Fungus

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Fungi are a kingdom of organisms that includes yeasts, molds and mushrooms. They play important roles in decomposition and nutrient cycling in ecosystems.

Fungi are classified as a separate kingdom from plants and animals. They have chitin in their cell walls and acquire nutrients by absorbing dissolved molecules from their environment.

Fungi are important decomposers in ecosystems and play roles in decomposition, nutrient cycling and exchange. They are also used as a food source and in various food and industrial processes.

Fungus

A fungus (plural: fungi[3] or funguses[4]) 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
Fungi
the other eukaryotic life kingdoms of plants and animals. Temporal range: Early Devonian–Present
(but see text) 410–0 Ma
A characteristic that places fungi in a different kingdom from plants, bacteria, and some protists is chitin in their cell walls. Similar to
PreЄ Є O S D C P T J K PgN
animals, fungi are heterotrophs; they acquire their food by absorbing dissolved molecules, typically by secreting digestive enzymes into
their environment. Fungi do not photosynthesize. 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 the 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 to plants.

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
Clockwise from top left:
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.
Amanita muscaria, a basidiomycete;
Many species produce bioactive compounds called mycotoxins, such as alkaloids and polyketides, that are toxic to animals including Sarcoscypha coccinea, an ascomycete;
humans. The fruiting structures of a few species contain psychotropic compounds and are consumed recreationally or in traditional
bread covered in mold;
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
a chytrid;
supplies and local economies. an Aspergillus conidiophore.

The fungus kingdom encompasses an enormous diversity of taxa with varied ecologies, life cycle strategies, and morphologies ranging Scientific classification
from unicellular aquatic chytrids to large mushrooms. However, little is known of the true biodiversity of Kingdom Fungi, which has (unranked): Opisthokonta
been estimated at 2.2 million to 3.8 million species.[5] Of these, only about 120,000 have been described, with over 8,000 species
(unranked): Holomycota
known to be detrimental to plants and at least 300 that can be pathogenic to humans.[6] Ever since the pioneering 18th and 19th century
taxonomical works of Carl Linnaeus, Christian Hendrik Persoon, and Elias Magnus Fries, fungi have been classified according to their (unranked): Zoosporia
morphology (e.g., characteristics such as spore color or microscopic features) or physiology. Advances in molecular genetics have
Kingdom: Fungi
opened the way for DNA analysis to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on
(L.) R.T.Moore[1]
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 subphyla. Subkingdoms/Phyla/Subphyla[2]

Blastocladiomycota
Contents Chytridiomycota
Glomeromycota
Etymology
Microsporidia
Characteristics
Neocallimastigomycota
Diversity
Mycology Dikarya (inc. Deuteromycota)
History
Morphology Ascomycota
Microscopic structures
Macroscopic structures Pezizomycotina
Growth and physiology Saccharomycotina
Reproduction Taphrinomycotina
Asexual reproduction
Sexual reproduction
Basidiomycota
Spore dispersal
Homothallism
Other sexual processes
Agaricomycotina
Evolution
Pucciniomycotina
Taxonomy
Ustilaginomycotina
Taxonomic groups
Fungus-like organisms Subphyla incertae sedis

Ecology
Symbiosis
Entomophthoromycotina
With plants Kickxellomycotina
With algae and cyanobacteria Mucoromycotina
With insects
Zoopagomycotina
As pathogens and parasites
As targets of mycoparasites

Mycotoxins
Pathogenic mechanisms
Human use
Therapeutic uses
Modern chemotherapeutics
Traditional and folk medicine
Cultured foods
In food
Poisonous fungi
Pest control
Bioremediation
Model organisms
Others
See also
References
Cited literature
External links

Etymology
The English word fungus is directly adopted from the Latin fungus (mushroom), used in the writings of Horace and Pliny.[7] This in turn is derived from the Greek word sphongos (σφόγγος
"sponge"), which refers to the macroscopic structures and morphology of mushrooms and molds;[8] the root is also used in other languages, such as the German Schwamm ("sponge") and Schimmel
("mold").[9]

The word mycology is derived from the Greek mykes (μύκης "mushroom") and logos (λόγος "discourse").[10] It denotes the scientific study of fungi. The Latin adjectival form of "mycology"
(mycologicæ) appeared as early as 1796 in a book on the subject by Christiaan Hendrik Persoon.[11] The word appeared in English as early as 1824 in a book by Robert Kaye Greville.[12] In 1836
the English naturalist Miles Joseph Berkeley's publication The English Flora of Sir James Edward Smith, Vol. 5. also refers to mycology as the study of fungi.[8][13]

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 Ireland".[14]

Characteristics
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 (around the start of the Neoproterozoic Era).[15][16] 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:
Fungal hyphae cells
Shared features:
1 Hyphal wall · 2 Septum ·
3 Mitochondrion · 4 Vacuole ·
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, 5 Ergosterol crystal · 6 Ribosome ·
sterol-containing membranes, and ribosomes of the 80S type.[17] They have a characteristic range of soluble carbohydrates and 7 Nucleus · 8 Endoplasmic reticulum
storage compounds, including sugar alcohols (e.g., mannitol), disaccharides, (e.g., trehalose), and polysaccharides (e.g., · 9 Lipid body · 10 Plasma membrane
glycogen, which is also found in animals[18]). · 11 Spitzenkörper · 12 Golgi
With animals: Fungi lack chloroplasts and are heterotrophic organisms and so require preformed organic compounds as energy apparatus
sources.[19]
With plants: Fungi have a cell wall[20] and vacuoles.[21] 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.[22]
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.[23][24]
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.[25] Both fungi
and oomycetes grow as filamentous hyphal cells.[26] In contrast, similar-looking organisms, such as filamentous green algae, grow by repeated cell division within a chain of
cells.[18] There are also single-celled fungi (yeasts) that do not form hyphae, and some fungi have both hyphal and yeast forms.[27]
In common with some plant and animal species, more than 70 fungal species display bioluminescence.[28]
Unique features:

Some species grow as unicellular yeasts that reproduce by budding or fission. Dimorphic fungi can switch between a yeast phase and a hyphal phase in response to
environmental conditions.[27]
The fungal cell wall is composed of glucans and chitin; while glucans are also found in plants and chitin in the exoskeleton of arthropods,[29][30] 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.[31]
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,[32] 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.[33] Plants and some other organisms have an additional terpene biosynthesis pathway in their chloroplasts, a structure fungi and
animals do not have.[34] Fungi produce several secondary metabolites that are similar or identical in structure to those made by plants.[33] 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.[33][35]

Diversity
Fungi have a worldwide distribution, and grow in a wide range of habitats, including extreme environments such as deserts or areas with high salt
Omphalotus nidiformis, a
concentrations[36] or ionizing radiation,[37] as well as in deep sea sediments.[38] Some can survive the intense UV and cosmic radiation
bioluminescent mushroom
encountered during space travel.[39] 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.[40] Other
examples of aquatic fungi include those living in hydrothermal areas of the ocean.[41]

Around 120,000 species of fungi have been described by taxonomists,[42] but the global biodiversity of the fungus kingdom is not fully
understood.[42] A 2017 estimate suggests there may be between 2.2 and 3.8 million species.[5] 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.[43] 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 Bracket fungi on a tree stump
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.[44]

Mycology
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.[45]

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).[46] 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.[47]

History
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 publication of Pier Antonio Micheli's
1729 work Nova plantarum genera.[48] 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.[49] Extending the use of the binomial system of nomenclature introduced by Carl
Linnaeus in his 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 In 1729, Pier Antonio Micheli
centuries include Miles Joseph Berkeley, August Carl Joseph Corda, Anton de Bary, the brothers Louis René and Charles Tulasne, Arthur H. R. Buller, first published descriptions
Curtis G. Lloyd, and Pier Andrea Saccardo. The 20th century has seen a modernization of mycology that has come from advances in biochemistry, genetics, of fungi.
molecular biology, and biotechnology. The use of DNA sequencing technologies and phylogenetic analysis has provided new insights into fungal
relationships and biodiversity, and has challenged traditional morphology-based groupings in fungal taxonomy.[50]

Morphology

Microscopic structures
Most fungi grow as hyphae, which are cylindrical, thread-like structures 2–10 µm in diameter and up to several centimeters in
length. 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.[51]
Hyphae also sometimes fuse when they come into contact, a process called hyphal fusion (or anastomosis). These growth
processes lead to the development of a mycelium, an interconnected network of hyphae.[27] 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.[52] 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.[53] Coenocytic hyphae are in essence
multinucleate supercells.[54]
An environmental isolate of Penicillium
Many species have developed specialized hyphal structures for nutrient uptake from living hosts; examples include haustoria 1 hypha · 2 conidiophore · 3 phialide · 4 conidia · 5 septa
in plant-parasitic species of most fungal phyla, and arbuscules of several mycorrhizal fungi, which penetrate into the host cells
to consume nutrients.[55]

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.[56] 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.[57]

Macroscopic structures
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.[58] Some individual fungal colonies can reach extraordinary dimensions and ages as in the case of a clonal colony of Armillaria solidipes,
which extends over an area of more than 900 ha (3.5 square miles), with an estimated age of nearly 9,000 years.[59]

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.[60] The fruit bodies of the basidiomycetes (basidiocarps) and some
ascomycetes can sometimes grow very large, and many are well known as mushrooms.
Armillaria solidipes

Growth and physiology


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.[61] Hyphae are specifically adapted for growth on solid surfaces, and to invade
substrates and tissues.[62] They can exert large penetrative mechanical forces; for example, many plant pathogens, including Magnaporthe grisea, form
a structure called an appressorium that evolved to puncture plant tissues.[63] The pressure generated by the appressorium, directed against the plant
epidermis, can exceed 8 megapascals (1,200 psi).[63] The filamentous fungus Paecilomyces lilacinus uses a similar structure to penetrate the eggs of
nematodes.[64]

The mechanical pressure exerted by the appressorium is generated from physiological processes that increase intracellular turgor by producing
osmolytes such as glycerol.[65] 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.[66][67][68] The vast
Mold growth covering a decaying majority of filamentous fungi grow in a polar fashion (extending in one direction) by elongation at the tip (apex) of the hypha.[69] Other forms of
peach. The frames were taken fungal growth include intercalary extension (longitudinal expansion of hyphal compartments that are below the apex) as in the case of some endophytic
approximately 12 hours apart fungi,[70] or growth by volume expansion during the development of mushroom stipes and other large organs.[71] Growth of fungi as multicellular
over a period of six days.
structures consisting of somatic and reproductive cells—a feature independently evolved in animals and plants[72]—has several functions, including the
development of fruit bodies for dissemination of sexual spores (see above) and biofilms for substrate colonization and intercellular communication.[73]

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.[74][75] 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.[37] This process might bear similarity to CO2 fixation via visible light, but instead uses ionizing radiation as a
source of energy.[76]

Reproduction
Fungal reproduction is complex, reflecting the differences in lifestyles and genetic makeup within this diverse kingdom of organisms.[77] 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.[78] 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
Asexual reproduction occurs via vegetative spores (conidia) or through mycelial fragmentation. Mycelial fragmentation occurs when a fungal
Polyporus squamosus
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.[79] The "Fungi imperfecti" (fungi lacking
the perfect or sexual stage) or Deuteromycota comprise all the species that lack an observable sexual cycle.[80] Deuteromycota is not an accepted taxonomic clade, and is now taken to mean simply
fungi that lack a known sexual stage.

Sexual reproduction
Sexual reproduction with meiosis has been directly observed in all fungal phyla except Glomeromycota [81] (genetic analysis suggests meiosis in 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.[82][83] Mating experiments between fungal isolates may identify species on the basis of biological species concepts.[83] 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. Fungi employ two mating systems: heterothallic species allow mating only between individuals of opposite mating type, whereas homothallic species can mate, and
sexually reproduce, with any other individual or itself.[84]

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).[85]

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.[86]

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
The 8-spore asci of Morchella elata, clamp connection in the basidiomycetes is required for controlled transfer of nuclei during cell division, to maintain the dikaryotic stage with two
viewed with phase contrast genetically different nuclei in each hyphal compartment.[87] A basidiocarp is formed in which club-like structures known as basidia generate
microscopy
haploid basidiospores after karyogamy and meiosis.[88] The most commonly known basidiocarps are mushrooms, but they may also take other
forms (see Morphology section).

In fungi formerly classified as Zygomycota, 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 mycelia.[89]

Spore dispersal
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.
Specialized mechanical and physiological mechanisms, as well as spore surface structures (such as hydrophobins), enable efficient spore
ejection.[90] 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.[91] 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;[92] 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.[93] 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.[94] Another strategy is seen in the
stinkhorns, a group of fungi with lively colors and putrid odor that attract insects to disperse their spores.[95]

The most common means of spore dispersal is by wind - species using this form of dispersal often produce dry or hydrophobic spores which do not The bird's nest fungus Cyathus
absorb water and are readily scattered by raindrops, for example. Most of the researched species of fungus are transported by wind.[96][97] stercoreus

Homothallism
In homothallic sexual reproduction, two haploid nuclei derived form the same individual fuse to form a zygote that can then undergo meiosis. Homothallic fungi include species with an aspergillus-
like asexual stage (anamorphs) occurring in numerous different genera,[98] several species of the ascomycete genus Cochliobolus,[99] and the ascomycete Pneumocystis jiroveccii.[100]
Heitman[101] reviewed evidence bearing on the evolution of sexual reproduction in the fungi and concluded that the earliest mode of sexual reproduction among eukaryotes was likely
homothallism, that is, self-fertile unisexual reproduction.

Other sexual processes


Besides regular sexual reproduction with meiosis, certain fungi, such as those in the genera Penicillium and Aspergillus, may exchange genetic material via parasexual processes, initiated by
anastomosis between hyphae and plasmogamy of fungal cells.[102] 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[103] and is likely required for hybridization between species, which has been associated with major events in fungal evolution.[104]

Evolution
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.[105] 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.[106] Researchers study compression fossils by dissolving the
surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details.[107]

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.[108] 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.[109]
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.[110] 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.[111] Recent (2009) studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent
lichenization events have occurred multiple times.[112]

In May 2019, scientists reported the discovery of a fossilized fungus, named Ourasphaira giraldae, in the Canadian Arctic, that may have grown on land a billion years ago, well before plants were
living on land.[113][114][115] Earlier, it had been presumed that the fungi colonized the land during the Cambrian (542–488.3 Ma), also long before land plants.[116] Fossilized hyphae and spores
recovered from the Ordovician of Wisconsin (460 Ma) resemble modern-day Glomerales, and existed at a time when the land flora likely consisted of only non-vascular bryophyte-like plants.[117]
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, mostly as Zygomycota and Chytridiomycota.[116][118][119] At about this same time, approximately 400 Ma, the Ascomycota and
Basidiomycota diverged,[120] and all modern classes of fungi were present by the Late Carboniferous (Pennsylvanian, 318.1–299 Ma).[121]

Lichen-like fossils have been found in the Doushantuo Formation in southern China dating back to 635–551 Ma.[122] Lichens formed a component of the early terrestrial ecosystems, and the
estimated age of the oldest terrestrial lichen fossil is 400 Ma;[123] this date corresponds to the age of the oldest known sporocarp fossil, a Paleopyrenomycites species found in the Rhynie
Chert.[124] The oldest fossil with microscopic features resembling modern-day basidiomycetes is Palaeoancistrus, found permineralized with a fern from the Pennsylvanian.[125] 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 Ma.[126][127]

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.[128] However, the relative proportion of fungal spores relative to spores formed by
algal species is difficult to assess,[129] the spike did not appear worldwide,[130][131] and in many places it did not fall on the Permian–Triassic boundary.[132]

65 million years ago, immediately after the Cretaceous–Paleogene extinction event that famously killed off most dinosaurs, there is a dramatic increase in evidence of fungi, apparently the death of
most plant and animal species leading to a huge fungal bloom like "a massive compost heap".[133]

Taxonomy
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.[134] Analyses using molecular phylogenetics support a monophyletic origin of fungi.[44] The taxonomy of 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 matings.[135]

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.[44][136] 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.[44] 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,[137] "The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied
Research"[138] and Tedersoo et al. 2018.[139] The lengths of the branches are not proportional to evolutionary distances.

Rozellomycota Rozellomycetes

Mitosporidium

Paramicrosporidium
Rozellomyceta
Microsporidiomycota Nucleophaga

Metchnikovellea

Microsporea

Aphelidiomyceta Aphelidiomycota Aphelidiomycetes

Eumycota Neocallimastigomycota Neocallimastigomycetes

Hyaloraphidiomycetes

Monoblepharomycotina Monoblepharidomycetes
Chytridiomyceta
Chytridiomycota Sanchytriomycetes

Mesochytriomycetes
Chytridiomycotina
Chytridiomycetes

Blastocladiomyceta Blastocladiomycota Blastocladiomycetes

Physodermatomycetes

Basidiobolomycetes
Basidiobolomycota
Zoosporia Olpidiomycetes

Neozygitomycetes
Entomophthoromycota
Zoopagomyceta Entomophthoromycetes

Zoopagomycotina Zoopagomycete

Kickxellomycota Dimargaritomy
Kickxellomycotina
Kickxellomyce

Mortierellomycota Mortierellomycetes

Calcarisporiellomycota Calcarisporiellomycetes
Amastigomycota
Mucoromyceta Umbelopsidomycetes
Mucoromycota
Mucoromycetes

Paraglomeromycetes

Glomeromycota Archaeosporomycetes

Glomeromycetes
Symbiomycota
Entorrhizomycota Entorrhizomycetes

Dikarya Basidiomycota

Ascomycota

Basidiomycota Pucciniomycotina
Tritirachiomycetes

Mixiomycetes

Agaricostilbomycetes
Cystobasidiomycetes

Classiculaceae

Microbotryomycetes

Cryptomycocolacomycetes

Atractiellomycetes

Pucciniomycetes

Monilielliomycetes

Malasseziomycetes
Ustilaginomycotina
Ustilaginomycetes

Exobasidiomycetes

?Geminibasidiomycetes
Orthomycotina
?Wallemiomycetes

Bartheletiomycetes
Agaricomycotina
Tremellomycetes

Dacrymycetes

Agaricomycetes

Ascomycota
Neolectomycetes

Taphrinomycetes

Taphrinomycotina Archaeorhizomycetes

Schizosaccharomyceta Pneumocystidomycetes

Schizosaccharomycetes

Saccharomycotina Saccharomycetes

Pezizomycotina ?Thelocarpales

?Vezdaeales

?Lahmiales

?Triblidiales

Orbiliomycetes

Pezizomycetes

Leotiomyceta Xylonomycetes

Geoglossomycetes

Sordariomyceta Leotiomycetes
Saccharomyceta
Laboulbeniomycetes

Sordariomycetes

Coniocybomycetes

Lichinomycetes

Eurotiomycetes

Dothideomyceta Lecanoromycetes
Collemopsidiomycetes

Arthoniomycetes

Dothideomycetes

Taxonomic groups
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.[44]

Phylogenetic analysis has demonstrated that the Microsporidia, unicellular parasites of animals and protists, are
fairly recent and highly derived endobiotic fungi (living within the tissue of another species).[110][140] 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.[141] Hibbett and colleagues suggest that this analysis does not clash with their
classification of the Fungi, and although the Microsporidia are elevated to phylum status, it is acknowledged that
further analysis is required to clarify evolutionary relationships within this group.[44]

The Chytridiomycota are commonly known as chytrids. These fungi are distributed worldwide. Chytrids and
their close relatives Neocallimastigomycota and 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 Main groups of fungi.
sequences in ribosomes, suggest that the Chytrids are a basal group divergent from the other fungal phyla,
consisting of four major clades with suggestive evidence for paraphyly or possibly polyphyly.[142]

The 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 Dikarya (Ascomycota 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 meiosis.[110]

The 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).[143] They lack mitochondria but contain hydrogenosomes of mitochondrial
origin. As in the related chrytrids, neocallimastigomycetes form zoospores that are posteriorly uniflagellate or polyflagellate.[44]

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. All known Glomeromycota species reproduce asexually.[81] The symbiotic
association between the Glomeromycota and plants is ancient, with evidence dating to 400 million years ago.[144] 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.[145] Fungi that were placed in the Zygomycota are now being reassigned to the Glomeromycota, or the subphyla incertae sedis
Mucoromycotina, Kickxellomycotina, the Zoopagomycotina and the Entomophthoromycotina.[44] 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.[146] Medically relevant genera include Mucor, Rhizomucor, and Rhizopus.

The Ascomycota, commonly known as sac fungi or ascomycetes, constitute the largest taxonomic group within the Eumycota.[43] These fungi form Arbuscular mycorrhiza seen under
meiotic spores called ascospores, which are enclosed in a special sac-like structure called an ascus. This phylum includes morels, a few mushrooms microscope. Flax root cortical cells
containing paired arbuscules.
and truffles, unicellular yeasts (e.g., of the genera Saccharomyces, Kluyveromyces, 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 Aspergillus,
Penicillium, 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.[147] 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).[148]

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,[149] human commensal species of the genus Malassezia,[150] and the
opportunistic human pathogen, Cryptococcus neoformans.[151]

Fungus-like organisms Diagram of an apothecium (the


typical cup-like reproductive structure
Because of similarities in morphology and lifestyle, the slime molds (mycetozoans, plasmodiophorids, acrasids, Fonticula and labyrinthulids, now of Ascomycetes) showing sterile
in Amoebozoa, Rhizaria, Excavata, Opisthokonta and Stramenopiles, respectively), water molds (oomycetes) and hyphochytrids (both tissues as well as developing and
Stramenopiles) were formerly classified in the kingdom Fungi, in groups like Mastigomycotina, Gymnomycota and Phycomycetes. The slime mature asci.
molds were studied also as protozoans, leading to an 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 research literature.[152]

The Eccrinales and Amoebidiales are opisthokont protists, previously thought to be zygomycete fungi. Other groups now in Opisthokonta (e.g., Corallochytrium, Ichthyosporea) were also at given
time classified as fungi. The genus Blastocystis, now in Stramenopiles, was originally classified as a yeast. Ellobiopsis, now in Alveolata, was considered a chytrid. The bacteria were also included
in fungi in some classifications, as the group Schizomycetes.

The 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 of fungi.
The nucleariids may be the next sister group to the eumycete clade, and as such could be included in an expanded fungal kingdom.[134]

Ecology
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[153]
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 organisms.[154][155]

Symbiosis
Many fungi have important symbiotic relationships with organisms from most if not all Kingdoms.[156][157][158] These interactions can be A pin mold decomposing a peach
mutualistic or antagonistic in nature, or in the case of commensal fungi are of no apparent benefit or detriment to the host.[159][160][161]

With plants
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.[162]

The mycorrhizal symbiosis is ancient, dating to at least 400 million years ago.[144] 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.[154][163] The fungal partners may also mediate plant-to-plant transfer
of carbohydrates and other nutrients.[164] Such mycorrhizal communities are called "common mycorrhizal networks".[165][166] A special case of mycorrhiza
is myco-heterotrophy, whereby the plant parasitizes the fungus, obtaining all of its nutrients from its fungal symbiont.[167] Some fungal species inhabit the
tissues inside roots, stems, and leaves, in which case they are called endophytes.[168] 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 return.[169]

With algae and cyanobacteria


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
The dark filaments are basidiomycetes.[170] Lichens occur in every ecosystem on all continents, play a key role in soil formation
hyphae of the endophytic and the initiation of biological succession,[171] and are prominent in some extreme environments, including
fungus Neotyphodium
polar, alpine, and semiarid desert regions.[172] They are able to grow on inhospitable surfaces, including
coenophialum in the
bare soil, rocks, tree bark, wood, shells, barnacles and leaves.[173] As in mycorrhizas, the photobiont
intercellular spaces of tall
fescue leaf sheath tissue 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 The lichen Lobaria pulmonaria, a
that they function almost as a single organism; in most cases the resulting organism differs greatly from the symbiosis of fungal, algal, and
individual components. Lichenization is a common mode of nutrition for fungi; around 20% of fungi—between 17,500 and 20,000 described cyanobacterial species
species—are lichenized.[174] 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 habitat.[175]

With insects
Many insects also engage in mutualistic relationships with fungi. Several groups of ants cultivate fungi in the order Agaricales as their primary food source, while ambrosia beetles cultivate various
species of fungi in the bark of trees that they infest.[176] 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.[177] 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.[178] Termites on the African savannah are also
known to cultivate fungi,[156] 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.[179] The larvae of many families of fungicolous flies, particularly those within the superfamily Sciaroidea such as the Mycetophilidae and some Keroplatidae feed on
fungal fruiting bodies and sterile mycorrhizae.[180]

As pathogens and parasites


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,[181] tree pathogens such as Ophiostoma ulmi and
Ophiostoma novo-ulmi causing Dutch elm disease[182] and Cryphonectria parasitica responsible for chestnut blight,[183] and plant pathogens in the
genera Fusarium, Ustilago, Alternaria, and Cochliobolus.[160] 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.[184] Many fungi that are plant pathogens,
such as Magnaporthe oryzae, can switch from being biotrophic (parasitic on living plants) to being necrotrophic (feeding on the dead tissues of
plants they have killed).[185] This same principle is applied to fungi-feeding parasites, including Asterotremella albida, which feeds on the fruit
bodies of other fungi both while they are living and after they are dead.[186]

The plant pathogen Aecidium


Some fungi can cause serious diseases in humans, several of which may be fatal if untreated. These include aspergillosis, candidiasis,
magellanicum causes calafate rust,
coccidioidomycosis, cryptococcosis, histoplasmosis, mycetomas, and paracoccidioidomycosis. Furthermore, persons with immuno-deficiencies are
seen here on a Berberis shrub in
particularly susceptible to disease by genera such as Aspergillus, Candida, Cryptoccocus,[161][187][188] Histoplasma,[189] and Pneumocystis.[190] Chile.
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.[191] Fungal spores are also a cause of allergies, and fungi from different taxonomic groups can evoke allergic
reactions.[192]
As targets of mycoparasites
The organisms which parasitize fungi are known as mycoparasitic organisms. Certain species of the genus Pythium, which are oomycetes, have potential as biocontrol agents against certain
fungi.[193] Fungi can also act as mycoparasites or antagonists of other fungi, such as Hypomyces chrysospermus, which grows on bolete mushrooms.

Mycotoxins
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 fungus, Claviceps purpurea.[194]
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 trichothecenes (e.g., T-2 mycotoxin) and
fumonisins, which have significant impact on human food supplies or animal livestock.[195]

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 fungi.[33] Mycotoxins may provide fitness benefits in terms of physiological
Ergotamine, a major mycotoxin
adaptation, competition with other microbes and fungi, and protection from consumption (fungivory).[196][197] Many fungal secondary metabolites
produced by Claviceps species,
(or derivatives) are used medically, as described under Human Use below. which if ingested can cause
gangrene, convulsions, and
Pathogenic mechanisms hallucinations

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.[198] Furthermore, U. maydis has a well-established recombinational DNA repair system which acts during mitosis and meiosis.[199] The system may assist the pathogen in
surviving DNA damage arising from the host plant's oxidative defensive response to infection.[200]

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.[201] 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.[201] 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.[202] 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.[200][202]

Human use
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,[203] enabling metabolic engineering of fungal species. For example, genetic modification of yeast species[204]—
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.[205]

Therapeutic uses Saccharomyces cerevisiae


cells shown with DIC
microscopy
Modern chemotherapeutics
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 (produced by 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.[206] 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.[207] 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.[208]
Other drugs produced by fungi include griseofulvin isolated from Penicillium griseofulvum, used to treat fungal infections,[209] and statins (HMG-
The mould Penicillium chrysogenum
CoA reductase inhibitors), used to inhibit cholesterol synthesis. Examples of statins found in fungi include mevastatin from Penicillium citrinum
was the source of penicillin G.
and lovastatin from Aspergillus terreus and the oyster mushroom.[210] Fungi produce compounds that inhibit viruses[211][212] and cancer
cells.[213][214] 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.[215][216] In Europe and Japan, polysaccharide-K (brand name
Krestin), a chemical derived from Trametes versicolor, is an approved adjuvant for cancer therapy.[217]

Traditional and folk medicine


Certain mushrooms enjoy usage as therapeutics in folk medicines, such as Traditional Chinese medicine. Notable medicinal mushrooms with a well-documented history of use include Agaricus
subrufescens,[213][218] Ganoderma lucidum,[219] Psilocybe and Ophiocordyceps sinensis.[220]
Cultured foods
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.[221] Yeast species of the genus
Saccharomyces are also used to produce alcoholic beverages through fermentation.[222] Shoyu
koji mold (Aspergillus oryzae) is an essential ingredient in brewing Shoyu (soy sauce) and sake,
and the preparation of miso,[223] while Rhizopus species are used for making tempeh.[224] Several The medicinal fungi Ganoderma lucidum (left) and Ophiocordyceps sinensis (right)
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.[225] Quorn, a meat substitute, is made from Fusarium venenatum.[226]

In food
Edible mushrooms include commercially raised and wild-harvested fungi. 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 (Flammulina spp.).[227]

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
A selection of edible mushrooms eaten in Asia
boletes) demand a high price on the market. They are often used in gourmet
dishes.[228]

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 Penicillium roqueforti.[229] 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.[230] Stilton cheese veined with
Penicillium roqueforti

Poisonous fungi
Many mushroom species are poisonous to humans and cause a range of reactions including slight digestive problems, allergic reactions, hallucinations,
severe organ failure, and death. Genera with mushrooms containing deadly toxins include Conocybe, Galerina, Lepiota, and, the most infamous,
Amanita.[231] The latter genus includes the destroying angel (A. virosa) and the death cap (A. phalloides), the most common cause of deadly mushroom
poisoning.[232] The false morel (Gyromitra esculenta) is occasionally considered a delicacy when cooked, yet can be highly toxic when eaten raw.[233]
Tricholoma equestre was considered edible until it was implicated in serious poisonings causing rhabdomyolysis.[234] 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 Siberia.[235]

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.[236][237]

Amanita phalloides accounts


Pest control for the majority of fatal
mushroom poisonings
In agriculture, fungi may be useful if they actively compete for nutrients and space with pathogenic worldwide. It sometimes
microorganisms such as bacteria or other fungi via the competitive exclusion principle,[238] or if they are lacks the greenish color
parasites of these pathogens. For example, certain species may be used to eliminate or suppress the growth seen here.
of harmful plant pathogens, such as insects, mites, weeds, nematodes, and other fungi that cause diseases of
important crop plants.[239] 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.[240] Examples
that have been used as biological insecticides are Beauveria bassiana, Metarhizium spp, Hirsutella spp, Paecilomyces (Isaria) spp, and
Lecanicillium lecanii.[241][242] 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.[243] 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,
Grasshoppers killed by Beauveria
while being non-toxic to livestock.[244]
bassiana

Bioremediation
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.[245] Fungi have been shown to biomineralize uranium oxides, suggesting they may have application in the bioremediation of radioactively polluted sites.[246][247][248]

Model organisms
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.[249] Other important model fungi are Aspergillus nidulans and
the yeasts Saccharomyces cerevisiae and 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,[250] Magnaporthe grisea, a plant pathogen,[251] and Pichia pastoris, a yeast widely used for eukaryotic
protein production.[252]
Others
Fungi are used extensively to produce industrial chemicals like citric, gluconic, lactic, and malic acids,[253] and industrial enzymes, such as lipases used in biological detergents,[254] cellulases used
in making cellulosic ethanol[255] and stonewashed jeans,[256] and amylases,[257] invertases, proteases and xylanases.[258]

See also
Conservation of fungi Mycosis
DPVweb Outline of fungi
Marine fungi Plant pathology
MycoBank

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Cited literature
Ainsworth GC (1976). Introduction to the History of Mycology. Cambridge, UK: Hanson JR (2008). The Chemistry of Fungi. Royal Society Of Chemistry.
Cambridge University Press. ISBN 978-0-521-11295-6. ISBN 978-0-85404-136-7.
Alexopoulos CJ, Mims CW, Blackwell M (1996). Introductory Mycology. John Wiley Jennings DH, Lysek G (1996). Fungal Biology: Understanding the Fungal Lifestyle.
and Sons. ISBN 978-0-471-52229-4. Guildford, UK: Bios Scientific Publishers Ltd. ISBN 978-1-85996-150-6.
Chandler PJ (2010). A Dipterist's Handbook (2nd ed.). The Amateur Entomologists' Kirk PM, Cannon PF, Minter DW, Stalpers JA (2008). Dictionary of the Fungi (10th
Society. pp. 1–525. ed.). Wallingford, UK: CAB International. ISBN 978-0-85199-826-8.
Deacon J (2005). Fungal Biology. Cambridge, Massachusetts: Blackwell Taylor EL, Taylor TN (1993). The Biology and Evolution of Fossil Plants.
Publishers. ISBN 978-1-4051-3066-0. Englewood Cliffs, New Jersey: Prentice Hall. ISBN 978-0-13-651589-0.
Hall IR (2003). Edible and Poisonous Mushrooms of the World. Portland, Oregon:
Timber Press. ISBN 978-0-88192-586-9.

External links
Tree of Life web project: Fungi (http://tolweb.org/fungi)
Mushroom Observer (mushroomobserver.org (https://mushroomobserver.org)), a collaborative fungus recording and identification project
FUNGI (http://www.botanical-dermatology-database.info/BotDermFolder/FUNGI.html) in BoDD – Botanical Dermatology Database (http://www.botanical-dermatology-databas
e.info/index.html)

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