhizal contribution to plant productivity, plant nutrition and
soil structure in experimental grassland". The New Phytologist. 172
(4): 739–52. doi :10.1111/j.1469-8137.2006.01862.x . PMID 17096799 .
* ^ Selosse MA, Richard F, He X, Simard SW (November 2006).
"Mycorrhizal networks: des liaisons dangereuses?". Trends in
Evolution. 21 (11): 621–8. doi :10.1016/j.tree.2006.07.003 . PMID
* ^ Merckx V, Bidartondo MI, Hynson NA (December 2009).
"Myco-heterotrophy: when fungi host plants" . Annals of Botany. 104
(7): 1255–61. doi :10.1093/aob/mcp235 . PMC 2778383 . PMID
* ^ Schulz B, Boyle C (June 2005). "The endophytic continuum".
Mycological Research. 109 (Pt 6): 661–86. doi
:10.1017/S095375620500273X . PMID 16080390 .
* ^ Clay K, Schardl C (October 2002). "
Evolutionary origins and
ecological consequences of endophyte symbiosis with grasses". The
American Naturalist. 160 Suppl 4 (suppl. 4): S99–S127. doi
:10.1086/342161 . PMID 18707456 .
* ^ Brodo IM, Sharnoff SD (2001).
Lichens of North America. New
Haven, Connecticut: Yale University Press. ISBN 0-300-08249-5 .
* ^ Raven PH, Evert RF, Eichhorn, SE (2005). "14—Fungi". Biology
Plants (7 ed.). W. H. Freeman. p. 290. ISBN 978-0-7167-1007-3 .
* ^ Deacon, p. 267.
* ^ Purvis W (2000). Lichens. Washington, D.C.: Smithsonian
Institution Press in association with the Natural History Museum,
London. pp. 49–75. ISBN 1-56098-879-7 .
* ^ Kirk et al., p. 378.
* ^ Deacon, pp. 267–276.
* ^ Douglas AE (November 1989). "Mycetocyte symbiosis in insects".
Biological Reviews of the Cambridge Philosophical Society. 64 (4):
409–34. doi :10.1111/j.1469-185X.1989.tb00682.x . PMID 2696562 .
* ^ Deacon, p. 277.
* ^ "Entomologists: Brazilian Stingless Bee Must Cultivate Special
Fungus to Survive". Sci-News.com. 23 October 2015. Retrieved
25 October 2015.
* ^ Nguyen NH, Suh SO, Blackwell M (2007). "Five novel Candida
species in insect-associated yeast clades isolated from
other insects". Mycologia. 99 (6): 842–58. doi
:10.3852/mycologia.99.6.842 . PMID 18333508 .
* ^ A B Filipiak, Michał; Weiner, January (2016-09-01).
"Nutritional dynamics during the development of xylophagous beetles
related to changes in the stoichiometry of 11 elements". Physiological
Entomology. 42: n/a–n/a. doi :10.1111/phen.12168 . ISSN 1365-3032
* ^ Filipiak, Michał; Sobczyk, Łukasz; Weiner, January
(2016-04-09). "Fungal Transformation of Tree Stumps into a Suitable
Resource for Xylophagous Beetles via Changes in Elemental Ratios".
Insects. 7 (2): 13. doi :10.3390/insects7020013 .
* ^ Chandler PJ (2010). A Dipterist's Handbook (2nd Edition). U.K.:
The Amateur Entomologists' Society. pp. 1–525.
* ^ Talbot NJ (2003). "On the trail of a cereal killer: Exploring
the biology of Magnaporthe grisea". Annual Review of Microbiology. 57:
177–202. doi :10.1146/annurev.micro.57.030502.090957 . PMID 14527276
* ^ Paoletti M, Buck KW, Brasier CM (January 2006). "Selective
acquisition of novel mating type and vegetative incompatibility genes
via interspecies gene transfer in the globally invading eukaryote
Ophiostoma novo-ulmi". Molecular Ecology. 15 (1): 249–62. doi
:10.1111/j.1365-294X.2005.02728.x . PMID 16367844 .
* ^ Gryzenhout M, Wingfield BD, Wingfield MJ (May 2006). "New
taxonomic concepts for the important forest pathogen Cryphonectria
parasitica and related fungi". FEMS
Microbiology Letters. 258 (2):
161–72. doi :10.1111/j.1574-6968.2006.00170.x . PMID 16640568 .
* ^ Yang Y, Yang E, An Z, Liu X (May 2007). "
nematode-trapping cells of predatory fungi of the Orbiliaceae based on
evidence from rRNA-encoding DNA and multiprotein sequences" .
Proceedings of the National Academy of Sciences of the United States
of America. 104 (20): 8379–84.
Bibcode :2007PNAS..104.8379Y. doi
:10.1073/pnas.0702770104 . PMC 1895958 . PMID 17494736 .
* ^ Nielsen K, Heitman J (2007). "Sex and virulence of human
pathogenic fungi". Advances in Genetics. Advances in Genetics. 57:
143–73. doi :10.1016/S0065-2660(06)57004-X . ISBN 978-0-12-017657-1
. PMID 17352904 .
* ^ Brakhage AA (December 2005). "Systemic fungal infections caused
Aspergillus species: epidemiology, infection process and virulence
determinants". Current Drug Targets. 6 (8): 875–86. doi
:10.2174/138945005774912717 . PMID 16375671 .
* ^ Kauffman CA (January 2007). "Histoplasmosis: a clinical and
laboratory update" . Clinical
Microbiology Reviews. 20 (1): 115–32.
doi :10.1128/CMR.00027-06 . PMC 1797635 . PMID 17223625 .
* ^ Cushion MT, Smulian AG, Slaven BE, Sesterhenn T, Arnold J,
Staben C, Porollo A, Adamczak R, Meller J (2007). "Transcriptome of
Pneumocystis carinii during fulminate infection: carbohydrate
metabolism and the concept of a compatible parasite" . PLoS One. 2
Bibcode :2007PLoSO...2..423C. doi
:10.1371/journal.pone.0000423 . PMC 1855432 . PMID 17487271 .
* ^ Cook GC, Zumla AI (2008). Manson's Tropical Diseases: Expert
Consult. Edinburgh, Scotland: Saunders Ltd. p. 347. ISBN 1-4160-4470-1
* ^ Simon-Nobbe B, Denk U, Pöll V, Rid R, Breitenbach M (2008).
"The spectrum of fungal allergy". International Archives of Allergy
and Immunology. 145 (1): 58–86. doi :10.1159/000107578 . PMID
* ^ Le Floch G, Rey P, Benizri E, Benhamou N, Tirilly Y (2003).
"Impact of auxin-compounds produced by the antagonistic fungus Pythium
oligandrum or the minor pathogen
Pythium group F on plant growth".
Plant Soil. 257 (2): 459–470. doi :10.1023/a:1027330024834 .
* ^ Schardl CL, Panaccione DG, Tudzynski P (2006). "Ergot
alkaloids--biology and molecular biology". The Alkaloids. Chemistry
and Biology. The Alkaloids: Chemistry and Biology. 63: 45–86. doi
:10.1016/S1099-4831(06)63002-2 . ISBN 978-0-12-469563-4 . PMID
* ^ van Egmond HP, Schothorst RC, Jonker MA (September 2007).
"Regulations relating to mycotoxins in food: perspectives in a global
and European context". Analytical and Bioanalytical Chemistry. 389
(1): 147–57. doi :10.1007/s00216-007-1317-9 . PMID 17508207 .
* ^ Demain AL, Fang A (2000). "The natural functions of secondary
metabolites". Advances in Biochemical Engineering/Biotechnology.
Advances in Biochemical Engineering/Biotechnology. 69: 1–39. doi
:10.1007/3-540-44964-7_1 . ISBN 978-3-540-67793-2 . PMID 11036689 .
* ^ Rohlfs M, Albert M, Keller NP, Kempken F (October 2007).
"Secondary chemicals protect mould from fungivory" .
3 (5): 523–5. doi :10.1098/rsbl.2007.0338 . PMC 2391202 . PMID
* ^ Molina L, Kahmann R (July 2007). "An
Ustilago maydis gene
involved in H2O2 detoxification is required for virulence" . The Plant
Cell. 19 (7): 2293–309. doi :10.1105/tpc.107.052332 . PMC 1955693
. PMID 17616735 .
* ^ Kojic M, Zhou Q, Lisby M, Holloman WK (January 2006). "Rec2
interplay with both Brh2 and Rad51 balances recombinational repair in
Ustilago maydis" . Molecular and Cellular Biology. 26 (2): 678–88.
doi :10.1128/MCB.26.2.678-688.2006 . PMC 1346908 . PMID 16382157 .
* ^ A B Michod RE, Bernstein H, Nedelcu AM (May 2008). "Adaptive
value of sex in microbial pathogens" (PDF). Infection,
Evolution. 8 (3): 267–85. doi :10.1016/j.meegid.2008.01.002 . PMID
* ^ A B Fan W, Kraus PR, Boily MJ, Heitman J (August 2005).
Cryptococcus neoformans gene expression during murine macrophage
infection" . Eukaryotic Cell. 4 (8): 1420–33. doi
:10.1128/EC.4.8.1420-1433.2005 . PMC 1214536 . PMID 16087747 .
* ^ A B Lin X, Hull CM, Heitman J (April 2005). "Sexual
reproduction between partners of the same mating type in Cryptococcus
neoformans". Nature. 434 (7036): 1017–21. Bibcode
:2005Natur.434.1017L. doi :10.1038/nature03448 . PMID 15846346 .
* ^ Fincham JR (March 1989). "Transformation in fungi" .
Microbiological Reviews. 53 (1): 148–70. PMC 372721 . PMID
* ^ Hawkins KM, Smolke CD (September 2008). "Production of
benzylisoquinoline alkaloids in
Saccharomyces cerevisiae" . Nature
Chemical Biology. 4 (9): 564–73. doi :10.1038/nchembio.105 . PMC
2830865 . PMID 18690217 .
* ^ Huang B, Guo J, Yi B, Yu X, Sun L, Chen W (July 2008).
"Heterologous production of secondary metabolites as pharmaceuticals
Biotechnology Letters. 30 (7):
1121–37. doi :10.1007/s10529-008-9663-z . PMID 18512022 .
* ^ Brakhage AA, Spröte P, Al-Abdallah Q, Gehrke A, Plattner H,
Tüncher A (2004). "Regulation of penicillin biosynthesis in
filamentous fungi". Advances in Biochemical Engineering/Biotechnology.
Advances in Biochemical Engineering/Biotechnology. 88: 45–90. doi
:10.1007/b99257 . ISBN 978-3-540-22032-9 . PMID 15719552 .
* ^ Pan A, Lorenzotti S, Zoncada A (January 2008). "Registered and
investigational drugs for the treatment of methicillin-resistant
Staphylococcus aureus infection". Recent Patents on Anti-Infective
Drug Discovery. 3 (1): 10–33. doi :10.2174/157489108783413173 . PMID
* ^ Fajardo A, Martínez JL (April 2008). "Antibiotics as signals
that trigger specific bacterial responses". Current Opinion in
Microbiology. 11 (2): 161–7. doi :10.1016/j.mib.2008.02.006 . PMID
* ^ Loo DS (2006). "Systemic antifungal agents: an update of
established and new therapies". Advances in Dermatology. 22: 101–24.
doi :10.1016/j.yadr.2006.07.001 . PMID 17249298 .
* ^ Manzoni M, Rollini M (April 2002). "
biotechnological production of statins by filamentous fungi and
application of these cholesterol-lowering drugs". Applied Microbiology
and Biotechnology. 58 (5): 555–64. doi :10.1007/s00253-002-0932-9 .
PMID 11956737 .
* ^ el-Mekkawy S, Meselhy MR, Nakamura N, Tezuka Y, Hattori M,
Kakiuchi N, Shimotohno K, Kawahata T, Otake T (November 1998).
"Anti-HIV-1 and anti-HIV-1-protease substances from Ganoderma
lucidum". Phytochemistry. 49 (6): 1651–7. doi
:10.1016/S0031-9422(98)00254-4 . PMID 9862140 .
* ^ El Dine RS, El Halawany AM, Ma CM, Hattori M (June 2008).
"Anti-HIV-1 protease activity of lanostane triterpenes from the
vietnamese mushroom Ganoderma colossum". Journal of Natural Products.
71 (6): 1022–6. doi :10.1021/np8001139 . PMID 18547117 .
* ^ A B Hetland G, Johnson E, Lyberg T, Bernardshaw S, Tryggestad
AM, Grinde B (October 2008). "Effects of the medicinal mushroom
Agaricus blazei Murill on immunity, infection and cancer".
Scandinavian Journal of Immunology. 68 (4): 363–70. doi
:10.1111/j.1365-3083.2008.02156.x . PMID 18782264 .
* ^ Yuen JW, Gohel MD (2005). "Anticancer effects of Ganoderma
lucidum: a review of scientific evidence". Nutrition and Cancer. 53
(1): 11–7. doi :10.1207/s15327914nc5301_2 . PMID 16351502 .
* ^ Sullivan R, Smith JE, Rowan NJ (2006). "Medicinal mushrooms and
cancer therapy: translating a traditional practice into Western
medicine". Perspectives in
Biology and Medicine. 49 (2): 159–70. doi
:10.1353/pbm.2006.0034 . PMID 16702701 .
* ^ Halpern GM, Miller A (2002). Medicinal Mushrooms: Ancient
Remedies for Modern Ailments. New York, New York: M. Evans and Co. p.
116. ISBN 0-87131-981-0 .
* ^ Fisher M, Yang LX (2002). "Anticancer effects and mechanisms of
polysaccharide-K (PSK): implications of cancer immunotherapy".
Anticancer Research. 22 (3): 1737–54. PMID 12168863 .
* ^ Firenzuoli F, Gori L, Lombardo G (March 2008). "The Medicinal
Mushroom Agaricus blazei Murrill: Review of Literature and
Pharmaco-Toxicological Problems" . Evidence-Based Complementary and
Alternative Medicine. 5 (1): 3–15. doi :10.1093/ecam/nem007 . PMC
2249742 . PMID 18317543 .
* ^ Paterson RR (September 2006). "Ganoderma - a therapeutic fungal
biofactory". Phytochemistry. 67 (18): 1985–2001. doi
:10.1016/j.phytochem.2006.07.004 . PMID 16905165 .
* ^ Paterson RR (May 2008). "Cordyceps: a traditional Chinese
medicine and another fungal therapeutic biofactory?". Phytochemistry.
69 (7): 1469–95. doi :10.1016/j.phytochem.2008.01.027 . PMID
* ^ Kulp K (2000). Handbook of
Cereal Science and Technology. CRC
Press. ISBN 0-8247-8294-1 .
* ^ Piskur J, Rozpedowska E, Polakova S, Merico A, Compagno C
(April 2006). "How did
Saccharomyces evolve to become a good brewer?".
Trends in Genetics. 22 (4): 183–6. doi :10.1016/j.tig.2006.02.002 .
PMID 16499989 .
* ^ Abe K, Gomi K, Hasegawa F, Machida M (September 2006). "Impact
Aspergillus oryzae genomics on industrial production of
metabolites". Mycopathologia. 162 (3): 143–53. doi
:10.1007/s11046-006-0049-2 . PMID 16944282 .
* ^ Hachmeister KA, Fung DY (1993). "Tempeh: a mold-modified
indigenous fermented food made from soybeans and/or cereal grains".
Critical Reviews in Microbiology. 19 (3): 137–88. doi
:10.3109/10408419309113527 . PMID 8267862 .
* ^ Jørgensen TR (December 2007). "Identification and toxigenic
potential of the industrially important fungi,
Aspergillus oryzae and
Aspergillus sojae". Journal of Food Protection. 70 (12): 2916–34.
PMID 18095455 .
* ^ O'Donnell K, Cigelnik E, Casper HH (February 1998). "Molecular
phylogenetic, morphological, and mycotoxin data support
reidentification of the Quorn mycoprotein fungus as Fusarium
Genetics and Biology. 23 (1): 57–67. doi
:10.1006/fgbi.1997.1018 . PMID 9501477 .
* ^ Stamets P (2000). Growing Gourmet and Medicinal Mushrooms = .
Berkeley, California: Ten Speed Press. pp. 233–248. ISBN
* ^ Hall, pp. 13–26.
* ^ Kinsella JE, Hwang DH (November 1976). "Enzymes of Penicillium
roqueforti involved in the biosynthesis of cheese flavor". CRC
Critical Reviews in Food Science and Nutrition. 8 (2): 191–228. doi
:10.1080/10408397609527222 . PMID 21770 .
* ^ Erdogan A, Gurses M, Sert S (August 2003). "Isolation of moulds
capable of producing mycotoxins from blue mouldy Tulum cheeses
produced in Turkey". International Journal of Food Microbiology. 85
(1–2): 83–5. doi :10.1016/S0168-1605(02)00485-3 . PMID 12810273 .
* ^ Orr DB, Orr RT (1979). Mushrooms of Western North America.
Berkeley, California: University of California Press. p. 17. ISBN
* ^ Vetter J (January 1998). "Toxins of
Toxicon. 36 (1): 13–24. doi :10.1016/S0041-0101(97)00074-3 . PMID
* ^ Leathem AM, Dorran TJ (March 2007). "Poisoning due to raw
Gyromitra esculenta (false morels) west of the Rockies". Cjem. 9 (2):
127–30. doi :10.1017/s1481803500014937 . PMID 17391587 .
* ^ Karlson-Stiber C, Persson H (September 2003). "Cytotoxic
fungi--an overview". Toxicon. 42 (4): 339–49. doi
:10.1016/S0041-0101(03)00238-1 . PMID 14505933 .
* ^ Michelot D, Melendez-Howell LM (February 2003). "Amanita
muscaria: chemistry, biology, toxicology, and ethnomycology".
Mycological Research. 107 (Pt 2): 131–46. doi
:10.1017/S0953756203007305 . PMID 12747324 .
* ^ Hall, p. 7.
* ^ Ammirati JF, McKenny M, Stuntz DE (1987). The New Savory Wild
Mushroom. Seattle, Washington: University of Washington Press. pp.
xii–xiii. ISBN 0-295-96480-4 .
* ^ López-Gómez J, Molina-Meyer M (February 2006). "The
competitive exclusion principle versus biodiversity through
competitive segregation and further adaptation to spatial
heterogeneities". Theoretical Population Biology. 69 (1): 94–109.
doi :10.1016/j.tpb.2005.08.004 . PMID 16223517 .
* ^ Becker H (1998). "Setting the Stage To Screen Biocontrol
Fungi". United States Department of Agriculture, Agricultural Research
Service. Retrieved 23 February 2009.
* ^ Keiller TS. "Whey-based fungal microfactory technology for
enhanced biological pest management using fungi" (PDF). UVM
Innovations. Archived from the original (PDF) on 29 October 2011.
Retrieved 29 October 2011.
* ^ Deshpande MV (1999). "Mycopesticide production by fermentation:
potential and challenges". Critical Reviews in Microbiology. 25 (3):
229–43. doi :10.1080/10408419991299220 . PMID 10524330 .
* ^ Thomas MB, Read AF (May 2007). "Can fungal biopesticides
Nature Reviews. Microbiology. 5 (5): 377–83. doi
:10.1038/nrmicro1638 . PMID 17426726 .
* ^ Bush LP, Wilkinson HH, Schardl CL (May 1997). "Bioprotective
Alkaloids of Grass-Fungal
Endophyte Symbioses" .
Plant Physiology. 114
(1): 1–7. doi :10.1104/pp.114.1.1 . PMC 158272 . PMID 12223685 .
* ^ Bouton JH, Latch GC, Hill NS, Hoveland CS, McCann MA, Watson
RH, Parish JA, Hawkins LL, Thompson FN (2002). "Reinfection of Tall
Fescue Cultivars with Non-Ergot Alkaloid–Producing Endophytes".
Agronomy Journal . 94 (3): 567–574. doi :10.2134/agronj2002.5670 .
Review by Parish JA, McCann MA, Watson RH, Hoveland CS, Hawkins LL,
Hill NS, Bouton JH (May 2003). "Use of nonergot alkaloid-producing
endophytes for alleviating tall fescue toxicosis in sheep". Journal of
Animal Science. 81 (5): 1316–22. doi :10.2527/2003.8151316x . PMID
* ^ Christian V, Shrivastava R, Shukla D, Modi HA, Vyas BR (April
2005). "Degradation of xenobiotic compounds by lignin-degrading
white-rot fungi: enzymology and mechanisms involved". Indian Journal
of Experimental Biology. 43 (4): 301–12. PMID 15875713 .
* ^ "Fungi to fight \'toxic war zones\'". BBC News. 5 May 2008.
* ^ Fomina M, Charnock JM, Hillier S, Alvarez R, Gadd GM (July
2007). "Fungal transformations of uranium oxides". Environmental
Microbiology. 9 (7): 1696–710. doi :10.1111/j.1462-2920.2007.01288.x
. PMID 17564604 .
* ^ Fomina M, Charnock JM, Hillier S, Alvarez R, Livens F, Gadd GM
(May 2008). "Role of fungi in the biogeochemical fate of depleted
uranium". Current Biology. 18 (9): R375–7. doi
:10.1016/j.cub.2008.03.011 . PMID 18460315 .
* ^ Beadle GW, Tatum EL (November 1941). "Genetic Control of
Biochemical Reactions in Neurospora" . Proceedings of the National
Academy of Sciences of the United States of America. 27 (11):
Bibcode :1941PNAS...27..499B. doi :10.1073/pnas.27.11.499 .
PMC 1078370 . PMID 16588492 .
* ^ Datta A, Ganesan K, Natarajan K (1989). "Current trends in
Candida albicans research". Advances in Microbial Physiology. Advances
in Microbial Physiology. 30: 53–88. doi
:10.1016/S0065-2911(08)60110-1 . ISBN 978-0-12-027730-8 . PMID 2700541
* ^ Dean RA, Talbot NJ, Ebbole DJ, Farman ML, Mitchell TK, Orbach
MJ, et al. (April 2005). "The genome sequence of the rice blast fungus
Magnaporthe grisea". Nature. 434 (7036): 980–6. Bibcode
:2005Natur.434..980D. doi :10.1038/nature03449 . PMID 15846337 .
* ^ Daly R, Hearn MT (2005). "Expression of heterologous proteins
Pichia pastoris: a useful experimental tool in protein engineering
and production". Journal of Molecular Recognition. 18 (2): 119–38.
doi :10.1002/jmr.687 . PMID 15565717 .
* ^ Schlegel HG (1993). General Microbiology. Cambridge, UK:
Cambridge University Press. p. 360. ISBN 0-521-43980-9 .
* ^ Joseph B, Ramteke PW, Thomas G (2008). "Cold active microbial
lipases: some hot issues and recent developments". Biotechnology
Advances. 26 (5): 457–70. doi :10.1016/j.biotechadv.2008.05.003 .
PMID 18571355 .
* ^ Kumar R, Singh S, Singh OV (May 2008). "Bioconversion of
lignocellulosic biomass: biochemical and molecular perspectives".
Journal of Industrial
Microbiology & Biotechnology. 35 (5): 377–91.
doi :10.1007/s10295-008-0327-8 . PMID 18338189 .
* ^ "Trichoderma spp., including T. harzianum, T. viride, T.
koningii, T. hamatum and other spp. Deuteromycetes, Moniliales
(asexual classification system)". Biological Control: A Guide to
Natural Enemies in North America. Archived from the original on 14
April 2011. Retrieved 10 July 2007.
* ^ Olempska-
Beer ZS, Merker RI, Ditto MD, DiNovi MJ (July 2006).
"Food-processing enzymes from recombinant microorganisms--a review".
Regulatory Toxicology and Pharmacology. 45 (2): 144–158. doi
:10.1016/j.yrtph.2006.05.001 . PMID 16769167 .
* ^ Polizeli ML, Rizzatti AC, Monti R, Terenzi HF, Jorge JA, Amorim
DS (June 2005). "Xylanases from fungi: properties and industrial
Microbiology and Biotechnology. 67 (5):
577–91. doi :10.1007/s00253-005-1904-7 . PMID 15944805 .
* Ainsworth GC (1976). Introduction to the History of Mycology.
Cambridge, UK: Cambridge University Press. ISBN 0-521-11295-8 .
* Alexopoulos CJ, Mims CW, Blackwell M (1996). Introductory
Mycology. John Wiley and Sons. ISBN 0-471-52229-5 .
* Chandler PJ (2010). A Dipterist's Handbook (2nd ed.). The Amateur
Entomologists' Society. pp. 1–525.
* Deacon J (2005). Fungal Biology. Cambridge, Massachusetts:
Blackwell Publishers. ISBN 1-4051-3066-0 .
* Hall IR (2003). Edible and Poisonous Mushrooms of the World.
Portland, Oregon: Timber Press. ISBN 0-88192-586-1 .
* Hanson JR (2008). The Chemistry of Fungi. Royal Society Of
Chemistry. ISBN 0-85404-136-2 .
* Jennings DH, Lysek G (1996). Fungal Biology: Understanding the
Fungal Lifestyle. Guildford, UK: Bios Scientific Publishers Ltd. ISBN
* Kirk PM, Cannon PF, Minter DW, Stalpers JA (2008). Dictionary of
the Fungi (10th ed.). Wallingford, UK: CAB International. ISBN
* Taylor EL, Taylor TN (1993). The
Evolution of Fossil
Plants. Englewood Cliffs, New Jersey: Prentice Hall. ISBN
Look up FUNGUS in Wiktionary, the free dictionary.
Find more aboutFUNGUSat's sister projects
* Definitions from Wiktionary
* Media from Wikimedia Commons
* News from Wikinews
* Quotations from Wikiquote
* Texts from Wikisource
* Textbooks from Wikibooks
* Learning resources from Wikiversity
* Taxonomy from Wikispecies
Library resources about
* Online books
* Resources in your library
* Resources in other libraries
* Tree of
Life web project: Fungi
Fungus at the Encyclopedia