A fungus ( : fungi or funguses) is any member of the group of
eukaryotic
Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
organisms that includes microorganisms such as
yeast
Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constit ...
s and
mold
A mold () or mould () is one of the structures certain fungi can form. The dust-like, colored appearance of molds is due to the formation of spores containing fungal secondary metabolites. The spores are the dispersal units of the fungi. Not ...
s, as well as the more familiar
mushroom
A mushroom or toadstool is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground, on soil, or on its food source. ''Toadstool'' generally denotes one poisonous to humans.
The standard for the name "mushroom" is ...
s. These organisms are classified as a kingdom, separately from the other eukaryotic kingdoms, which by one traditional classification include
Plant
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
ae,
Animal
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage ...
ia,
Protozoa
Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
, and
Chromista
Chromista is a biological kingdom consisting of single-celled and multicellular eukaryotic species that share similar features in their photosynthetic organelles ( plastids). It includes all protists whose plastids contain chlorophyll ''c'', ...
.
A characteristic that places fungi in a different kingdom from plants, bacteria, and some protists is
chitin
Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
in their
cell wall
A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mec ...
s. Fungi, like animals, are
heterotroph
A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
s; they acquire their food by absorbing dissolved molecules, typically by secreting
digestive enzyme
Digestive enzymes are a group of enzymes that break down polymeric macromolecules into their smaller building blocks, in order to facilitate their absorption into the cells of the body. Digestive enzymes are found in the digestive tracts of anima ...
s into their environment. Fungi do not photosynthesize. Growth is their means of
mobility
Mobility may refer to:
Social sciences and humanities
* Economic mobility, ability of individuals or families to improve their economic status
* Geographic mobility, the measure of how populations and goods move over time
* Mobilities, a conte ...
, except for
spore
In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
s (a few of which are
flagellate
A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and thei ...
d), 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''), that share a common ancestor (i.e. they form a ''
monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gr ...
group''), an interpretation that is also strongly supported by
molecular phylogenetics
Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
. This fungal group is distinct from the structurally similar myxomycetes (slime molds) and
oomycete
Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the resul ...
s (water molds). The discipline of
biology
Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary ...
devoted to the study of fungi is known as
mycology
Mycology is the branch of biology concerned with the study of fungi, including their genetic and biochemical properties, their taxonomy and their use to humans, including as a source for tinder, traditional medicine, food, and entheogen ...
(from the
Greek
Greek may refer to:
Greece
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group.
*Greek language, a branch of the Indo-European language family.
**Proto-Greek language, the assumed last common ancestor ...
', mushroom). In the past, mycology was regarded as a branch of
botany
Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...
, 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
Cryptic may refer to:
In science:
* Cryptic species complex, a group of species that are very difficult to distinguish from one another
* Crypsis, the ability of animals to blend in to avoid observation
* Cryptic era, earliest period of the Earth ...
lifestyles in soil or on dead matter. Fungi include
symbionts
Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
of plants, animals, or other fungi and also
parasites
Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
. 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
Cycling, also, when on a two-wheeled bicycle, called bicycling or biking, is the use of cycles for transport, recreation, exercise or sport. People engaged in cycling are referred to as "cyclists", "bicyclists", or "bikers". Apart from ...
and exchange in the environment. They have long been used as a direct source of human food, in the form of mushrooms and
truffles
A truffle is the fruiting body of a subterranean ascomycete fungus, predominantly one of the many species of the genus ''Tuber''. In addition to ''Tuber'', many other genera of fungi are classified as truffles including '' Geopora'', '' Pe ...
; as a
leavening agent
In cooking, a leavening agent () or raising agent, also called a leaven () or leavener, is any one of a number of substances used in doughs and batters that cause a foaming action (gas bubbles) that lightens and softens the mixture. An altern ...
for bread; and in the
fermentation
Fermentation is a metabolic process that produces chemical changes in organic substrates through the action of enzymes. In biochemistry, it is narrowly defined as the extraction of energy from carbohydrates in the absence of oxygen. In food p ...
of various food products, such as
wine
Wine is an alcoholic drink typically made from Fermentation in winemaking, fermented grapes. Yeast in winemaking, Yeast consumes the sugar in the grapes and converts it to ethanol and carbon dioxide, releasing heat in the process. Different ...
,
beer
Beer is one of the oldest and the most widely consumed type of alcoholic drink in the world, and the third most popular drink overall after water and tea. It is produced by the brewing and fermentation of starches, mainly derived from ce ...
, and
soy sauce
Soy sauce (also called simply soy in American English and soya sauce in British English) is a liquid condiment of Chinese origin, traditionally made from a fermented paste of soybeans, roasted grain, brine, and ''Aspergillus oryzae'' or '' As ...
. Since the 1940s, fungi have been used for the production of
antibiotic
An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention ...
s, and, more recently, various
enzyme
Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
biological pesticide
A Biopesticide is a biological substance or organism that damages, kills, or repels organisms seens as pests. Biological pest management intervention involves predatory, parasitic, or chemical relationships.
They are obtained from organisms inclu ...
s to control weeds, plant diseases, and insect pests. Many species produce
bioactive compound
A bioactive compound is a compound that has an effect on a living organism, tissue or cell, usually demonstrated by basic research in vitro or in vivo in the laboratory. While dietary nutrients are essential to life, bioactive compounds have not ...
s called
mycotoxin
A mycotoxin (from the Greek μύκης , "fungus" and τοξίνη , "toxin") is a toxic secondary metabolite produced by organisms of kingdom Fungi and is capable of causing disease and death in both humans and other animals. The term 'mycotoxin' ...
s, such as
alkaloid
Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of simila ...
s and
polyketide
Polyketides are a class of natural products derived from a precursor molecule consisting of a chain of alternating ketone (or reduced forms of a ketone) and methylene groups: (-CO-CH2-). First studied in the early 20th century, discovery, biosynth ...
s, that are toxic to animals, including humans. The fruiting structures of a few species contain
psychotropic
A psychoactive drug, psychopharmaceutical, psychoactive agent or psychotropic drug is a chemical substance, that changes functions of the nervous system, and results in alterations in perception, mood, consciousness, cognition or behavior.
Th ...
compounds and are consumed recreationally or in traditional spiritual ceremonies. Fungi can break down manufactured materials and buildings, and become significant
pathogens
In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a ger ...
of humans and other animals. Losses of crops due to fungal diseases (e.g.,
rice blast disease
''Magnaporthe grisea'', also known as rice blast fungus, rice rotten neck, rice seedling blight, blast of rice, oval leaf spot of graminea, pitting disease, ryegrass blast, Johnson spot, neck blast, wheat blast, and Imochi ( Japanese:稲熱) is ...
) or
food spoilage
Food spoilage is the process where a food product becomes unsuitable to ingest by the consumer. The cause of such a process is due to many outside factors as a side-effect of the type of product it is, as well as how the product is packaged and s ...
can have a large impact on human food supplies and local economies.
The fungus kingdom encompasses an enormous diversity of
taxa
In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
with varied ecologies, life cycle strategies, and morphologies ranging from unicellular aquatic
chytrid
Chytridiomycota are a division of zoosporic organisms in the kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoöspores. Chytrid ...
s to large mushrooms. However, little is known of the true
biodiversity
Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
of the fungus kingdom, which has been estimated at 2.2 million to 3.8 million species. Of these, only about 148,000 have been described, with over 8,000 species known to be detrimental to plants and at least 300 that can be pathogenic to humans. Ever since the pioneering 18th and 19th century taxonomical works of
Carl Linnaeus
Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...
,
Christiaan Hendrik Persoon
Christiaan Hendrik Persoon (1 February 1761 – 16 November 1836) was a German mycologist who made additions to Linnaeus' mushroom taxonomy.
Early life
Persoon was born in South Africa at the Cape of Good Hope, the third child of an i ...
, and
Elias Magnus Fries
Elias Magnus Fries (15 August 1794 – 8 February 1878) was a Swedish mycologist and botanist.
Career
Fries was born at Femsjö ( Hylte Municipality), Småland, the son of the pastor there. He attended school in Växjö.
He acquire ...
, fungi have been classified according to their morphology (e.g., characteristics such as spore color or microscopic features) or
physiology
Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemic ...
. Advances in
molecular genetics
Molecular genetics is a sub-field of biology that addresses how differences in the structures or expression of DNA molecules manifests as variation among organisms. Molecular genetics often applies an "investigative approach" to determine the ...
have opened the way for
DNA analysis
Genetic testing, also known as DNA testing, is used to identify changes in DNA sequence or chromosome structure. Genetic testing can also include measuring the results of genetic changes, such as RNA analysis as an output of gene expression, or ...
to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits.
Phylogenetic
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
studies published in the first decade of the 21st century have helped reshape the classification within the fungi kingdom, which is divided into one subkingdom, seven
phyla Phyla, the plural of ''phylum'', may refer to:
* Phylum, a biological taxon between Kingdom and Class
* by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another
Phy ...
The English word ''fungus'' is directly adopted from the
Latin
Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
''fungus'' (mushroom), used in the writings of
Horace
Quintus Horatius Flaccus (; 8 December 65 – 27 November 8 BC), known in the English-speaking world as Horace (), was the leading Roman lyric poet during the time of Augustus (also known as Octavian). The rhetorician Quintilian regarded his ' ...
Greek
Greek may refer to:
Greece
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group.
*Greek language, a branch of the Indo-European language family.
**Proto-Greek language, the assumed last common ancestor ...
word ''sphongos'' (σφόγγος 'sponge'), which refers to the
macroscopic
The macroscopic scale is the length scale on which objects or phenomena are large enough to be visible with the naked eye, without magnifying optical instruments. It is the opposite of microscopic.
Overview
When applied to physical phenomena a ...
structures and morphology of mushrooms and molds;Ainsworth, p. 2. the
root
In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the su ...
is also used in other languages, such as the German '' Schwamm'' ('sponge') and ''
Schimmel
Schimmel is a German and Dutch surname. Notable people with the surname include:
* Annemarie Schimmel (1922–2003), German Islam scholar
* Corrie Schimmel (born 1939), Dutch swimmer
* Hendrik Jan Schimmel (1823–1906), Dutch poet and novelist
...
'' ('mold').
The word ''
mycology
Mycology is the branch of biology concerned with the study of fungi, including their genetic and biochemical properties, their taxonomy and their use to humans, including as a source for tinder, traditional medicine, food, and entheogen ...
'' is derived from the Greek (μύκης 'mushroom') and ''logos'' (λόγος 'discourse'). 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
Christiaan Hendrik Persoon (1 February 1761 – 16 November 1836) was a German mycologist who made additions to Linnaeus' mushroom taxonomy.
Early life
Persoon was born in South Africa at the Cape of Good Hope, the third child of an i ...
. The word appeared in English as early as 1824 in a book by
Robert Kaye Greville
Dr. Robert Kaye Greville FRSE FLS LLD (13 December 1794 – 4 June 1866) was an English mycologist, bryologist, and botanist. He was an accomplished artist and illustrator of natural history. In addition to art and science he was interest ...
. In 1836 the English naturalist
Miles Joseph Berkeley
Miles Joseph Berkeley (1 April 1803 – 30 July 1889) was an English cryptogamist and clergyman, and one of the founders of the science of plant pathology.
Life
Berkeley was born at Biggin Hall, Benefield, Northamptonshire, and educated at R ...
's publication ''The English Flora of Sir James Edward Smith, Vol. 5.'' also refers to mycology as the study of fungi.
A group of all the fungi present in a particular region is known as ''
mycobiota
Mycobiota (plural noun, no singular) are a group of all the fungi present in a particular geographic region (e.g. "the mycobiota of Ireland") or habitat type (e.g. "the mycobiota of cocoa").
Human mycobiota
Mycobiota exist on the surface and in th ...
'' (plural noun, no singular). The term ''mycota'' is often used for this purpose, but many authors use it as a synonym of Fungi. The word ''
funga
In life sciences ''funga'' is a recent term for the kingdom fungi similar to the longstanding fauna for animals and flora for plants. The term was considered to be urgently needed in order to simplify projects oriented toward implementation of edu ...
'' has been proposed as a less ambiguous term morphologically similar to
fauna
Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is '' flora'', and for fungi, it is '' funga''. Flora, fauna, funga and other forms of life are collectively referred to as '' biota''. ...
and
flora
Flora is all the plant life present in a particular region or time, generally the naturally occurring ( indigenous) native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' gut flora'' or '' skin flora''.
...
International Union for Conservation of Nature
The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
(IUCN) in August 2021 asked that the phrase ''fauna and flora'' be replaced by ''fauna, flora, and funga''.
Characteristics
Before the introduction of
molecular methods
Molecular biology is the branch of biology that seeks to understand the molecular basis of biological activity in and between cells, including biomolecular synthesis, modification, mechanisms, and interactions. The study of chemical and physica ...
for phylogenetic analysis,
taxonomists
In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...
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. Although inaccurate, the common misconception that fungi are plants persists among the general public due to their historical classification, as well as several similarities. Like plants, fungi often grow in soil and, in the case of
mushroom
A mushroom or toadstool is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground, on soil, or on its food source. ''Toadstool'' generally denotes one poisonous to humans.
The standard for the name "mushroom" is ...
s, form conspicuous fruit bodies, which sometimes resemble plants such as
mosses
Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hornw ...
. 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
The Neoproterozoic Era is the unit of geologic time from 1 billion to 538.8 million years ago.
It is the last era of the Precambrian Supereon and the Proterozoic Eon; it is subdivided into the Tonian, Cryogenian, and Ediacaran periods. It is prec ...
Era). 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:
Shared features:
* With other
eukaryote
Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacter ...
chromosomes
A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins ar ...
intron
An intron is any Nucleic acid sequence, nucleotide sequence within a gene that is not expressed or operative in the final RNA product. The word ''intron'' is derived from the term ''intragenic region'', i.e. a region inside a gene."The notion of ...
s and coding regions called
exons
An exon is any part of a gene that will form a part of the final mature RNA produced by that gene after introns have been removed by RNA splicing. The term ''exon'' refers to both the DNA sequence within a gene and to the corresponding seque ...
. Fungi have membrane-bound cytoplasmic
organelles
In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
such as
mitochondria
A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used ...
,
sterol
Sterol is an organic compound with formula , whose molecule is derived from that of gonane by replacement of a hydrogen atom in position 3 by a hydroxyl group. It is therefore an alcohol of gonane. More generally, any compounds that contain the go ...
-containing membranes, and ribosomes of the 80S type. They have a characteristic range of soluble carbohydrates and storage compounds, including
sugar alcohol
Sugar alcohols (also called polyhydric alcohols, polyalcohols, alditols or glycitols) are organic compounds, typically derived from sugars, containing one hydroxyl group (–OH) attached to each carbon atom. They are white, water-soluble solids ...
s (e.g.,
mannitol
Mannitol is a type of sugar alcohol used as a sweetener and medication. It is used as a low calorie sweetener as it is poorly absorbed by the intestines. As a medication, it is used to decrease pressure in the eyes, as in glaucoma, and to lo ...
),
disaccharide
A disaccharide (also called a double sugar or ''biose'') is the sugar formed when two monosaccharides are joined by glycosidic linkage. Like monosaccharides, disaccharides are simple sugars soluble in water. Three common examples are sucrose, la ...
s, (e.g.,
trehalose
Trehalose (from Turkish '' tıgala'' – a sugar derived from insect cocoons + -ose) is a sugar consisting of two molecules of glucose. It is also known as mycose or tremalose. Some bacteria, fungi, plants and invertebrate animals synthesize it ...
), and
polysaccharide
Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with w ...
s (e.g.,
glycogen
Glycogen is a multibranched polysaccharide of glucose that serves as a form of energy storage in animals, fungi, and bacteria. The polysaccharide structure represents the main storage form of glucose in the body.
Glycogen functions as one of ...
, which is also found in animals).
* With animals: Fungi lack
chloroplast
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it i ...
s and are
heterotroph
A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
ic organisms and so require preformed
organic compound
In chemistry, organic compounds are generally any chemical compounds that contain carbon-hydrogen or carbon-carbon bonds. Due to carbon's ability to catenate (form chains with other carbon atoms), millions of organic compounds are known. Th ...
s as energy sources.
* With plants: Fungi have a cell wall and
vacuole
A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic m ...
s. They reproduce by both sexual and asexual means, and like basal plant groups (such as
fern
A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
s and
moss
Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta ('' sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and ...
es) produce
spore
In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
s. Similar to mosses and algae, fungi typically have
haploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respective ...
nuclei.
* With euglenoids and bacteria: Higher fungi, euglenoids, and some bacteria produce the
amino acid
Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha ...
L-lysine in specific
biosynthesis
Biosynthesis is a multi-step, enzyme-catalyzed process where substrates are converted into more complex products in living organisms. In biosynthesis, simple compounds are modified, converted into other compounds, or joined to form macromolecul ...
steps, called the α-aminoadipate pathway.
* The cells of most fungi grow as tubular, elongated, and thread-like (filamentous) structures called
hypha
A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium.
Structure
A hypha consists of one or ...
e, which may contain multiple nuclei and extend by growing at their tips. Each tip contains a set of aggregated
vesicles
Vesicle may refer to:
; In cellular biology or chemistry
* Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane
* Synaptic vesicle
; In human embryology
* Vesicle (embryology), bulge-like features o ...
—cellular structures consisting of
protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s,
lipid
Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids in ...
s, and other organic molecules—called the
Spitzenkörper
The Spitzenkörper (German for pointed body, SPK) is a structure found in fungal hyphae that is the organizing center for hyphal growth and morphogenesis. It consists of many small vesicles and is present in growing hyphal tips, during spore ger ...
. Both fungi and
oomycete
Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the resul ...
s grow as filamentous hyphal cells. In contrast, similar-looking organisms, such as filamentous
green algae
The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/ Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga ...
, grow by repeated cell division within a chain of cells. There are also single-celled fungi (
yeast
Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constit ...
s) 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
Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some fungi, microorganisms including some b ...
.
Unique features:
* Some species grow as unicellular yeasts that reproduce by
budding
Budding or blastogenesis is a type of asexual reproduction in which a new organism develops from an outgrowth or bud due to cell division at one particular site. For example, the small bulb-like projection coming out from the yeast cell is kno ...
Dimorphic fungi
Dimorphic fungi are fungi that can exist in the form of both mold and yeast. This is usually brought about by change in temperature and the fungi are also described as thermally dimorphic fungi. An example is '' Talaromyces marneffei'', a huma ...
can switch between a yeast phase and a hyphal phase in response to environmental conditions.
* The fungal cell wall is made of a
chitin-glucan complex Chitin-glucan complex (CGC) is a copolymer (polysaccharide) that makes up fungal cell walls, consisting of covalently-bonded chitin and branched 1,3/1,6-ß-D-glucan. CGCs are alkaline-insoluble. Different species of fungi have different structur ...
; while glucans are also found in plants and chitin in the
exoskeleton
An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton ( endoskeleton) in for example, a human. In usage, some of the ...
of
arthropods
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chitin, ...
, 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.
Most fungi lack an efficient system for the long-distance transport of water and nutrients, such as the
xylem
Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
and
phloem
Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is c ...
in many plants. To overcome this limitation, some fungi, such as ''
Armillaria
''Armillaria'' is a genus of fungi that includes the ''Armillaria mellea, A. mellea'' species known as honey fungi that live on trees and woody shrubs. It includes about 10 species formerly categorized summarily as ''A. mellea''. ''Armill ...
'', form rhizomorphs, which resemble and perform functions similar to the
root
In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the su ...
s of plants. As eukaryotes, fungi possess a
biosynthetic pathway
Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
for producing
terpene
Terpenes () are a class of natural products consisting of compounds with the formula (C5H8)n for n > 1. Comprising more than 30,000 compounds, these unsaturated hydrocarbons are produced predominantly by plants, particularly conifers. Terpenes ...
s that uses
mevalonic acid
Mevalonic acid (MVA) is a key organic compound in biochemistry; the name is a contraction of dihydroxymethylvalerolactone. The carboxylate anion of mevalonic acid, which is the predominant form in biological environments, is known as ''mevalonat ...
and
pyrophosphate
In chemistry, pyrophosphates are phosphorus oxyanions that contain two phosphorus atoms in a P–O–P linkage. A number of pyrophosphate salts exist, such as disodium pyrophosphate (Na2H2P2O7) and tetrasodium pyrophosphate (Na4P2O7), among othe ...
as chemical building blocks. Plants and some other organisms have an additional terpene biosynthesis pathway in their chloroplasts, a structure that fungi and animals do not have. Fungi produce several
secondary metabolite
Secondary metabolites, also called specialised metabolites, toxins, secondary products, or natural products, are organic compounds produced by any lifeform, e.g. bacteria, fungi, animals, or plants, which are not directly involved in the norma ...
s 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
In mathematics, a sequence is an enumerated collection of objects in which repetitions are allowed and order matters. Like a set, it contains members (also called ''elements'', or ''terms''). The number of elements (possibly infinite) is called ...
and other characteristics, which indicates separate origins and
convergent evolution
Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
of these enzymes in the fungi and plants.
Diversity
Fungi have a worldwide distribution, and grow in a wide range of habitats, including extreme environments such as
deserts
A desert is a barren area of landscape where little precipitation occurs and, consequently, living conditions are hostile for plant and animal life. The lack of vegetation exposes the unprotected surface of the ground to denudation. About on ...
or areas with high salt concentrations or
ionizing radiation
Ionizing radiation (or ionising radiation), including nuclear radiation, consists of subatomic particles or electromagnetic waves that have sufficient energy to ionize atoms or molecules by detaching electrons from them. Some particles can travel ...
, as well as in
deep sea
The deep sea is broadly defined as the ocean depth where light begins to fade, at an approximate depth of 200 metres (656 feet) or the point of transition from continental shelves to continental slopes. Conditions within the deep sea are a combin ...
cosmic radiation
Cosmic rays are high-energy particles or clusters of particles (primarily represented by protons or atomic nuclei) that move through space at nearly the speed of light. They originate from the Sun, from outside of the Solar System in our ow ...
encountered during space travel. Most grow in terrestrial environments, though several species live partly or solely in aquatic habitats, such as the
chytrid
Chytridiomycota are a division of zoosporic organisms in the kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoöspores. Chytrid ...
fungi ''
Batrachochytrium dendrobatidis
''Batrachochytrium dendrobatidis'' ( ), also known as ''Bd'' or the amphibian chytrid fungus, is a fungus that causes the disease chytridiomycosis in amphibians.
Since its discovery in 1998 by Lee Berger, the disease devastated amphibian popula ...
parasite
Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson h ...
s that have been responsible for a worldwide decline in
amphibian
Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...
populations. These organisms spend part of their life cycle as a motile zoospore, enabling them to propel itself through water and enter their amphibian host. Other examples of aquatic fungi include those living in
hydrothermal
Hydrothermal circulation in its most general sense is the circulation of hot water (Ancient Greek ὕδωρ, ''water'',Liddell, H.G. & Scott, R. (1940). ''A Greek-English Lexicon. revised and augmented throughout by Sir Henry Stuart Jones. with th ...
areas of the ocean.
As of 2020, around 148,000 species of fungi have been described by
taxonomists
In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...
, 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. The number of new fungi species discovered yearly has increased from 1,000 to 1,500 per year about 10 years ago, to about 2000 with a peak of more than 2,500 species in 2016. In the year 2019, 1882 new species of fungi were described, and it was estimated that more than 90% of fungi remain unknown. The following year, 2905 new species were described—the highest annual record of new fungus names. 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.Kirk ''et al''., p. 489. Species may also be distinguished by their
biochemical
Biochemistry or biological chemistry is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology an ...
and
physiological
Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemica ...
characteristics, such as their ability to metabolize certain biochemicals, or their reaction to
chemical tests
A chemical substance is a form of matter having constant chemical composition and characteristic properties. Some references add that chemical substance cannot be separated into its constituent elements by physical separation methods, i.e., wit ...
. The
biological species concept
The species problem is the set of questions that arises when biologists attempt to define what a species is. Such a definition is called a species concept; there are at least 26 recognized species concepts. A species concept that works well for se ...
discriminates species based on their ability to mate. The application of
molecular
A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bio ...
tools, such as
DNA sequencing
DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. T ...
and phylogenetic analysis, to study diversity has greatly enhanced the resolution and added robustness to estimates of
genetic diversity
Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species, it ranges widely from the number of species to differences within species and can be attributed to the span of survival for a species. It is dis ...
within various taxonomic groups.
Mycology
Mycology
Mycology is the branch of biology concerned with the study of fungi, including their genetic and biochemical properties, their taxonomy and their use to humans, including as a source for tinder, traditional medicine, food, and entheogen ...
is the branch of
biology
Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary ...
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
Plant pathology (also phytopathology) is the scientific study of diseases in plants caused by pathogens (infectious organisms) and environmental conditions (physiological factors). Organisms that cause infectious disease include fungi, oomy ...
, the study of plant diseases, is closely related because many plant pathogens are fungi.
The use of fungi by humans dates back to prehistory;
Ötzi the Iceman
Ötzi, also called the Iceman, is the natural mummy of a man who lived some time between 3350 and 3105 BC, discovered in September 1991 in the Ötztal Alps (hence the nickname "Ötzi") on the border between Austria and Italy.
Ötzi is believed to ...
, a well-preserved mummy of a 5,300-year-old
Neolithic
The Neolithic period, or New Stone Age, is an Old World archaeological period and the final division of the Stone Age. It saw the Neolithic Revolution, a wide-ranging set of developments that appear to have arisen independently in several pa ...
man found frozen in the Austrian Alps, carried two species of polypore mushrooms that may have been used as
tinder
Tinder is easily combustible material used to start a fire. Tinder is a finely divided, open material which will begin to glow under a shower of sparks. Air is gently wafted over the glowing tinder until it bursts into flame. The flaming tinder i ...
(''
Fomes fomentarius
''Fomes fomentarius'' (commonly known as the tinder fungus, false tinder fungus, hoof fungus, tinder conk, tinder polypore or ice man fungus) is a species of fungal plant pathogen found in Europe, Asia, Africa and North America. The species produ ...
''), 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.
History
Mycology became a systematic science after the development of the
microscope
A microscope () is a laboratory instrument used to examine objects that are too small to be seen by the naked eye. Microscopy is the science of investigating small objects and structures using a microscope. Microscopic means being invisi ...
in the 17th century. Although fungal spores were first observed by
Giambattista della Porta
Giambattista della Porta (; 1535 – 4 February 1615), also known as Giovanni Battista Della Porta, was an Italian scholar, polymath and playwright who lived in Naples at the time of the Renaissance, Scientific Revolution and Reformation.
Giamb ...
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''. 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
Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...
in his ''
Species plantarum
' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genera. It is the first work to consistently apply binomial names and was the ...
'' (1753), the Dutch
Christiaan Hendrik Persoon
Christiaan Hendrik Persoon (1 February 1761 – 16 November 1836) was a German mycologist who made additions to Linnaeus' mushroom taxonomy.
Early life
Persoon was born in South Africa at the Cape of Good Hope, the third child of an i ...
(1761–1836) established the first classification of mushrooms with such skill as to be considered a founder of modern mycology. Later,
Elias Magnus Fries
Elias Magnus Fries (15 August 1794 – 8 February 1878) was a Swedish mycologist and botanist.
Career
Fries was born at Femsjö ( Hylte Municipality), Småland, the son of the pastor there. He attended school in Växjö.
He acquire ...
(1794–1878) further elaborated the
classification Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood.
Classification is the grouping of related facts into classes.
It may also refer to:
Business, organizat ...
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
Miles Joseph Berkeley (1 April 1803 – 30 July 1889) was an English cryptogamist and clergyman, and one of the founders of the science of plant pathology.
Life
Berkeley was born at Biggin Hall, Benefield, Northamptonshire, and educated at R ...
,
August Carl Joseph Corda
August Carl Joseph Corda (1809–1849) was a Czech physician and mycologist. This botanist is denoted by the author abbreviation when citing a botanical name.
Early life and education
Corda was born in Reichenberg (now Liberec), Bohemia on No ...
,
Anton de Bary
Heinrich Anton de Bary (26 January 183119 January 1888) was a German surgeon, botanist, microbiologist, and mycologist (fungal systematics and physiology).
He is considered a founding father of plant pathology (phytopathology) as well as the fou ...
Charles Tulasne
Charles Tulasne (5 September 1816 – 28 August 1884) was a French physician, mycologist and illustrator born in Langeais in the département of Indre-et-Loire.
He received his medical doctorate in 1840 and practiced medicine in Paris until 18 ...
Pier Andrea Saccardo
Pier Andrea Saccardo (23 April 1845 in Treviso, Treviso – 12 February 1920 in Padua) was an Italian botanist and mycologist.
Life
Saccardo studied at the Lyceum in Venice, and then at the Technical Institute of the University of Padua wher ...
. In the 20th and 21st centuries, advances in
biochemistry
Biochemistry or biological chemistry is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology and ...
,
genetics
Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar work ...
,
molecular biology
Molecular biology is the branch of biology that seeks to understand the molecular basis of biological activity in and between cells, including biomolecular synthesis, modification, mechanisms, and interactions. The study of chemical and phys ...
,
biotechnology
Biotechnology is the integration of natural sciences and engineering sciences in order to achieve the application of organisms, cells, parts thereof and molecular analogues for products and services. The term ''biotechnology'' was first used ...
,
DNA sequencing
DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. T ...
and phylogenetic analysis has provided new insights into fungal relationships and
biodiversity
Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
, and has challenged traditional morphology-based groupings in fungal
taxonomy
Taxonomy is the practice and science of categorization or classification.
A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
.
Morphology
Microscopic structures
Most fungi grow as
hypha
A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium.
Structure
A hypha consists of one or ...
e, which are cylindrical, thread-like structures 2–10
µm
The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer ( American spelling), also commonly known as a micron, is a unit of length in the International System of Uni ...
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. Hyphae also sometimes fuse when they come into contact, a process called hyphal fusion (or
anastomosis
An anastomosis (, plural anastomoses) is a connection or opening between two things (especially cavities or passages) that are normally diverging or branching, such as between blood vessels, leaf veins, or streams. Such a connection may be norm ...
). These growth processes lead to the development of a
mycelium
Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates ...
, 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 angle
In geometry and trigonometry, a right angle is an angle of exactly 90 degrees or radians corresponding to a quarter turn. If a ray is placed so that its endpoint is on a line and the adjacent angles are equal, then they are right angles. Th ...
s 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
In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. ...
,
organelle
In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' th ...
s, and sometimes nuclei to pass through; an example is the
dolipore septum
Dolipore septa are specialized dividing walls between cells (septa) found in almost all species of fungi in the phylum Basidiomycota. Unlike most fungal septa, they have a barrel-shaped swelling around their central pore, which is about 0.1–0. ...
in fungi of the phylum Basidiomycota. Coenocytic hyphae are in essence
multinucleate
Multinucleate cells (also known as multinucleated or polynuclear cells) are eukaryotic cells that have more than one nucleus per cell, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordina ...
supercells.
Many species have developed specialized hyphal structures for nutrient uptake from living hosts; examples include
haustoria
In botany and mycology, a haustorium (plural haustoria) is a rootlike structure that grows into or around another structure to absorb water or nutrients. For example, in mistletoe or members of the broomrape family, the structure penetrates th ...
in plant-parasitic species of most fungal phyla, and arbuscules of several
mycorrhiza
A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plan ...
l fungi, which penetrate into the host cells to consume nutrients.
Although fungi are
opisthokont
The opisthokonts () are a broad group of eukaryotes, including both the animal and fungus kingdoms. The opisthokonts, previously called the "Fungi/Metazoa group", are generally recognized as a clade. Opisthokonts together with Apusomonadida and ...
s—a grouping of evolutionarily related organisms broadly characterized by a single posterior
flagellum
A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates.
A microorganism may have f ...
—all phyla except for the
chytrids
Chytridiomycota are a division of zoosporic organisms in the kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoöspores. Chytr ...
have lost their posterior flagella. Fungi are unusual among the eukaryotes in having a cell wall that, in addition to
glucan
A glucan is a polysaccharide derived from D-glucose, linked by glycosidic bonds. Glucans are noted in two forms: alpha glucans and beta glucans. Many beta-glucans are medically important. They represent a drug target for antifungal medications o ...
s (e.g., β-1,3-glucan) and other typical components, also contains the
biopolymer
Biopolymers are natural polymers produced by the cells of living organisms. Like other polymers, biopolymers consist of monomeric units that are covalently bonded in chains to form larger molecules. There are three main classes of biopolymers, ...
chitin.
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
mold
A mold () or mould () is one of the structures certain fungi can form. The dust-like, colored appearance of molds is due to the formation of spores containing fungal secondary metabolites. The spores are the dispersal units of the fungi. Not ...
s. Mycelia grown on solid
agar
Agar ( or ), or agar-agar, is a jelly-like substance consisting of polysaccharides obtained from the cell walls of some species of red algae, primarily from ogonori (''Gracilaria'') and "tengusa" (''Gelidiaceae''). As found in nature, agar i ...
media in laboratory
petri dish
A Petri dish (alternatively known as a Petri plate or cell-culture dish) is a shallow transparent lidded dish that biologists use to hold growth medium in which cells can be cultured,R. C. Dubey (2014): ''A Textbook Of Biotechnology For Class-X ...
es are usually referred to as
colonies
In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
. 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 colony of ''
Armillaria solidipes
''Armillaria ostoyae'' (synonym ''Armillaria solidipes'') is a species of fungus (mushroom), pathogenic to trees, in the family Physalacriaceae. In the western United States, it is the most common variant of the group of species under the na ...
'', which extends over an area of more than 900 ha (3.5 square miles), with an estimated age of nearly 9,000years.
The apothecium—a specialized structure important in
sexual reproduction
Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...
in the ascomycetes—is a cup-shaped fruit body that is often macroscopic and holds the
hymenium
The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in others so ...
, a layer of tissue containing the spore-bearing cells. The fruit bodies of the basidiomycetes (
basidiocarp
In fungi, a basidiocarp, basidiome, or basidioma () is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do no ...
s) and some ascomycetes can sometimes grow very large, and many are well known as
mushroom
A mushroom or toadstool is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground, on soil, or on its food source. ''Toadstool'' generally denotes one poisonous to humans.
The standard for the name "mushroom" is ...
s.
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. 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 pathogen
Plant pathology (also phytopathology) is the scientific study of diseases in plants caused by pathogens (infectious organisms) and environmental conditions (physiological factors). Organisms that cause infectious disease include fungi, oomy ...
s, including ''
Magnaporthe grisea
''Magnaporthe grisea'', also known as rice blast fungus, rice rotten neck, rice seedling blight, blast of rice, oval leaf spot of graminea, pitting disease, ryegrass blast, Johnson spot, neck blast, wheat blast, and Imochi ( Japanese:稲熱) is ...
'', form a structure called an
appressorium
An appressorium is a specialized cell typical of many fungal plant pathogens that is used to infect host plants. It is a flattened, hyphal "pressing" organ, from which a minute infection peg grows and enters the host, using turgor pressure capabl ...
that evolved to puncture plant tissues. The pressure generated by the appressorium, directed against the plant
epidermis
The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rel ...
, can exceed . The filamentous fungus '' Paecilomyces lilacinus'' uses a similar structure to penetrate the eggs of
nematode
The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant- parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...
s.
The mechanical pressure exerted by the appressorium is generated from physiological processes that increase intracellular
turgor
Turgor pressure is the force within the cell that pushes the plasma membrane against the cell wall.
It is also called ''hydrostatic pressure'', and is defined as the pressure in a fluid measured at a certain point within itself when at equilibri ...
by producing
osmolyte
Osmolytes are low-molecular weight organic compounds that influence the properties of biological fluids. Their primary role is to maintain the integrity of cells by affecting the viscosity, melting point, and ionic strength of the aqueous solution. ...
s such as
glycerol
Glycerol (), also called glycerine in British English and glycerin in American English, is a simple triol compound. It is a colorless, odorless, viscous liquid that is sweet-tasting and non-toxic. The glycerol backbone is found in lipids known ...
. Adaptations such as these are complemented by
hydrolytic enzymes
Hydrolase is a class of enzyme that commonly perform as biochemical catalysts that use water to break a chemical bond, which typically results in dividing a larger molecule into smaller molecules. Some common examples of hydrolase enzymes are este ...
secreted into the environment to digest large organic molecules—such as
polysaccharide
Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with w ...
s,
protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s, and
lipid
Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids in ...
s—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
An endophyte is an endosymbiont, often a bacterium or fungus, that lives within a plant for at least part of its life cycle without causing apparent disease. Endophytes are ubiquitous and have been found in all species of plants studied to date; h ...
fungi, or growth by volume expansion during the development of mushroom
stipes
Crucifixion is a method of capital punishment in which the victim is tied or nailed to a large wooden cross or beam and left to hang until eventual death from exhaustion and asphyxiation. It was used as a punishment by the Persians, Carthagi ...
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
biofilm
A biofilm comprises any syntrophic consortium of microorganisms in which cells stick to each other and often also to a surface. These adherent cells become embedded within a slimy extracellular matrix that is composed of extracellular po ...
s for substrate colonization and
intercellular communication
"Intercellular communication" refers to the varying ways and structures biological cells use to communicate with each other directly or through their environment. Not all cells use all of the proteins or mechanisms and there are likely to be more. ...
.
The fungi are traditionally considered
heterotroph
A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
s, organisms that rely solely on carbon fixed by other organisms for
metabolism
Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run ...
. Fungi have
evolved
Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variati ...
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
Nitrate is a polyatomic ion with the chemical formula . Salts containing this ion are called nitrates. Nitrates are common components of fertilizers and explosives. Almost all inorganic nitrates are soluble in water. An example of an insolu ...
,
ammonia
Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous ...
,
acetate
An acetate is a salt formed by the combination of acetic acid with a base (e.g. alkaline, earthy, metallic, nonmetallic or radical base). "Acetate" also describes the conjugate base or ion (specifically, the negatively charged ion called ...
, or
ethanol
Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an alcohol with the chemical formula . Its formula can be also written as or (an ethyl group linked to a ...
. In some species the pigment
melanin
Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
may play a role in extracting energy from
ionizing radiation
Ionizing radiation (or ionising radiation), including nuclear radiation, consists of subatomic particles or electromagnetic waves that have sufficient energy to ionize atoms or molecules by detaching electrons from them. Some particles can travel ...
, such as
gamma radiation
A gamma ray, also known as gamma radiation (symbol γ or \gamma), is a penetrating form of electromagnetic radiation arising from the radioactive decay of atomic nuclei. It consists of the shortest wavelength electromagnetic waves, typically s ...
. 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
Light or visible light is electromagnetic radiation that can be perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nanometres (nm), corresponding to frequencies of 750–420 t ...
, but instead uses ionizing radiation as a source of energy.
Reproduction
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 (sexual reproduction) and the
anamorph
In mycology, the terms teleomorph, anamorph, and holomorph apply to portions of the life cycles of fungi in the phyla Ascomycota and Basidiomycota:
*Teleomorph: the sexual reproductive stage (morph), typically a fruiting body.
*Anamorph: an a ...
(asexual reproduction). 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
Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the ...
occurs via vegetative spores (
conidia
A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non- motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to t ...
) 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
Niche may refer to:
Science
*Developmental niche, a concept for understanding the cultural context of child development
* Ecological niche, a term describing the relational position of an organism's species
*Niche differentiation, in ecology, the ...
, and allow more rapid dispersal than sexual reproduction. The "Fungi imperfecti" (fungi lacking the perfect or sexual stage) or Deuteromycota comprise all the species that lack an observable sexual cycle. Deuteromycota (alternatively known as Deuteromycetes, conidial fungi, or mitosporic fungi) 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
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately ...
has been directly observed in all fungal phyla except
Glomeromycota
Glomeromycota (often referred to as glomeromycetes, as they include only one class, Glomeromycetes) are one of eight currently recognized divisions within the kingdom Fungi,
with approximately 230 described species. Members of the Glomeromyco ...
(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. 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 ASCI or Asci may refer to:
* Advertising Standards Council of India
* Asci, the plural of ascus, in fungal anatomy
* Accelerated Strategic Computing Initiative
* American Society for Clinical Investigation
* Argus Sour Crude Index
* Association of ...
and
basidia
A basidium () is a microscopic sporangium (a spore-producing structure) found on the hymenophore of fruiting bodies of basidiomycete fungi which are also called tertiary mycelium, developed from secondary mycelium. Tertiary mycelium is highly- ...
, can be used in the identification of ascomycetes and basidiomycetes, respectively. Fungi employ two
mating system
A mating system is a way in which a group is structured in relation to sexual behaviour. The precise meaning depends upon the context. With respect to animals, the term describes which males and females mate under which circumstances. Recognised ...
s: heterothallic species allow mating only between individuals of the opposite
mating type
Mating types are the microorganism equivalent to sexes in multicellular lifeforms and are thought to be the ancestor to distinct sexes. They also occur in macro-organisms such as fungi.
Definition
Mating types are the microorganism equivalent to ...
, whereas homothallic species can mate, and sexually reproduce, with any other individual or itself.
Most fungi have both a
haploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respective ...
and a
diploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectiv ...
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
An anastomosis (, plural anastomoses) is a connection or opening between two things (especially cavities or passages) that are normally diverging or branching, such as between blood vessels, leaf veins, or streams. Such a connection may be norm ...
, 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
A heterokaryon is a multinucleate cell that contains genetically different nuclei. Heterokaryotic and heterokaryosis are derived terms. This is a special type of syncytium. This can occur naturally, such as in the mycelium of fungi during sexual ...
).
In ascomycetes, dikaryotic hyphae of the
hymenium
The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in others so ...
(the spore-bearing tissue layer) form a characteristic ''hook'' (crozier) at the hyphal septum. During
cell division
Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukaryotes, there ...
, the 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
An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mos ...
, or fruiting body. Karyogamy in the asci is followed immediately by meiosis and the production of
ascospore
An ascus (; ) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some genera ...
s. 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
A clamp connection is a hook-like structure formed by growing hyphal cells of certain fungi. It is a characteristic feature of Basidiomycetes fungi. It is created to ensure that each cell, or segment of hypha separated by septa (cross walls), rec ...
, 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
In fungi, a basidiocarp, basidiome, or basidioma () is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do no ...
is formed in which club-like structures known as
basidia
A basidium () is a microscopic sporangium (a spore-producing structure) found on the hymenophore of fruiting bodies of basidiomycete fungi which are also called tertiary mycelium, developed from secondary mycelium. Tertiary mycelium is highly- ...
generate haploid
basidiospores
A basidiospore is a reproductive spore produced by Basidiomycete fungi, a grouping that includes mushrooms, shelf fungi, rusts, and smuts. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are pro ...
after karyogamy and meiosis. The most commonly known basidiocarps are mushrooms, but they may also take other forms (see Morphology section).
In fungi formerly classified as
Zygomycota
Zygomycota, or zygote fungi, is a former division or phylum of the kingdom Fungi. The members are now part of two phyla: the Mucoromycota and Zoopagomycota. Approximately 1060 species are known. They are mostly terrestrial in habitat, living ...
, haploid hyphae of two individuals fuse, forming a gametangium, a specialized cell structure that becomes a fertile
gamete
A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce ...
-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
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately ...
, 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.
Spore dispersal
The spores of most of the researched species of fungi are transported by wind. Such species often produce dry or
hydrophobic
In chemistry, hydrophobicity is the physical property of a molecule that is seemingly repelled from a mass of water (known as a hydrophobe). In contrast, hydrophiles are attracted to water.
Hydrophobic molecules tend to be nonpolar and, ...
spores that do not absorb water and are readily scattered by raindrops, for example. In other species, 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. 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 0.01–0.02cm, sufficient distance for it to fall through the
gills
A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
or pores into the air below. Other fungi, like the
puffballs
Puffballs are a type of fungus featuring a ball-shaped fruit body that bursts on impact, releasing a cloud of dust-like spores when mature. Puffballs belong to the division Basidiomycota and encompass several genera, including ''Calvatia'', '' ...
, rely on alternative mechanisms for spore release, such as external mechanical forces. The hydnoid fungi (tooth fungi) produce spores on pendant, tooth-like or spine-like projections. 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.
sexual reproduction
Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...
, two
haploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respective ...
nuclei derived from the same individual fuse to form a
zygote
A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism.
In multicell ...
that can then undergo
meiosis
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately ...
. Homothallic fungi include species with an ''Aspergillus''-like asexual stage (anamorphs) occurring in numerous different genera, several species of the
ascomycete
Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The defi ...
genus ''
Cochliobolus
The fungal genus ''Cochliobolus'' includes 55 species, including the following plant pathogenic species: '' C. carbonum'', '' C. heterostrophus'', '' C. miyabeanus'', '' C. sativus'' and '' C. lunatus''.
Heterothallism and homothallism
Those ...
'', and the ascomycete ''
Pneumocystis jirovecii
''Pneumocystis jirovecii'' (previously ''P. carinii'') is a yeast-like fungus of the genus ''Pneumocystis''. The causative organism of ''Pneumocystis'' pneumonia, it is an important human pathogen, particularly among immunocompromised hosts. P ...
Besides regular sexual reproduction with meiosis, certain fungi, such as those in the genera ''
Penicillium
''Penicillium'' () is a genus of ascomycetous fungi that is part of the mycobiome of many species and is of major importance in the natural environment, in food spoilage, and in food and drug production.
Some members of the genus produce pe ...
'' and ''
Aspergillus
' () is a genus consisting of several hundred mold species found in various climates worldwide.
''Aspergillus'' was first catalogued in 1729 by the Italian priest and biologist Pier Antonio Micheli. Viewing the fungi under a microscope, Mic ...
'', 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.
Evolution
In contrast to
plants
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclude ...
and
animals
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
, 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
Extant is the opposite of the word extinct. It may refer to:
* Extant hereditary titles
* Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English
* Extant taxon, a taxon which is not extinct, ...
fungi. Often recovered from a
permineralized
Permineralization is a process of fossilization of bones and tissues in which mineral deposits form internal casts of organisms. Carried by water, these minerals fill the spaces within organic tissue. Because of the nature of the casts, perminera ...
plant or animal host, these samples are typically studied by making thin-section preparations that can be examined with
light microscopy
Microscopy is the technical field of using microscopes to view objects and areas of objects that cannot be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of micr ...
or
transmission electron microscopy
Transmission electron microscopy (TEM) is a microscopy technique in which a beam of electrons is transmitted through a specimen to form an image. The specimen is most often an ultrathin section less than 100 nm thick or a suspension on a ...
. 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
The Paleoproterozoic Era (;, also spelled Palaeoproterozoic), spanning the time period from (2.5–1.6 Ga), is the first of the three sub-divisions ( eras) of the Proterozoic Eon. The Paleoproterozoic is also the longest era of the Earth's ...
benthic
The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning " ...
organisms had filamentous structures capable of
anastomosis
An anastomosis (, plural anastomoses) is a connection or opening between two things (especially cavities or passages) that are normally diverging or branching, such as between blood vessels, leaf veins, or streams. Such a connection may be norm ...
. Other studies (2009) estimate the arrival of fungal organisms at about 760–1060Ma on the basis of comparisons of the rate of evolution in closely related groups. For much of the
Paleozoic
The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon.
The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838
by combining the Greek words ''palaiós'' (, "old") and ...
Era (542–251Ma), the fungi appear to have been aquatic and consisted of organisms similar to the extant
chytrid
Chytridiomycota are a division of zoosporic organisms in the kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoöspores. Chytrid ...
s 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
Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
mycorrhiza
A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plan ...
and lichenization. Studies suggest that the ancestral ecological state of the
Ascomycota
Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The defi ...
was saprobism, and that independent
lichen
A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
ized fungus, named ''
Ourasphaira giraldae
''Ourasphaira giraldae'' is an extinct process-bearing multicellular eukaryotic microorganism. Corentin Loron argues that it was an early fungus. It existed approximately a billion years ago during the time of the transition from the Mesoprotero ...
'', in the
Canadian Arctic
Northern Canada, colloquially the North or the Territories, is the vast northernmost region of Canada variously defined by geography and politics. Politically, the term refers to the three territories of Canada: Yukon, Northwest Territories and ...
, that may have grown on land a billion years ago, well before
plant
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
microfossil
A microfossil is a fossil that is generally between 0.001 mm and 1 mm in size, the visual study of which requires the use of light or electron microscopy. A fossil which can be studied with the naked eye or low-powered magnification, ...
s preserved in the basal Ediacaran Doushantuo Formation (~635 Ma) have been reported in South China. Earlier, it had been presumed that the fungi colonized the land during the
Cambrian
The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
(542–488.3Ma), also long before land plants. Fossilized hyphae and spores recovered from the
Ordovician
The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya.
T ...
of Wisconsin (460Ma) resemble modern-day Glomerales, and existed at a time when the land flora likely consisted of only non-vascular
bryophyte
The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited in s ...
-like plants.Prototaxites, which was probably a fungus or lichen, would have been the tallest organism of the late
Silurian
The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleoz ...
and early
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
. Fungal fossils do not become common and uncontroversial until the early
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
(416–359.2Ma), when they occur abundantly in the
Rhynie chert
The Rhynie chert is a Lower Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness (a Lagerstätte). It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is located ...
, mostly as
Zygomycota
Zygomycota, or zygote fungi, is a former division or phylum of the kingdom Fungi. The members are now part of two phyla: the Mucoromycota and Zoopagomycota. Approximately 1060 species are known. They are mostly terrestrial in habitat, living ...
and
Chytridiomycota
Chytridiomycota are a division of zoosporic organisms in the kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoöspores. Chytri ...
. At about this same time, approximately 400Ma, the Ascomycota and Basidiomycota diverged, and all modern classes of fungi were present by the Late
Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferou ...
( Pennsylvanian, 318.1–299Ma).
Lichens formed a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 415Ma; this date roughly corresponds to the age of the oldest known sporocarp fossil, a ''
Paleopyrenomycites
''Paleopyrenomycites'' is a Devonian genus of fungus of uncertain phylogenetic affinity within the Pezizomycotina total group, known from the Rhynie chert
The Rhynie chert is a Lower Devonian sedimentary deposit exhibiting extraordinary fos ...
'' species found in the Rhynie Chert. The oldest fossil with microscopic features resembling modern-day basidiomycetes is ''Palaeoancistrus'', found permineralized with a
fern
A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
from the Pennsylvanian. Rare in the fossil record are the Homobasidiomycetes (a
taxon
In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
roughly equivalent to the mushroom-producing species of the
Agaricomycetes
The Agaricomycetes are a class of fungi in the division Basidiomycota. The taxon is roughly identical to that defined for the Homobasidiomycetes (alternatively called holobasidiomycetes) by Hibbett & Thorn, with the inclusion of Auriculariales ...
). Two
amber
Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In M ...
-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species ''
Archaeomarasmius leggetti
''Archaeomarasmius'' is an extinct genus of gilled fungus in the Agaricales family Tricholomataceae, containing the single species ''Archaeomarasmius leggetti''. It is known from two fruit bodies recovered from amber, one consisting of a complete ...
'') appeared during the late
Cretaceous
The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
, 90Ma.
Some time after the
Permian–Triassic extinction event
The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, ...
(251.4Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in
sediment
Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sand ...
s) formed, suggesting that fungi were the dominant life form at this time, representing nearly 100% of the available
fossil record
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
for this period. However, the relative proportion of fungal spores relative to spores formed by
algal
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular micr ...
species is difficult to assess, the spike did not appear worldwide, and in many places it did not fall on the Permian–Triassic boundary.
Sixty-five million years ago, immediately after the
Cretaceous–Paleogene extinction event
The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
that famously killed off most dinosaurs, there was a dramatic increase in evidence of fungi; apparently the death of most plant and animal species led to a huge fungal bloom like "a massive compost heap".
Taxonomy
Although commonly included in botany curricula and textbooks, fungi are more closely related to
animal
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage ...
s than to plants and are placed with the animals in the
monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gr ...
group of
opisthokont
The opisthokonts () are a broad group of eukaryotes, including both the animal and fungus kingdoms. The opisthokonts, previously called the "Fungi/Metazoa group", are generally recognized as a clade. Opisthokonts together with Apusomonadida and ...
s. Analyses using
molecular phylogenetics
Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
support a
monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gr ...
origin of fungi. The
taxonomy
Taxonomy is the practice and science of categorization or classification.
A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
of fungi is in a state of constant flux, especially due to 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
mating
In biology, mating is the pairing of either opposite- sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reprod ...
s.
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
Nomenclature (, ) is a system of names or terms, or the rules for forming these terms in a particular field of arts or sciences. The principles of naming vary from the relatively informal conventions of everyday speech to the internationally ag ...
. Until relatively recent (2012) changes to the
International Code of Nomenclature for algae, fungi and plants
The ''International Code of Nomenclature for algae, fungi, and plants'' (ICN) is the set of rules and recommendations dealing with the formal botanical names that are given to plants, fungi and a few other groups of organisms, all those "trad ...
, fungal species could also have multiple scientific names depending on their life cycle and mode (sexual or asexual) of reproduction. Web sites such as
Index Fungorum
''Index Fungorum'' is an international project to index all formal names (scientific names) in the fungus kingdom. the project is based at the Royal Botanic Gardens, Kew, one of three partners along with Landcare Research and the Institute of M ...
and
MycoBank
MycoBank is an online database, documenting new mycological names and combinations, eventually combined with descriptions and illustrations. It is run by the Westerdijk Fungal Biodiversity Institute in Utrecht.
Each novelty, after being screened ...
are officially recognized
nomenclatural
Nomenclature (, ) is a system of names or terms, or the rules for forming these terms in a particular field of arts or sciences. The principles of naming vary from the relatively informal conventions of everyday speech to the internationally agre ...
repositories and list current names of fungal species (with cross-references to older
synonyms
A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are ...
).
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 Phyla, the plural of ''phylum'', may refer to:
* Phylum, a biological taxon between Kingdom and Class
* by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another
Phy ...
, two of which—the Ascomycota and the Basidiomycota—are contained within a branch representing subkingdom
Dikarya
Dikarya is a subkingdom of Fungi that includes the divisions Ascomycota and Basidiomycota, both of which in general produce dikaryons, may be filamentous or unicellular, but are always without flagella. The Dikarya are most of the so-called " ...
, 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
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
depicts the major fungal
taxa
In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
and their relationship to opisthokont and unikont organisms, based on the work of Philippe Silar, "The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research" and Tedersoo et al. 2018. The lengths of the branches are not proportional to
evolutionary
Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variati ...
distances.
Taxonomic groups
The major
phyla Phyla, the plural of ''phylum'', may refer to:
* Phylum, a biological taxon between Kingdom and Class
* by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another
Phy ...
(sometimes called divisions) of fungi have been classified mainly on the basis of characteristics of their sexual
reproductive
The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are a ...
structures. , nine major lineages have been identified: Opisthosporidia, Chytridiomycota, Neocallimastigomycota, Blastocladiomycota, Zoopagomycota, Mucoromycota, Glomeromycota, Ascomycota and Basidiomycota.
Phylogenetic analysis has demonstrated that the
Microsporidia
Microsporidia are a group of spore-forming unicellular parasites. These spores contain an extrusion apparatus that has a coiled polar tube ending in an anchoring disc at the apical part of the spore. They were once considered protozoans or pr ...
, unicellular parasites of animals and protists, are fairly recent and highly derived
biologically active
In pharmacology, biological activity or pharmacological activity describes the beneficial or adverse effects of a drug on living matter. When a drug is a complex chemical mixture, this activity is exerted by the substance's active ingredient or ...
compounds, several of which are
toxic
Toxicity is the degree to which a chemical substance or a particular mixture of substances can damage an organism. Toxicity can refer to the effect on a whole organism, such as an animal, bacterium, or plant, as well as the effect on a sub ...
to animals or plants and are therefore called
mycotoxins
A mycotoxin (from the Greek μύκης , "fungus" and τοξίνη , "toxin") is a toxic secondary metabolite produced by organisms of kingdom Fungi and is capable of causing disease and death in both humans and other animals. The term 'mycotoxin' ...
. Of particular relevance to humans are mycotoxins produced by molds causing food spoilage, and poisonous mushrooms (see above). Particularly infamous are the lethal
amatoxin Amatoxin is the collective name of a subgroup of at least nine related toxic compounds found in three genera of poisonous mushrooms (''Amanita'', '' Galerina'' and '' Lepiota'') and one species ( Conocybe filaris) of the genus '' Conocybe''. Amatoxi ...
s in some ''
Amanita
The genus ''Amanita'' contains about 600 species of agarics, including some of the most toxic known mushrooms found worldwide, as well as some well-regarded edible species. This genus is responsible for approximately 95% of the fatalities resul ...
'' mushrooms, and ergot alkaloids, which have a long history of causing serious epidemics of
ergotism
Ergotism (pron. ) is the effect of long-term ergot poisoning, traditionally due to the ingestion of the alkaloids produced by the '' Claviceps purpurea'' fungus—from the Latin "club" or clavus "nail" and for "head", i.e. the purple club-he ...
(St Anthony's Fire) in people consuming rye or related
cereal
A cereal is any grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, germ, and bran. Cereal grain crops are grown in greater quantities and provide more food ...
s contaminated with sclerotia of the ergot fungus, ''
Claviceps purpurea
''Claviceps purpurea'' is an ergot fungus that grows on the ears of rye and related cereal and forage plants. Consumption of grains or seeds contaminated with the survival structure of this fungus, the ergot sclerotium, can cause ergotism in h ...
''. Other notable mycotoxins include the
aflatoxin
Aflatoxins are various poisonous carcinogens and mutagens that are produced by certain molds, particularly ''Aspergillus'' species. The fungi grow in soil, decaying vegetation and various staple foodstuffs and commodities such as hay, sweetcorn ...
carcinogenic
A carcinogen is any substance, radionuclide, or radiation that promotes carcinogenesis (the formation of cancer). This may be due to the ability to damage the genome or to the disruption of cellular metabolic processes. Several radioactive sub ...
metabolites produced by certain ''
Aspergillus
' () is a genus consisting of several hundred mold species found in various climates worldwide.
''Aspergillus'' was first catalogued in 1729 by the Italian priest and biologist Pier Antonio Micheli. Viewing the fungi under a microscope, Mic ...
'' species often growing in or on grains and nuts consumed by humans,
ochratoxin
Ochratoxins are a group of mycotoxins produced by some ''Aspergillus'' species (mainly ''A. ochraceus'' and A. ''carbonarius'', but also by 33% of ''A. niger'' industrial strains) and some ''Penicillium'' species, especially ''P. verrucosum''. ...
s,
patulin
Patulin is an organic compound classified as a polyketide. It is a white powder soluble in acidic water and in organic solvents. It is a lactone that is heat-stable, so it is not destroyed by pasteurization or thermal denaturation.http://www.s ...
livestock
Livestock are the domesticated animals raised in an agricultural setting to provide labor and produce diversified products for consumption such as meat, eggs, milk, fur, leather, and wool. The term is sometimes used to refer solely to ani ...
.
Mycotoxins are secondary metabolites (or
natural product
A natural product is a natural compound or substance produced by a living organism—that is, found in nature. In the broadest sense, natural products include any substance produced by life. Natural products can also be prepared by chemical sy ...
s), and research has established the existence of biochemical pathways solely for the purpose of producing mycotoxins and other natural products in fungi. 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.
Pathogenic mechanisms
''
Ustilago maydis
Corn smut is a plant disease caused by the pathogenic fungus ''Ustilago maydis'' that causes smut on maize and teosinte. The fungus forms galls on all above-ground parts of corn species. It is edible, and is known in Mexico as the delica ...
'' is a pathogenic plant fungus that causes smut disease in maize and
teosinte
''Zea'' is a genus of flowering plants in the grass family. The best-known species is ''Z. mays'' (variously called maize, corn, or Indian corn), one of the most important crops for human societies throughout much of the world. The four wild spe ...
. 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
In chemistry, reactive oxygen species (ROS) are highly reactive chemicals formed from diatomic oxygen (). Examples of ROS include peroxides, superoxide, hydroxyl radical, singlet oxygen, and alpha-oxygen.
The reduction of molecular oxygen () p ...
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
YAP1 (yes-associated protein 1), also known as YAP or YAP65, is a protein that acts as a transcription coregulator that promotes transcription of genes involved in cellular proliferation and suppressing apoptotic genes. YAP1 is a component in th ...
''. The response protects ''U. maydis'' from the host defense, and is necessary for the pathogen's virulence. Furthermore, ''U. maydis'' has a well-established recombinational
DNA repair
DNA repair is a collection of processes by which a cell identifies and corrects damage to the DNA molecules that encode its genome. In human cells, both normal metabolic activities and environmental factors such as radiation can cause DNA d ...
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 infection.
''
Cryptococcus neoformans
''Cryptococcus neoformans'' is an encapsulated yeast belonging to the class Tremellomycetes and an obligate aerobe that can live in both plants and animals. Its teleomorph is a filamentous fungus, formerly referred to ''Filobasidiella 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 macrophage
An alveolar macrophage, pulmonary macrophage, (or dust cell) is a type of macrophage, a professional phagocyte, found in the airways and at the level of the alveoli in the lungs, but separated from their walls.
Activity of the alveolar macropha ...
s. 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
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately ...
. 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
blastospore
A blastospore is an asexual fungal spore produced by budding. Produced by fungi within the phylum Glomeromycota and others. It is also known as a blastoconidium (plural = blastoconidia).
An example of a fungus that forms blastospores is '' ...
s. 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
Meiotic recombination protein DMC1/LIM15 homolog is a protein that in humans is encoded by the ''DMC1'' gene.
Meiotic recombination protein Dmc1 is a homolog of the bacterial strand exchange protein RecA. Dmc1 plays the central role in homologou ...
'', which is a conserved homologue of genes ''
recA
RecA is a 38 kilodalton protein essential for the repair and maintenance of DNA. A RecA structural and functional homolog has been found in every species in which one has been seriously sought and serves as an archetype for this class of homolog ...
'' in bacteria and ''
RAD51
DNA repair protein RAD51 homolog 1 is a protein encoded by the gene ''RAD51''. The enzyme encoded by this gene is a member of the RAD51 protein family which assists in repair of DNA double strand breaks. RAD51 family members are homologous to th ...
'' 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.
Human use
The human use of fungi for food preparation or preservation and other purposes is extensive and has a long history.
Mushroom farming
Fungiculture is the cultivation of fungi such as mushrooms. Cultivating fungi can yield foods (which include mostly mushrooms), medicine, construction materials and other products. A ''mushroom farm'' is involved in the business of growing fun ...
and
mushroom gathering
Mushroom hunting, mushrooming, mushroom picking, mushroom foraging, and similar terms describe the activity of gathering mushrooms in the wild, typically for culinary use. This practice is popular throughout most of Europe, Australia, Japan, K ...
are large industries in many countries. The study of the historical uses and sociological impact of fungi is known as
ethnomycology
Ethnomycology is the study of the historical uses and sociological impact of fungi and can be considered a subfield of ethnobotany or ethnobiology. Although in theory the term includes fungi used for such purposes as tinder, medicine (medicinal ...
. Because of the capacity of this group to produce an enormous range of
natural products
A natural product is a natural compound or substance produced by a living organism—that is, found in nature. In the broadest sense, natural products include any substance produced by life. Natural products can also be prepared by chemical sy ...
with
antimicrobial
An antimicrobial is an agent that kills microorganisms or stops their growth. Antimicrobial medicines can be grouped according to the microorganisms they act primarily against. For example, antibiotics are used against bacteria, and antifungals ar ...
or other biological activities, many species have long been used or are being developed for industrial
production of antibiotics
Production of antibiotics is a naturally occurring event, that thanks to advances in science can now be replicated and improved upon in laboratory settings. Due to the discovery of penicillin by Alexander Flemming, and the efforts of Florey and ...
, vitamins, and
anti-cancer
Cancer is a group of diseases involving abnormal cell growth with the potential to invade or spread to other parts of the body. These contrast with benign tumors, which do not spread. Possible signs and symptoms include a lump, abnormal ble ...
genetic engineering
Genetic engineering, also called genetic modification or genetic manipulation, is the modification and manipulation of an organism's genes using technology. It is a set of technologies used to change the genetic makeup of cells, including t ...
of fungi, enabling
metabolic engineering
Metabolic engineering is the practice of optimizing genetic and regulatory processes within cells to increase the cell's production of a certain substance. These processes are chemical networks that use a series of biochemical reactions and enz ...
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
A medication (also called medicament, medicine, pharmaceutical drug, medicinal drug or simply drug) is a drug used to diagnose, cure, treat, or prevent disease. Drug therapy (pharmacotherapy) is an important part of the medical field and re ...
production that are potentially more efficient than production by the original source organisms. Fungi-based industries are sometimes considered to be a major part of a growing
bioeconomy
Biobased economy, bioeconomy or biotechonomy is economic activity involving the use of biotechnology and biomass in the production of goods, services, or energy. The terms are widely used by regional development agencies, national and international ...
, with applications under
research and development
Research and development (R&D or R+D), known in Europe as research and technological development (RTD), is the set of innovative activities undertaken by corporations or governments in developing new services or products, and improving existi ...
including use for textiles,
meat
Meat is animal flesh that is eaten as food. Humans have hunted, farmed, and scavenged animals for meat since prehistoric times. The establishment of settlements in the Neolithic Revolution allowed the domestication of animals such as chic ...
substitution and general fungal biotechnology.
Therapeutic uses
Modern chemotherapeutics
Many species produce metabolites that are major sources of pharmacologically active drugs.
= Antibiotics
=
Particularly important are the antibiotics, including the
penicillin
Penicillins (P, PCN or PEN) are a group of β-lactam antibiotics originally obtained from ''Penicillium'' moulds, principally '' P. chrysogenum'' and '' P. rubens''. Most penicillins in clinical use are synthesised by P. chrysogenum using ...
s, a structurally related group of
β-lactam antibiotic
β-lactam antibiotics (beta-lactam antibiotics) are antibiotics that contain a beta-lactam ring in their chemical
structure. This includes penicillin derivatives (penams), cephalosporins and cephamycins (cephems), monobactams, carbapenems and ...
s that are synthesized from small
peptide
Peptides (, ) are short chains of amino acids linked by peptide bonds. Long chains of amino acids are called proteins. Chains of fewer than twenty amino acids are called oligopeptides, and include dipeptides, tripeptides, and tetrapeptides.
A ...
s. Although naturally occurring penicillins such as
penicillin G
Benzylpenicillin, also known as penicillin G (PenG) or BENPEN, and in military slang "Peanut Butter Shot" is an antibiotic used to treat a number of bacterial infections. This includes pneumonia, strep throat, syphilis, necrotizing enteroco ...
(produced by ''
Penicillium chrysogenum
''Penicillium chrysogenum'' (formerly known as ''Penicillium notatum'') is a species of fungus in the genus ''Penicillium''. It is common in temperate and subtropical regions and can be found on salted food products, but it is mostly found in in ...
'') have a relatively narrow spectrum of biological activity, a wide range of other penicillins can be produced by
chemical modification
Chemical modification refers to a number of various processes involving the alteration of the chemical constitution or structure of molecules.
In chemistry
Chemical modification describes the conversion of macromolecules through a chemical react ...
of the natural penicillins. Modern penicillins are
semisynthetic
Semisynthesis, or partial chemical synthesis, is a type of chemical synthesis that uses chemical compounds isolated from natural sources (such as microbial cell cultures or plant material) as the starting materials to produce novel compounds with ...
compounds, obtained initially from
fermentation
Fermentation is a metabolic process that produces chemical changes in organic substrates through the action of enzymes. In biochemistry, it is narrowly defined as the extraction of energy from carbohydrates in the absence of oxygen. In food ...
cultures, but then structurally altered for specific desirable properties. Other antibiotics produced by fungi include:
ciclosporin
Ciclosporin, also spelled cyclosporine and cyclosporin, is a calcineurin inhibitor, used as an immunosuppressant medication. It is a natural product. It is taken orally or intravenously for rheumatoid arthritis, psoriasis, Crohn's disease, ...
, commonly used as an
immunosuppressant
Immunosuppressive drugs, also known as immunosuppressive agents, immunosuppressants and antirejection medications, are drugs that inhibit or prevent activity of the immune system.
Classification
Immunosuppressive drugs can be classified in ...
during
transplant surgery
Organ transplantation is a medical procedure in which an organ is removed from one body and placed in the body of a recipient, to replace a damaged or missing organ. The donor and recipient may be at the same location, or organs may be transpor ...
tuberculosis
Tuberculosis (TB) is an infectious disease usually caused by '' Mycobacterium tuberculosis'' (MTB) bacteria. Tuberculosis generally affects the lungs, but it can also affect other parts of the body. Most infections show no symptoms, in ...
,
syphilis
Syphilis () is a sexually transmitted infection caused by the bacterium ''Treponema pallidum'' subspecies ''pallidum''. The signs and symptoms of syphilis vary depending in which of the four stages it presents (primary, secondary, latent, an ...
,
leprosy
Leprosy, also known as Hansen's disease (HD), is a long-term infection by the bacteria ''Mycobacterium leprae'' or ''Mycobacterium lepromatosis''. Infection can lead to damage of the nerves, respiratory tract, skin, and eyes. This nerve damag ...
, 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
The rhizosphere is the narrow region of soil or substrate that is directly influenced by root secretions and associated soil microorganisms known as the root microbiome. Soil pores in the rhizosphere can contain many bacteria and other microor ...
, and at low concentrations as
quorum-sensing
In biology, quorum sensing or quorum signalling (QS) is the ability to detect and respond to cell population density by gene regulation. As one example, QS enables bacteria to restrict the expression of specific genes to the high cell densities at ...
molecules for intra- or interspecies signaling.
= Other
=
Other drugs produced by fungi include
griseofulvin
Griseofulvin is an antifungal medication used to treat a number of types of dermatophytoses (ringworm). This includes fungal infections of the nails and scalp, as well as the skin when antifungal creams have not worked. It is taken by mouth.
C ...
isolated from ''
Penicillium griseofulvum
''Penicillium griseofulvum'' is a species of the genus of ''Penicillium'' which produces patulin, penifulvin A, cyclopiazonic acid, roquefortine C, shikimic acid, griseofulvin, and 6-Methylsalicylic acid (via a polyketide synthase
Polyketides ...
'', used to treat fungal infections, and
statin
Statins, also known as HMG-CoA reductase inhibitors, are a class of lipid-lowering medications that reduce illness and mortality in those who are at high risk of cardiovascular disease. They are the most common cholesterol-lowering drugs.
Low- ...
cholesterol synthesis
The mevalonate pathway, also known as the isoprenoid pathway or HMG-CoA reductase pathway is an essential metabolic pathway present in eukaryotes, archaea, and some bacteria. The pathway produces two five-carbon building blocks called isopenten ...
. Examples of statins found in fungi include
mevastatin
Mevastatin (compactin, ML-236B) is a hypolipidemic agent that belongs to the statins class.
It was isolated from the mold ''Penicillium citrinum'' by Akira Endo in the 1970s, and he identified it as a HMG-CoA reductase inhibitor, i.e., a statin. ...
from ''
Penicillium citrinum
''Penicillium citrinum'' is an anamorph, mesophilic fungus species of the genus of ''Penicillium'' which produces tanzawaic acid A-D, ACC, Mevastatin, Quinocitrinine A, Quinocitrinine B, and nephrotoxic citrinin. ''Penicillium citrinum'' ...
'' and
lovastatin
Lovastatin, sold under the brand name Mevacor among others, is a statin medication, to treat hypercholesterolemia, high blood cholesterol and reduce the risk of cardiovascular disease. Its use is recommended together with lifestyle changes. It ...
from ''
Aspergillus terreus
''Aspergillus terreus'', also known as ''Aspergillus terrestris'', is a fungus (mold) found worldwide in soil. Although thought to be strictly asexual until recently, ''A. terreus'' is now known to be capable of sexual reproduction. This saprotr ...
'' and the
oyster mushroom
''Pleurotus'' is a genus of gilled mushrooms which includes one of the most widely eaten mushrooms, '' P. ostreatus''. Species of ''Pleurotus'' may be called oyster, abalone, or tree mushrooms, and are some of the most commonly cultivated edib ...
.
Psilocybin
Psilocybin ( , ) is a naturally occurring psychedelic prodrug compound produced by more than 200 species of fungi. The most potent are members of the genus ''Psilocybe'', such as '' P. azurescens'', '' P. semilanceata'', and '' P.&nbs ...
from
fungi
A fungus ( : 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, separately from ...
network
Network, networking and networked may refer to:
Science and technology
* Network theory, the study of graphs as a representation of relations between discrete objects
* Network science, an academic field that studies complex networks
Mathematics
...
integration
Integration may refer to:
Biology
*Multisensory integration
*Path integration
* Pre-integration complex, viral genetic material used to insert a viral genome into a host genome
*DNA integration, by means of site-specific recombinase technology, ...
. Fungi produce compounds that inhibit
virus
A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea.
Since Dmitri Ivanovsky's 1 ...
es and cancer cells. Specific metabolites, such as
polysaccharide-K
Polysaccharide-K (Krestin, PSK) is a protein-bound polysaccharide isolated from the fruitbody of ''Trametes versicolor''.
Research summary
PSK has been studied in patients with gastric cancer, breast cancer, colorectal cancer, and lung cancer. T ...
,
ergotamine
Ergotamine, sold under the brand names Cafergot (with caffeine) and Ergomar among others, is an ergopeptine and part of the ergot family of alkaloids; it is structurally and biochemically closely related to ergoline. It possesses structural simil ...
, and
β-lactam antibiotics
β-lactam antibiotics (beta-lactam antibiotics) are antibiotics that contain a beta-lactam ring in their chemical
structure. This includes penicillin derivatives (penams), cephalosporins and cephamycins (cephems), monobactams, carbapenems and ...
, are routinely used in clinical medicine. The
shiitake
The shiitake (alternate form shitake) (; ''Lentinula edodes'') is an edible mushroom native to East Asia, which is now cultivated and consumed around the globe. It is considered a Medicinal fungi, medicinal mushroom in some forms of tradition ...
mushroom is a source of
lentinan
Lentinan is a polysaccharide isolated from the fruit body of shiitake mushroom (''Lentinula edodes'' mycelium).
Chemistry
Lentinan is a β-1,3 beta-glucan with β-1,6 branching. It has a molecular weight of 500,000 Da and specific rotation of +1 ...
, a clinical drug approved for use in cancer treatments in several countries, including
Japan
Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
. In Europe and Japan,
polysaccharide-K
Polysaccharide-K (Krestin, PSK) is a protein-bound polysaccharide isolated from the fruitbody of ''Trametes versicolor''.
Research summary
PSK has been studied in patients with gastric cancer, breast cancer, colorectal cancer, and lung cancer. T ...
(brand name Krestin), a chemical derived from ''
Trametes versicolor
''Trametes versicolor''also known as ''Coriolus versicolor'' and ''Polyporus versicolor''is a common polypore mushroom found throughout the world. Meaning 'of several colors', ''versicolor'' reliably describes this fungus that displays a varie ...
'', is an approved
adjuvant In pharmacology, an adjuvant is a drug or other substance, or a combination of substances, that is used to increase the efficacy or potency of certain drugs. Specifically, the term can refer to:
* Adjuvant therapy in cancer management
* Analgesic ...
traditional Chinese medicine
Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of action ...
. Mushrooms with a history of such use include ''
Agaricus subrufescens
''Agaricus subrufescens'' (syn. ''Agaricus blazei'', ''Agaricus brasiliensis'' or ''Agaricus rufotegulis'') is a species of mushroom, commonly known as almond mushroom, mushroom of the sun, God's mushroom, mushroom of life, royal sun agaricus, '' ...
Ophiocordyceps sinensis
''Ophiocordyceps sinensis'' (formerly known as ''Cordyceps sinensis''), known colloquially as caterpillar fungus, is an entomopathogenic fungus (a fungus that grows on insects) in the family Ophiocordycipitaceae. It is mainly found in the mead ...
''.
Cultured foods
Baker's yeast
Baker's yeast is the common name for the strains of yeast commonly used in baking bread and other bakery products, serving as a leavening agent which causes the bread to rise (expand and become lighter and softer) by converting the fermentable ...
or ''
Saccharomyces cerevisiae
''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungus microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have been o ...
'', a unicellular fungus, is used to make
bread
Bread is a staple food prepared from a dough of flour (usually wheat) and water, usually by baking. Throughout recorded history and around the world, it has been an important part of many cultures' diet. It is one of the oldest human-made f ...
and other wheat-based products, such as pizza dough and dumplings. Yeast species of the genus ''
Saccharomyces
''Saccharomyces'' is a genus of fungi that includes many species of yeasts. ''Saccharomyces'' is from Greek σάκχαρον (sugar) and μύκης (fungus) and means ''sugar fungus''. Many members of this genus are considered very important in f ...
'' are also used to produce alcoholic beverages through fermentation. Shoyu koji mold (''Aspergillus oryzae'') is an essential ingredient in brewing Shoyu (
soy sauce
Soy sauce (also called simply soy in American English and soya sauce in British English) is a liquid condiment of Chinese origin, traditionally made from a fermented paste of soybeans, roasted grain, brine, and ''Aspergillus oryzae'' or '' As ...
) and sake, and the preparation of miso, while '' Rhizopus'' species are used for making tempeh. Several of these fungi are Domestication, domesticated species that were Breeding program, bred or selected according to their capacity to ferment food without producing harmful mycotoxins (see below), which are produced by very closely related ''Aspergillus flavus, Aspergilli''. Quorn (food product), Quorn, a Meat analogue, meat substitute, is made from ''Fusarium venenatum''.
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.).
Many other mushroom species are Mushroom hunting, harvested from the wild for personal consumption or commercial sale. Lactarius deliciosus, Milk mushrooms,
morel
''Morchella'', the true morels, is a genus of edible sac fungi closely related to anatomically simpler cup fungi in the order Pezizales ( division Ascomycota). These distinctive fungi have a honeycomb appearance due to the network of ridges wi ...
s, chanterelles,
truffles
A truffle is the fruiting body of a subterranean ascomycete fungus, predominantly one of the many species of the genus ''Tuber''. In addition to ''Tuber'', many other genera of fungi are classified as truffles including '' Geopora'', '' Pe ...
, Craterellus, 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 cheese, blue color in cheeses such as Stilton cheese, Stilton or Roquefort, which are made by inoculation with ''Penicillium roqueforti''. 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.
Poisonous fungi
Many mushroom species are Mushroom poisoning, poisonous to humans and cause a range of reactions including slight digestive problems, allergy, allergic reactions, hallucinations, severe organ failure, and death. Genera with mushrooms containing deadly toxins include ''Conocybe'', ''Galerina'', ''Lepiota'' and the most infamous, ''
Amanita
The genus ''Amanita'' contains about 600 species of agarics, including some of the most toxic known mushrooms found worldwide, as well as some well-regarded edible species. This genus is responsible for approximately 95% of the fatalities resul ...
''. The latter genus includes the destroying angel ''(Amanita virosa, A.virosa)'' and the death cap ''(Amanita phalloides, 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. Amanita muscaria, Fly agaric mushrooms (''Amanita muscaria'') also cause occasional non-fatal poisonings, mostly as a result of ingestion for its Psychedelics, dissociatives and deliriants, hallucinogenic properties. Historically, fly agaric was used by different peoples in Europe and Asia and its present usage for religious or shamanism, shamanic purposes is reported from some ethnic groups such as the Koryaks, Koryak people of northeastern Siberia.
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.Hall, p. 7.
Pest control
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 parasitism, parasites of these pathogens. For example, certain species eliminate or suppress the growth of harmful plant pathogens, such as insects, mites, weeds,
nematodes
The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a broa ...
, 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'', ''Metarhizium'' spp., ''Hirsutella'' spp., ''Paecilomyces'' (''Isaria'') spp., and ''Lecanicillium lecanii''. Endophytic fungi of grasses of the genus ''Epichloë'', such as ''Epichloë coenophiala, E. coenophiala'', 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 ''Epichloë'' endophytes is one approach being used in plant breeding, 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 livestock.
Bioremediation
Certain fungi, in particular white rot, 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 biomineralization, biomineralize uranium#Oxides, uranium oxides, suggesting they may have application in the bioremediation of radioactively polluted sites.
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. Other important model fungi are ''Aspergillus nidulans'' and the yeasts ''
Saccharomyces cerevisiae
''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungus microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have been o ...
'' and ''Schizosaccharomyces pombe'', each of which with a long history of use to investigate issues in eukaryotic cell biology and
genetics
Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar work ...
, such as cell cycle regulation, chromatin structure, and gene regulation. Other fungal models have 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
''Magnaporthe grisea'', also known as rice blast fungus, rice rotten neck, rice seedling blight, blast of rice, oval leaf spot of graminea, pitting disease, ryegrass blast, Johnson spot, neck blast, wheat blast, and Imochi ( Japanese:稲熱) is ...
'', a plant pathogen, and ''Pichia pastoris'', a yeast widely used for eukaryotic protein production.
Others
Fungi are used extensively to produce industrial chemicals like citric acid, citric, gluconic acid, gluconic, lactic acid, lactic, and malic acid, 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.
See also
* Conservation of fungi
* Fantastic Fungi
* Glossary of fungi
* Marine fungi
* Mycosis
* Outline of fungi
References
Citations
{{Reflist, refs=
{{cite journal , vauthors=Aanen DK , title=As you reap, so shall you sow: coupling of harvesting and inoculating stabilizes the mutualism between termites and fungi , journal=Biology Letters , volume=2 , issue=2 , pages=209–12 , date=June 2006 , pmid=17148364 , pmc=1618886 , doi=10.1098/rsbl.2005.0424{{cite journal , vauthors=Abe K, Gomi K, Hasegawa F, Machida M , s2cid=36874528 , title=Impact of ''Aspergillus oryzae'' genomics on industrial production of metabolites , journal=Mycopathologia , volume=162 , issue=3 , pages=143–53 , date=September 2006 , pmid=16944282 , doi=10.1007/s11046-006-0049-2{{cite book , vauthors=Alcamo IE, Pommerville J , title=Alcamo's Fundamentals of Microbiology , url=https://archive.org/details/alcamosfundament0000pomm , url-access=registration , publisher=Jones and Bartlett , location=Boston, Massachusetts , year=2004 , pag 590 , isbn=978-0-7637-0067-6{{cite book , vauthors=Ammirati JF, McKenny M, Stuntz DE , title=The New Savory Wild Mushroom , publisher=University of Washington Press , location=Seattle, Washington , year=1987 , pages=xii–xiii , isbn=978-0-295-96480-5{{cite journal , last1=Aramayo , first1=Rodolfo , last2=Selker , first2=Erik U. , title=''Neurospora crassa'', a model system for epigenetics research , journal=Cold Spring Harbor Perspectives in Biology , volume=5 , issue=10 , year=2013 , pages=a017921 , doi=10.1101/cshperspect.a017921 , pmc=3783048 , pmid=24086046{{cite journal , vauthors=Arnold AE, Mejía LC, Kyllo D, Rojas EI, Maynard Z, Robbins N, Herre EA , title=Fungal endophytes limit pathogen damage in a tropical tree , journal=Proceedings of the National Academy of Sciences of the United States of America , volume=100 , issue=26 , pages=15649–54 , date=December 2003 , pmid=14671327 , pmc=307622 , doi=10.1073/pnas.2533483100 , bibcode=2003PNAS..10015649A , doi-access=free{{cite journal , last1=Baghban , first1=Roghayyeh , last2=Farajnia , first2=Safar , last3=Rajabibazl , first3=Masoumeh , last4=Ghasemi , first4=Younes , last5=Mafi , first5=AmirAli , last6=Hoseinpoor , first6=Reyhaneh , last7=Rahbarnia , first7=Leila , last8=Aria , first8=Maryam , title=Yeast expression systems: Overview and recent advances , journal=Molecular Biotechnology , volume=61 , issue=5 , year=2019 , pages=365–384 , doi=10.1007/s12033-019-00164-8 , pmid=30805909 , s2cid=73501127{{cite journal , vauthors=Baldauf SL, Palmer JD , title=Animals and fungi are each other's closest relatives: congruent evidence from multiple proteins , journal=Proceedings of the National Academy of Sciences of the United States of America , volume=90 , issue=24 , pages=11558–62 , date=December 1993 , pmid=8265589 , pmc=48023 , doi=10.1073/pnas.90.24.11558 , bibcode=1993PNAS...9011558B , doi-access=free{{cite journal , vauthors=Barea JM, Pozo MJ, Azcón R, Azcón-Aguilar C , title=Microbial co-operation in the rhizosphere , journal=Journal of Experimental Botany , volume=56 , issue=417 , pages=1761–78 , date=July 2005 , pmid=15911555 , doi=10.1093/jxb/eri197 , doi-access=free{{cite news , title=Fungi to fight 'toxic war zones' , date=5 May 2008 , url=http://news.bbc.co.uk/2/hi/uk_news/scotland/tayside_and_central/7384500.stm , work=BBC News , access-date=12 May 2008 , archive-url=https://web.archive.org/web/20170915195952/http://news.bbc.co.uk/2/hi/uk_news/scotland/tayside_and_central/7384500.stm , archive-date=15 September 2017 , url-status=live{{cite journal , vauthors=Beadle GW, Tatum EL , title=Genetic Control of Biochemical Reactions in Neurospora , journal=Proceedings of the National Academy of Sciences of the United States of America , volume=27 , issue=11 , pages=499–506 , date=November 1941 , pmid=16588492 , pmc=1078370 , doi=10.1073/pnas.27.11.499 , bibcode=1941PNAS...27..499B , doi-access=free{{cite book , vauthors=Blackwell M, Spatafora JW , veditors=Bills GF, Mueller GM, Foster MS , chapter=Fungi and their allies , title=Biodiversity of Fungi: Inventory and Monitoring Methods , publisher=Elsevier Academic Press , location=Amsterdam , year=2004 , pages=18–20 , isbn=978-0-12-509551-8{{cite journal , vauthors=Bonfante P , title=Plants, mycorrhizal fungi and endobacteria: a dialog among cells and genomes , journal=The Biological Bulletin , volume=204 , issue=2 , pages=215–20 , date=April 2003 , pmid=12700157 , doi=10.2307/1543562 , jstor=1543562 , s2cid=12377410 , url=https://www.biodiversitylibrary.org/part/9240{{cite journal, vauthors=Bouton JH, Latch GC, Hill NS, Hoveland CS, McCann MA, Watson RH, Parish JA, Hawkins LL, Thompson FN , year=2002 , title=Reinfection of Tall Fescue Cultivars with Non-Ergot Alkaloid–Producing Endophytes , journal=Agronomy Journal , volume=94 , issue=3 , pages=567–574 , doi=10.2134/agronj2002.5670 , url=https://dl.sciencesocieties.org/publications/aj/pdfs/94/3/567{{cite journal , last1=Bozkurt , first1=Tolga O. , last2=Kamoun , first2=Sophien , last3=Lennon-Duménil , first3=Ana-Maria , title=The plant–pathogen haustorial interface at a glance , journal=Journal of Cell Science , volume=133 , issue=5 , year=2020 , doi=10.1242/jcs.237958 , pmc=7075074 , pmid=32132107{{Cite book , vauthors=Brakhage AA, Spröte P, Al-Abdallah Q, Gehrke A, Plattner H, Tüncher A , volume=88 , pages=45–90 , year=2004 , pmid=15719552 , doi=10.1007/b99257 , series=Advances in Biochemical Engineering/Biotechnology , isbn=978-3-540-22032-9 , title=Molecular Biotechnology of Fungal beta-Lactam Antibiotics and Related Peptide Synthetases , chapter=Regulation of Penicillin Biosynthesis in Filamentous Fungi{{cite journal , vauthors=Brakhage AA , title=Systemic fungal infections caused by Aspergillus species: epidemiology, infection process and virulence determinants , journal=Current Drug Targets , volume=6 , issue=8 , pages=875–86 , date=December 2005 , pmid=16375671 , doi=10.2174/138945005774912717{{cite book , vauthors=Brodie HJ , title=The Bird's Nest Fungi , publisher=University of Toronto Press , location=Toronto, Ontario , year=1975 , isbn=978-0-8020-5307-7 , page=80{{cite book , title=Lichens of North America , vauthors=Brodo IM, Sharnoff SD , year=2001 , publisher=Yale University Press , location=New Haven, Connecticut , isbn=978-0-300-08249-4{{cite journal , vauthors=Brundrett MC , year=2002 , title=Coevolution of roots and mycorrhizas of land plants , journal=New Phytologist , volume=154 , issue=2 , pages=275–304 , doi=10.1046/j.1469-8137.2002.00397.x, pmid=33873429 , doi-access=free{{cite journal , vauthors=Bruns T , s2cid=648881 , title=Evolutionary biology: a kingdom revised , journal=Nature (journal), Nature , volume=443 , issue=7113 , pages=758–61 , date=October 2006 , pmid=17051197 , doi=10.1038/443758a , bibcode=2006Natur.443..758B , doi-access=free{{cite journal , vauthors=Celio GJ, Padamsee M, Dentinger BT, Bauer R, McLaughlin DJ , s2cid=23123595 , title=Assembling the Fungal Tree of Life: constructing the structural and biochemical database , journal=Mycologia , volume=98 , issue=6 , pages=850–9 , year=2006 , pmid=17486962 , doi=10.3852/mycologia.98.6.850{{cite book , title=Mushrooms: Cultivation, Nutritional Value, Medicinal Effect and Environmental Impact , vauthors=((Chang S-T)), Miles PG , year=2004 , publisher=CRC Press , location=Boca Raton, Florida , isbn=978-0-8493-1043-0{{cite journal , display-authors=6 , last1=Cheek , first1=Martin , last2=Nic Lughadha , first2=Eimear , last3=Kirk , first3=Paul , last4=Lindon , first4=Heather , last5=Carretero , first5=Julia , last6=Looney , first6=Brian , last7=Douglas , first7=Brian , last8=Haelewaters , first8=Danny , last9=Gaya , first9=Ester , last10=Llewellyn , first10=Theo , last11=Ainsworth , first11=A. Martyn , last12=Gafforov , first12=Yusufjon , last13=Hyde , first13=Kevin , last14=Crous , first14=Pedro , last15=Hughes , first15=Mark , last16=Walker , first16=Barnaby E. , last17=Campostrini Forzza , first17=Rafaela , last18=Wong , first18=Khoon Meng , last19=Niskanen , first19=Tuula , title=New scientific discoveries: Plants and fungi , journal=Plants, People, Planet , volume=2 , issue=5 , year=2020 , pages=371–388 , doi=10.1002/ppp3.10148 , doi-access=free{{cite journal , vauthors=Christensen MJ, Bennett RJ, Ansari HA, Koga H, Johnson RD, Bryan GT, Simpson WR, Koolaard JP, Nickless EM, Voisey CR , title=Epichloë endophytes grow by intercalary hyphal extension in elongating grass leaves , journal=Fungal Genetics and Biology , volume=45 , issue=2 , pages=84–93 , date=February 2008 , pmid=17919950 , doi=10.1016/j.fgb.2007.07.013{{cite journal , vauthors=Clay K, Schardl C , title=Evolutionary origins and ecological consequences of endophyte symbiosis with grasses , journal=The American Naturalist , volume=160 Suppl 4 , issue=suppl. 4 , pages=S99–S127 , date=October 2002 , pmid=18707456 , doi=10.1086/342161 , s2cid=23909652{{cite book , editor-last1=Lacey , editor-first1=Lawrence A. , last1=Chandler , first1=D. , chapter=Basic and Applied Research on Entomopathogenic Fungi , year=2017 , title=Microbial Control of Insect and Mite Pests , pages=69–89 , doi=10.1016/B978-0-12-803527-6.00005-6 , publisher=Academic Press , isbn=978-0-12-803527-6{{cite journal , last1=Chomicki , first1=Guillaume , last2=Renner , first2=Susanne S. , title=The interactions of ants with their biotic environment , journal=Proceedings of the Royal Society B: Biological Sciences , volume=284 , issue=1850 , year=2017 , pages=20170013 , doi=10.1098/rspb.2017.0013 , pmc=5360932 , pmid=28298352{{cite journal , vauthors=Cushion MT, Smulian AG, Slaven BE, Sesterhenn T, Arnold J, Staben C, Porollo A, Adamczak R, Meller J , title=Transcriptome of ''Pneumocystis carinii'' during fulminate infection: carbohydrate metabolism and the concept of a compatible parasite , journal=PLOS ONE , volume=2 , issue=5 , pages=e423 , year=2007 , pmid=17487271 , pmc=1855432 , doi=10.1371/journal.pone.0000423 , doi-access=free , bibcode=2007PLoSO...2..423C{{cite book , vauthors=Cook GC, Zumla AI , author-link2=Alimuddin Zumla , title=Manson's Tropical Diseases: Expert Consult , publisher=Saunders Ltd , location=Edinburgh, Scotland , year=2008 , page=347 , isbn=978-1-4160-4470-3{{cite journal , vauthors=Dadachova E, Bryan RA, Huang X, Moadel T, Schweitzer AD, Aisen P, Nosanchuk JD, Casadevall A , title=Ionizing radiation changes the electronic properties of melanin and enhances the growth of melanized fungi , journal=PLOS ONE , volume=2 , issue=5 , pages=e457 , year=2007 , pmid=17520016 , pmc=1866175 , doi=10.1371/journal.pone.0000457 , doi-access=free , bibcode=2007PLoSO...2..457D{{cite journal , vauthors=Dadachova E, Casadevall A , title=Ionizing radiation: how fungi cope, adapt, and exploit with the help of melanin , journal=Current Opinion in Microbiology , volume=11 , issue=6 , pages=525–31 , date=December 2008 , pmid=18848901 , pmc=2677413 , doi=10.1016/j.mib.2008.09.013{{cite journal , vauthors=Daniels KJ, Srikantha T, Lockhart SR, Pujol C, Soll DR , title=Opaque cells signal white cells to form biofilms in ''Candida albicans'' , journal=The EMBO Journal , volume=25 , issue=10 , pages=2240–52 , date=May 2006 , pmid=16628217 , pmc=1462973 , doi=10.1038/sj.emboj.7601099{{Cite book , vauthors=Datta A, Ganesan K, Natarajan K , title=Current trends in ''Candida albicans'' research , volume=30 , pages=53–88 , year=1989 , pmid=2700541 , doi=10.1016/S0065-2911(08)60110-1 , isbn=978-0-12-027730-8 , series=Advances in Microbial Physiology , chapter=Current Trends in Candida albicans Research{{cite journal , vauthors=Dean RA, Talbot NJ, Ebbole DJ, Farman ML, Mitchell TK, Orbach MJ, Thon M, Kulkarni R, Xu JR, Pan H, Read ND, Lee YH, Carbone I, Brown D, Oh YY, Donofrio N, Jeong JS, Soanes DM, Djonovic S, Kolomiets E, Rehmeyer C, Li W, Harding M, Kim S, Lebrun MH, Bohnert H, Coughlan S, Butler J, Calvo S, Ma LJ, Nicol R, Purcell S, Nusbaum C, Galagan JE, Birren BW , display-authors=6 , title=The genome sequence of the rice blast fungus ''Magnaporthe grisea'' , journal=Nature (journal), Nature , volume=434 , issue=7036 , pages=980–6 , date=April 2005 , pmid=15846337 , doi=10.1038/nature03449 , bibcode=2005Natur.434..980D , doi-access=free{{cite journal , vauthors=Dennis RL , year=1970 , title=A Middle Pennsylvanian basidiomycete mycelium with clamp connections , journal=Mycologia , volume=62 , issue=3 , pages=578–584 , doi=10.2307/3757529 , jstor=3757529 , url=http://www.cybertruffle.org.uk/cyberliber/59350/0062/003/0578.htm , access-date=5 July 2011 , archive-url=https://web.archive.org/web/20180929141016/http://www.cybertruffle.org.uk/cyberliber/59350/0062/003/0578.htm , archive-date=29 September 2018 , url-status=live{{Cite book , vauthors=Demain AL, Fang A , volume=69 , pages=1–39 , year=2000 , pmid=11036689 , doi=10.1007/3-540-44964-7_1 , isbn=978-3-540-67793-2 , series=Advances in Biochemical Engineering/Biotechnology , title=History of Modern Biotechnology I , chapter=The Natural Functions of Secondary Metabolites{{cite journal , vauthors=Desjardin DE, Perry BA, Lodge DJ, Stevani CV, Nagasawa E , s2cid=25377671 , title=Luminescent Mycena: new and noteworthy species , journal=Mycologia , volume=102 , issue=2 , pages=459–77 , year=2010 , pmid=20361513 , doi=10.3852/09-197 , url=http://producao.usp.br/handle/BDPI/16784 , access-date=11 November 2018 , archive-url=https://web.archive.org/web/20181111043819/http://www.producao.usp.br/handle/BDPI/16784 , archive-date=11 November 2018 , url-status=live{{cite journal , vauthors=Deshpande MV , title=Mycopesticide production by fermentation: potential and challenges , journal=Critical Reviews in Microbiology , volume=25 , issue=3 , pages=229–43 , year=1999 , pmid=10524330 , doi=10.1080/10408419991299220{{cite book , author1=Donoghue MJ, author2=Cracraft J, author-link2=Joel Cracraft , title=Assembling the Tree of Life , publisher=Oxford University Press , location=Oxford (Oxfordshire), UK , year=2004 , page=187 , isbn=978-0-19-517234-8{{cite journal , vauthors=Dotzler N, Walker C, Krings M, Hass H, Kerp H, Taylor TN, Agerer R , s2cid=1746303 , year=2009 , title=''Acaulosporoid glomeromycotan'' spores with a germination shield from the 400-million-year-old Rhynie chert , journal=Mycological Progress , volume=8 , issue=1 , pages=9–18 , doi=10.1007/s11557-008-0573-1, hdl=1808/13680 , url=https://kuscholarworks.ku.edu/bitstream/1808/13680/1/Taylor_et_al_2009.pdf , hdl-access=free{{cite journal , vauthors=El Dine RS, El Halawany AM, Ma CM, Hattori M , title=Anti-HIV-1 protease activity of lanostane triterpenes from the Vietnamese mushroom ''Ganoderma colossum'' , journal=Journal of Natural Products , volume=71 , issue=6 , pages=1022–6 , date=June 2008 , pmid=18547117 , doi=10.1021/np8001139{{cite journal , vauthors=el-Mekkawy S, Meselhy MR, Nakamura N, Tezuka Y, Hattori M, Kakiuchi N, Shimotohno K, Kawahata T, Otake T , title=Anti-HIV-1 and anti-HIV-1-protease substances from ''Ganoderma lucidum'' , journal=Phytochemistry , volume=49 , issue=6 , pages=1651–7 , date=November 1998 , pmid=9862140 , doi=10.1016/S0031-9422(98)00254-4{{cite journal , vauthors=Erdogan A, Gurses M, Sert S , title=Isolation of moulds capable of producing mycotoxins from blue mouldy Tulum cheeses produced in Turkey , journal=International Journal of Food Microbiology , volume=85 , issue=1–2 , pages=83–5 , date=August 2003 , pmid=12810273 , doi=10.1016/S0168-1605(02)00485-3{{cite journal , vauthors=Eshet Y, Rampino MR, Visscher H , s2cid=58937537 , year=1995 , title=Fungal event and palynological record of ecological crisis and recovery across the Permian-Triassic boundary , journal=Geology (journal), Geology , volume=23 , issue=1 , pages=967–970 , doi=10.1130/0091-7613(1995)023<0967:FEAPRO>2.3.CO;2, bibcode=1995Geo....23..967E{{cite journal , vauthors=Fan W, Kraus PR, Boily MJ, Heitman J , title=''Cryptococcus neoformans'' gene expression during murine macrophage infection , journal=Eukaryotic Cell , volume=4 , issue=8 , pages=1420–33 , date=August 2005 , pmid=16087747 , pmc=1214536 , doi=10.1128/EC.4.8.1420-1433.2005{{cite journal , vauthors=Farrar JF , title=Carbohydrate metabolism in biotrophic plant pathogens , journal=Microbiological Sciences , volume=2 , issue=10 , pages=314–7 , date=October 1985 , pmid=3939987{{cite journal , vauthors=Fajardo A, Martínez JL , title=Antibiotics as signals that trigger specific bacterial responses , journal=Current Opinion in Microbiology , volume=11 , issue=2 , pages=161–7 , date=April 2008 , pmid=18373943 , doi=10.1016/j.mib.2008.02.006{{cite journal , vauthors=Ferguson BA, Dreisbach TA, Parks CG, Filip GM, Schmitt CL , year=2003 , title=Coarse-scale population structure of pathogenic ''Armillaria'' species in a mixed-conifer forest in the Blue Mountains of northeast Oregon , journal=Canadian Journal of Forest Research , volume=33 , issue=4 , pages=612–623 , doi=10.1139/x03-065 , url=https://zenodo.org/record/1235905 , access-date=3 July 2019 , archive-url=https://web.archive.org/web/20190703164318/https://zenodo.org/record/1235905 , archive-date=3 July 2019 , url-status=live{{cite journal , last1=Fernandez , first1=Jessie , last2=Orth , first2=Kim , title=Rise of a cereal killer: the biology of ''Magnaporthe oryzae'' biotrophic growth , journal=Trends in Microbiology , volume=26 , issue=7 , year=2018 , pages=582–597 , doi=10.1016/j.tim.2017.12.007 , pmid=29395728 , pmc=6003838{{cite journal , vauthors=Fincham JR , title=Transformation in fungi , journal=Microbiological Reviews , volume=53 , issue=1 , pages=148–70 , date=March 1989 , pmid=2651864 , pmc=372721 , doi=10.1128/MMBR.53.1.148-170.1989{{cite journal , vauthors=Firenzuoli F, Gori L, Lombardo G , title=The Medicinal Mushroom ''Agaricus blazei'' Murrill: Review of Literature and Pharmaco-Toxicological Problems , journal=Evidence-Based Complementary and Alternative Medicine , volume=5 , issue=1 , pages=3–15 , date=March 2008 , pmid=18317543 , pmc=2249742 , doi=10.1093/ecam/nem007{{cite journal , last1=Fisher , first1=Matthew C. , last2=Garner , first2=Trenton W. J. , title=Chytrid fungi and global amphibian declines , journal=Nature Reviews Microbiology , volume=18 , issue=6 , year=2020 , pages=332–343 , doi=10.1038/s41579-020-0335-x , pmid=32099078 , hdl=10044/1/78596 , s2cid=211266075 , url=https://discovery.ucl.ac.uk/id/eprint/10092667/ , hdl-access=free{{cite journal , last1=Fritz , first1=Heidi , last2=Kennedy , first2=Deborah A. , last3=Ishii , first3=Mami , last4=Fergusson , first4=Dean , last5=Fernandes , first5=Rochelle , last6=Cooley , first6=Kieran , last7=Seely , first7=Dugald , title=Polysaccharide K and ''Coriolus versicolor'' extracts for lung cancer , journal=Integrative Cancer Therapies , volume=14 , issue=3 , year=2015 , pages=201–211 , doi=10.1177/1534735415572883 , pmid=25784670 , doi-access=free{{cite journal , vauthors=Fischer R, Zekert N, Takeshita N , s2cid=205365895 , title=Polarized growth in fungi--interplay between the cytoskeleton, positional markers and membrane domains , journal=Molecular Microbiology , volume=68 , issue=4 , pages=813–26 , date=May 2008 , pmid=18399939 , doi=10.1111/j.1365-2958.2008.06193.x{{cite journal , vauthors=Fomina M, Charnock JM, Hillier S, Alvarez R, Gadd GM, author5-link=Geoffrey Michael Gadd , title=Fungal transformations of uranium oxides , journal=Environmental Microbiology , volume=9 , issue=7 , pages=1696–710 , date=July 2007 , pmid=17564604 , doi=10.1111/j.1462-2920.2007.01288.x{{cite journal , vauthors=Foster CB, Stephenson MH, Marshall C, Logan GA, Greenwood PF , year=2002 , title=A revision of Reduviasporonites Wilson 1962: description, illustration, comparison and biological affinities , journal=Palynology , volume=26 , issue=1 , pages=35–58 , doi=10.2113/0260035{{cite journal , vauthors=Fomina M, Charnock JM, Hillier S, Alvarez R, Livens F, Gadd GM , s2cid=52805144 , title=Role of fungi in the biogeochemical fate of depleted uranium , journal=Current Biology , volume=18 , issue=9 , pages=R375–7 , date=May 2008 , pmid=18460315 , doi=10.1016/j.cub.2008.03.011 , doi-access=free{{cite journal , vauthors=Furlaneto MC, Pizzirani-Kleiner AA , title=Intraspecific hybridisation of ''Trichoderma pseudokoningii'' by anastomosis and by protoplast fusion , journal=FEMS Microbiology Letters , volume=69 , issue=2 , pages=191–5 , date=January 1992 , pmid=1537549 , doi=10.1111/j.1574-6968.1992.tb05150.x , doi-access=free{{cite journal , vauthors=Gadd GM , title=Geomycology: biogeochemical transformations of rocks, minerals, metals and radionuclides by fungi, bioweathering and bioremediation , journal=Mycological Research , volume=111 , issue=Pt 1 , pages=3–49 , date=January 2007 , pmid=17307120 , doi=10.1016/j.mycres.2006.12.001{{cite journal , last1=Garrido-Benavent , first1=Isaac , last2=Pérez-Ortega , first2=Sergio , title=Past, present, and future research in bipolar lichen-forming fungi and their photobionts , journal=American Journal of Botany , volume=104 , issue=11 , year=2017 , pages=1660–1674 , doi=10.3732/ajb.1700182 , doi-access=free{{cite journal , last1=Gow , first1=Neil A. R. , last2=Latge , first2=Jean-Paul , last3=Munro , first3=Carol A. , last4=Heitman , first4=Joseph , title=The fungal cell wall: Structure, biosynthesis, and function , journal=Microbiology Spectrum , volume=5 , issue=3 , year=2017 , doi=10.1128/microbiolspec.FUNK-0035-2016 , pmid=28513415 , hdl=2164/8941{{cite journal , vauthors=Guarro J, Stchigel AM , title=Developments in fungal taxonomy , journal=Clinical Microbiology Reviews , volume=12 , issue=3 , pages=454–500 , date=July 1999 , pmid=10398676 , pmc=100249 , doi=10.1128/CMR.12.3.454{{cite journal , last1=Guerre , first1=Philippe , title=Ergot alkaloids produced by endophytic fungi of the genus ''Epichloë'' , journal=Toxins , volume=7 , issue=3 , year=2015 , pages=773–790 , doi=10.3390/toxins7030773 , pmid=25756954 , pmc=4379524 , doi-access=free{{cite journal , vauthors=Hachmeister KA, Fung DY , title=Tempeh: a mold-modified indigenous fermented food made from soybeans and/or cereal grains , journal=Critical Reviews in Microbiology , volume=19 , issue=3 , pages=137–88 , year=1993 , pmid=8267862 , doi=10.3109/10408419309113527{{cite journal , last1=Han , first1=Bing , last2=Weiss , first2=Louis M. , last3=Heitman , first3=Joseph , last4=Stukenbrock , first4=Eva Holtgrewe , title=Microsporidia: Obligate intracellular pathogens within the fungal kingdom , journal=Microbiology Spectrum , volume=5 , issue=2 , year=2017 , doi=10.1128/microbiolspec.FUNK-0018-2016 , pmid=28944750 , pmc=5613672{{cite journal , vauthors=Harris SD , s2cid=2147525 , title=Branching of fungal hyphae: regulation, mechanisms and comparison with other branching systems , journal=Mycologia , volume=100 , issue=6 , pages=823–32 , year=2008 , pmid=19202837 , doi=10.3852/08-177 , url=http://www.cybertruffle.org.uk/cyberliber/59350/0100/006/0823.htm , access-date=5 July 2011 , archive-url=https://web.archive.org/web/20160412145515/http://www.cybertruffle.org.uk/cyberliber/59350/0100/006/0823.htm , archive-date=12 April 2016 , url-status=live{{cite book , vauthors=Halpern GM, Miller A , title=Medicinal Mushrooms: Ancient Remedies for Modern Ailments , publisher=M. Evans and Co , location=New York, New York , year=2002 , page=116 , isbn=978-0-87131-981-4{{cite journal , vauthors=Hawksworth DL , title=Pandora's mycological box: molecular sequences vs. morphology in understanding fungal relationships and biodiversity , journal=Revista Iberoamericana de Micología , volume=23 , issue=3 , pages=127–33 , date=September 2006 , pmid=17196017 , doi=10.1016/S1130-1406(06)70031-6{{cite journal , vauthors=van der Heijden MG, Streitwolf-Engel R, Riedl R, Siegrist S, Neudecker A, Ineichen K, Boller T, Wiemken A, Sanders IR , s2cid=17048094 , title=The mycorrhizal contribution to plant productivity, plant nutrition and soil structure in experimental grassland , journal=The New Phytologist , volume=172 , issue=4 , pages=739–52 , year=2006 , pmid=17096799 , doi=10.1111/j.1469-8137.2006.01862.x , doi-access=free{{cite journal , vauthors=Heitman J , s2cid=2898102 , title=Sexual reproduction and the evolution of microbial pathogens , journal=Current Biology , volume=16 , issue=17 , pages=R711–25 , date=September 2006 , pmid=16950098 , doi=10.1016/j.cub.2006.07.064 , doi-access=free{{cite journal , vauthors=Hetland G, Johnson E, Lyberg T, Bernardshaw S, Tryggestad AM, Grinde B , s2cid=3866471 , title=Effects of the medicinal mushroom ''Agaricus blazei'' Murill on immunity, infection and cancer , journal=Scandinavian Journal of Immunology , volume=68 , issue=4 , pages=363–70 , date=October 2008 , pmid=18782264 , doi=10.1111/j.1365-3083.2008.02156.x , doi-access=free{{cite journal , vauthors=Hynes MJ , s2cid=45815733 , title=Regulatory circuits of the amdS gene of ''Aspergillus nidulans'' , journal=Antonie van Leeuwenhoek , volume=65 , issue=3 , pages=179–82 , year=1994 , pmid=7847883 , doi=10.1007/BF00871944{{cite journal , vauthors=Hibbett DS, Grimaldi D, Donoghue MJ , s2cid=4346359 , year=1995 , title=Cretaceous mushrooms in amber , journal=Nature (journal), Nature , volume=377 , issue=6549 , page=487 , doi=10.1038/377487a0 , bibcode=1995Natur.377..487H, doi-access=free{{cite journal , vauthors=Hibbett DS, Binder M, Bischoff JF, Blackwell M, Cannon PF, Eriksson OE, Huhndorf S, James T, Kirk PM, Lücking R, Thorsten Lumbsch H, Lutzoni F, Matheny PB, McLaughlin DJ, Powell MJ, Redhead S, Schoch CL, Spatafora JW, Stalpers JA, Vilgalys R, Aime MC, Aptroot A, Bauer R, Begerow D, Benny GL, Castlebury LA, Crous PW, Dai YC, Gams W, Geiser DM, Griffith GW, Gueidan C, Hawksworth DL, Hestmark G, Hosaka K, Humber RA, Hyde KD, Ironside JE, Kõljalg U, Kurtzman CP, Larsson KH, Lichtwardt R, Longcore J, Miadlikowska J, Miller A, Moncalvo JM, Mozley-Standridge S, Oberwinkler F, Parmasto E, Reeb V, Rogers JD, Roux C, Ryvarden L, Sampaio JP, Schüssler A, Sugiyama J, Thorn RG, Tibell L, Untereiner WA, Walker C, Wang Z, Weir A, Weiss M, White MM, Winka K, Yao YJ, Zhang N , display-authors=6 , title=A higher-level phylogenetic classification of the Fungi , journal=Mycological Research , volume=111 , issue=Pt 5 , pages=509–47 , date=May 2007 , pmid=17572334 , doi=10.1016/j.mycres.2007.03.004 , url=http://www.clarku.edu/faculty/dhibbett/AFTOL/documents/AFTOL%20class%20mss%2023,%2024/AFTOL%20CLASS%20MS%20resub.pdf , url-status=dead , archive-url=https://web.archive.org/web/20090326135053/http://www.clarku.edu/faculty/dhibbett/AFTOL/documents/AFTOL%20class%20mss%2023%2C%2024/AFTOL%20CLASS%20MS%20resub.pdf , archive-date=26 March 2009 , citeseerx=10.1.1.626.9582 , s2cid=4686378{{cite journal , vauthors=Hibbett DS, Grimaldi D, Donoghue MJ , s2cid=22011469 , year=1997 , title=Fossil mushrooms from Miocene and Cretaceous ambers and the evolution of homobasidiomycetes , journal=American Journal of Botany , volume=84 , issue=7 , pages=981–991 , doi=10.2307/2446289 , jstor=2446289 , pmid=21708653 , doi-access=free{{cite journal , last1=Honegger , first1=Rosmarie , authorlink1=Rosmarie Honegger, last2=Edwards , first2=Dianne , last3=Axe , first3=Lindsey , title=The earliest records of internally stratified cyanobacterial and algal lichens from the Lower Devonian of the Welsh Borderland , journal=New Phytologist , volume=197 , issue=1 , year=2013 , pages=264–275 , doi=10.1111/nph.12009 , pmid=23110612{{cite journal , vauthors=Howard RJ, Ferrari MA, Roach DH, Money NP , title=Penetration of hard substrates by a fungus employing enormous turgor pressures , journal=Proceedings of the National Academy of Sciences of the United States of America , volume=88 , issue=24 , pages=11281–4 , date=December 1991 , pmid=1837147 , pmc=53118 , doi=10.1073/pnas.88.24.11281 , bibcode=1991PNAS...8811281H , doi-access=free{{cite journal , vauthors=Huang B, Guo J, Yi B, Yu X, Sun L, Chen W , s2cid=2222358 , title=Heterologous production of secondary metabolites as pharmaceuticals in ''Saccharomyces cerevisiae'' , journal=Biotechnology Letters , volume=30 , issue=7 , pages=1121–37 , date=July 2008 , pmid=18512022 , doi=10.1007/s10529-008-9663-z{{cite journal , vauthors=James TY, Letcher PM, Longcore JE, Mozley-Standridge SE, Porter D, Powell MJ, Griffith GW, Vilgalys R , title=A molecular phylogeny of the flagellated fungi (Chytridiomycota) and description of a new phylum (Blastocladiomycota) , journal=Mycologia , volume=98 , issue=6 , pages=860–71 , year=2006 , pmid=17486963 , doi=10.3852/mycologia.98.6.860 , url=http://www.cybertruffle.org.uk/cyberliber/59350/0098/006/0860.htm , access-date=5 July 2011 , archive-url=https://web.archive.org/web/20150923230802/http://www.cybertruffle.org.uk/cyberliber/59350/0098/006/0860.htm , archive-date=23 September 2015 , url-status=live{{cite journal , vauthors=Hube B , title=From commensal to pathogen: stage- and tissue-specific gene expression of ''Candida albicans'' , journal=Current Opinion in Microbiology , volume=7 , issue=4 , pages=336–41 , date=August 2004 , pmid=15288621 , doi=10.1016/j.mib.2004.06.003{{cite journal , last1=Jakovlev , first1=Jevgeni , year=2012 , title=Fungal hosts of mycetophilids (Diptera: Sciaroidea excluding Sciaridae): a review , journal=Mycology , volume=3 , issue=1 , pages=11–23 , doi=10.1080/21501203.2012.662533 , s2cid=82107953 , url=https://www.researchgate.net/publication/254268258{{cite journal , vauthors=James TY, Kauff F, Schoch CL, Matheny PB, Hofstetter V, Cox CJ, Celio G, Gueidan C, Fraker E, Miadlikowska J, Lumbsch HT, Rauhut A, Reeb V, Arnold AE, Amtoft A, Stajich JE, Hosaka K, Sung GH, Johnson D, O'Rourke B, Crockett M, Binder M, Curtis JM, Slot JC, Wang Z, Wilson AW, Schüssler A, Longcore JE, O'Donnell K, Mozley-Standridge S, Porter D, Letcher PM, Powell MJ, Taylor JW, White MM, Griffith GW, Davies DR, Humber RA, Morton JB, Sugiyama J, Rossman AY, Rogers JD, Pfister DH, Hewitt D, Hansen K, Hambleton S, Shoemaker RA, Kohlmeyer J, Volkmann-Kohlmeyer B, Spotts RA, Serdani M, Crous PW, Hughes KW, Matsuura K, Langer E, Langer G, Untereiner WA, Lücking R, Büdel B, Geiser DM, Aptroot A, Diederich P, Schmitt I, Schultz M, Yahr R, Hibbett DS, Lutzoni F, McLaughlin DJ, Spatafora JW, Vilgalys R , s2cid=4302864 , display-authors=6 , title=Reconstructing the early evolution of Fungi using a six-gene phylogeny , journal=Nature (journal), Nature , volume=443 , issue=7113 , pages=818–22 , date=October 2006 , pmid=17051209 , doi=10.1038/nature05110 , bibcode=2006Natur.443..818J{{cite journal , last1=Janik , first1=Edyta , last2=Niemcewicz , first2=Marcin , last3=Ceremuga , first3=Michal , last4=Stela , first4=Maksymilian , last5=Saluk-Bijak , first5=Joanna , last6=Siadkowski , first6=Adrian , last7=Bijak , first7=Michal , title=Molecular aspects of mycotoxins—a serious problem for human health , journal=International Journal of Molecular Sciences , volume=21 , issue=21 , year=2020 , pages=8187 , doi=10.3390/ijms21218187 , pmid=33142955 , pmc=7662353 , doi-access=free{{cite journal , vauthors=Jørgensen TR , title=Identification and toxigenic potential of the industrially important fungi, ''Aspergillus oryzae'' and ''Aspergillus sojae'' , journal=Journal of Food Protection , volume=70 , issue=12 , pages=2916–34 , date=December 2007 , pmid=18095455 , doi=10.4315/0362-028X-70.12.2916{{cite journal , vauthors=Joseph B, Ramteke PW, Thomas G , title=Cold active microbial lipases: some hot issues and recent developments , journal=Biotechnology Advances , volume=26 , issue=5 , pages=457–70 , year=2008 , pmid=18571355 , doi=10.1016/j.biotechadv.2008.05.003{{cite journal , last1=Joseph , first1=Ross , last2=Keyhani , first2=Nemat O. , title=Fungal mutualisms and pathosystems: life and death in the ambrosia beetle mycangia , journal=Applied Microbiology and Biotechnology , volume=105 , issue=9 , year=2021 , pages=3393–3410 , doi=10.1007/s00253-021-11268-0 , pmid=33837831, s2cid=233200379{{cite journal , last1=Karbalaei , first1=Mohsen , last2=Rezaee , first2=Seyed A. , last3=Farsiani , first3=Hadi , title=''Pichia pastoris'': A highly successful expression system for optimal synthesis of heterologous proteins , journal=Journal of Cellular Physiology , volume=235 , issue=9 , year=2020 , pages=5867–5881 , doi=10.1002/jcp.29583 , pmc=7228273 , pmid=32057111{{cite journal , vauthors=Karlson-Stiber C, Persson H , title=Cytotoxic fungi--an overview , journal=Toxicon , volume=42 , issue=4 , pages=339–49 , date=September 2003 , pmid=14505933 , doi=10.1016/S0041-0101(03)00238-1{{cite journal , vauthors=Kauffman CA , title=Histoplasmosis: a clinical and laboratory update , journal=Clinical Microbiology Reviews , volume=20 , issue=1 , pages=115–32 , date=January 2007 , pmid=17223625 , pmc=1797635 , doi=10.1128/CMR.00027-06{{cite journal , vauthors=Keller NP, Turner G, Bennett JW , s2cid=23537608 , title=Fungal secondary metabolism - from biochemistry to genomics , journal=Nature Reviews. Microbiology , volume=3 , issue=12 , pages=937–47 , date=December 2005 , pmid=16322742 , doi=10.1038/nrmicro1286{{cite journal , vauthors=Kinsella JE, Hwang DH , title=Enzymes of ''Penicillium roqueforti'' involved in the biosynthesis of cheese flavor , journal=Critical Reviews in Food Science and Nutrition , volume=8 , issue=2 , pages=191–228 , date=November 1976 , pmid=21770 , doi=10.1080/10408397609527222{{cite journal , vauthors=Kojic M, Zhou Q, Lisby M, Holloman WK , title=Rec2 interplay with both Brh2 and Rad51 balances recombinational repair in ''Ustilago maydis'' , journal=Molecular and Cellular Biology , volume=26 , issue=2 , pages=678–88 , date=January 2006 , pmid=16382157 , pmc=1346908 , doi=10.1128/MCB.26.2.678-688.2006{{cite journal , last1=Kuhar , first1=Francisco , last2=Furci , first2=Giuliana , last3=Drechsler-Santos , first3=Elisandro Ricardo , last4=Pfister , first4=Donald H. , title=Delimitation of Funga as a valid term for the diversity of fungal communities: the Fauna, Flora & Funga proposal (FF&F) , journal=IMA Fungus , volume=9 , issue=2 , year=2018 , pages=A71–A74 , doi=10.1007/BF03449441 , doi-access=free{{cite book , title=Handbook of Cereal Science and Technology , vauthors=Kulp K , year=2000 , publisher=CRC Press , isbn=978-0-8247-8294-8{{cite journal , vauthors=Kumar R, Singh S, Singh OV , s2cid=4830678 , title=Bioconversion of lignocellulosic biomass: biochemical and molecular perspectives , journal=Journal of Industrial Microbiology & Biotechnology , volume=35 , issue=5 , pages=377–91 , date=May 2008 , pmid=18338189 , doi=10.1007/s10295-008-0327-8{{cite journal , vauthors=Leathem AM, Dorran TJ , title=Poisoning due to raw ''Gyromitra esculenta'' (false morels) west of the Rockies , journal=Canadian Journal of Emergency Medicine , volume=9 , issue=2 , pages=127–30 , date=March 2007 , pmid=17391587 , doi=10.1017/s1481803500014937 , doi-access=free{{cite book , editor-last1=Stephenson , editor-first1=Steven L. , editor-last2=Rojas , editor-first2=Carlos , last1=Leontyev , first1=Dmitry V. , last2=Schnittler , first2=Martin , chapter=The phylogeny of Myxomycetes , title=Myxomycetes. Biology, Systematics, Biogeography, and Ecology , year=2017 , pages=83–106 , doi=10.1016/B978-0-12-805089-7.00003-2 , publisher=Academic Press , isbn=978-0-12-805089-7{{cite journal , vauthors=Lin X, Hull CM, Heitman J , s2cid=52857557 , title=Sexual reproduction between partners of the same mating type in ''Cryptococcus neoformans'' , journal=Nature (journal), Nature , volume=434 , issue=7036 , pages=1017–21 , date=April 2005 , pmid=15846346 , doi=10.1038/nature03448 , bibcode=2005Natur.434.1017L{{cite journal , vauthors=Linder MB, Szilvay GR, Nakari-Setälä T, Penttilä ME , title=Hydrophobins: the protein-amphiphiles of filamentous fungi , journal=FEMS Microbiology Reviews , volume=29 , issue=5 , pages=877–96 , date=November 2005 , pmid=16219510 , doi=10.1016/j.femsre.2005.01.004 , doi-access=free{{cite journal , vauthors=Lindahl BD, Ihrmark K, Boberg J, Susan Trumbore, Trumbore SE, Högberg P, Stenlid J, Finlay RD , title=Spatial separation of litter decomposition and mycorrhizal nitrogen uptake in a boreal forest , journal=The New Phytologist , volume=173 , issue=3 , pages=611–20 , year=2007 , pmid=17244056 , doi=10.1111/j.1469-8137.2006.01936.x , hdl=11858/00-001M-0000-0027-D56D-D , url=http://www.escholarship.org/uc/item/1r43h5sj , doi-access=free{{cite journal , vauthors=Lockhart RJ, Van Dyke MI, Beadle IR, Humphreys P, McCarthy AJ , title=Molecular biological detection of anaerobic gut fungi (Neocallimastigales) from landfill sites , journal=Applied and Environmental Microbiology , volume=72 , issue=8 , pages=5659–61 , date=August 2006 , pmid=16885325 , doi=10.1128/AEM.01057-06 , pmc=1538735 , bibcode=2006ApEnM..72.5659L{{cite journal , vauthors=Loo DS , title=Systemic antifungal agents: an update of established and new therapies , journal=Advances in Dermatology , volume=22 , pages=101–24 , year=2006 , pmid=17249298 , doi=10.1016/j.yadr.2006.07.001{{cite journal , vauthors=López-Gómez J, Taylor EL , title=Permian-Triassic transition in Spain: a multidisciplinary approach , journal=Palaeogeography, Palaeoclimatology, Palaeoecology , volume=229 , issue=1–2 , year=2005 , pages=1–2 , doi=10.1016/j.palaeo.2005.06.028{{cite journal , vauthors=Looy CV, Twitchett RJ, Dilcher DL, Van Konijnenburg-Van Cittert JH, Visscher H , title=Life in the end-Permian dead zone , journal=Proceedings of the National Academy of Sciences of the United States of America , volume=98 , issue=14 , pages=7879–83 , date=July 2001 , pmid=11427710 , pmc=35436 , doi=10.1073/pnas.131218098 , quote=See image 2 , bibcode=2001PNAS...98.7879L , doi-access=free{{cite journal , last1=Lu , first1=Jiahui , last2=He , first2=Rongjun , last3=Sun , first3=Peilong , last4=Zhang , first4=Fuming , last5=Linhardt , first5=Robert J. , last6=Zhang , first6=Anqiang , title=Molecular mechanisms of bioactive polysaccharides from ''Ganoderma lucidum'' (Lingzhi), a review , journal=International Journal of Biological Macromolecules , volume=150 , year=2020 , pages=765–774 , doi=10.1016/j.ijbiomac.2020.02.035 , pmid=32035956 , s2cid=211071754{{cite journal , vauthors=Lücking R, Huhndorf S, Pfister DH, Plata ER, Lumbsch HT , s2cid=6689439 , title=Fungi evolved right on track , journal=Mycologia , volume=101 , issue=6 , pages=810–22 , year=2009 , pmid=19927746 , doi=10.3852/09-016 , url=http://nrs.harvard.edu/urn-3:HUL.InstRepos:14168857{{Cite book , vauthors=Hawksworth DL, Lücking R , title=The Fungal Kingdom , chapter=Fungal Diversity Revisited: 2.2 to 3.8 Million Species , journal=Microbiology Spectrum , volume=5 , issue=4 , pages=79–95 , date=July 2017 , pmid=28752818 , doi=10.1128/microbiolspec.FUNK-0052-2016 , isbn=978-1-55581-957-6{{cite journal , vauthors=López-Gómez J, Molina-Meyer M , title=The competitive exclusion principle versus biodiversity through competitive segregation and further adaptation to spatial heterogeneities , journal=Theoretical Population Biology , volume=69 , issue=1 , pages=94–109 , date=February 2006 , pmid=16223517 , doi=10.1016/j.tpb.2005.08.004{{cite journal , vauthors=Manzoni M, Rollini M , s2cid=5761188 , title=Biosynthesis and biotechnological production of statins by filamentous fungi and application of these cholesterol-lowering drugs , journal=Applied Microbiology and Biotechnology , volume=58 , issue=5 , pages=555–64 , date=April 2002 , pmid=11956737 , doi=10.1007/s00253-002-0932-9{{cite journal , vauthors=Metzenberg RL, Glass NL , title=Mating type and mating strategies in Neurospora , journal=BioEssays , volume=12 , issue=2 , pages=53–9 , date=February 1990 , pmid=2140508 , doi=10.1002/bies.950120202 , s2cid=10818930{{cite journal , vauthors=Merckx V, Bidartondo MI, Hynson NA , title=Myco-heterotrophy: when fungi host plants , journal=Annals of Botany , volume=104 , issue=7 , pages=1255–61 , date=December 2009 , pmid=19767309 , pmc=2778383 , doi=10.1093/aob/mcp235{{cite journal , vauthors=Michod RE, Bernstein H, Nedelcu AM , title=Adaptive value of sex in microbial pathogens , journal=Infection, Genetics and Evolution , volume=8 , issue=3 , pages=267–85 , date=May 2008 , pmid=18295550 , doi=10.1016/j.meegid.2008.01.002 , url=http://www.hummingbirds.arizona.edu/Faculty/Michod/Downloads/IGE%20review%20sex.pdf , access-date=22 July 2013 , archive-url=https://web.archive.org/web/20170516235741/http://www.hummingbirds.arizona.edu/Faculty/Michod/Downloads/IGE%20review%20sex.pdf , archive-date=16 May 2017 , url-status=live{{cite journal , vauthors=Mihail JD, Bruhn JN , title=Foraging behaviour of ''Armillaria'' rhizomorph systems , journal=Mycological Research , volume=109 , issue=Pt 11 , pages=1195–207 , date=November 2005 , pmid=16279413 , doi=10.1017/S0953756205003606{{cite journal , vauthors=Michelot D, Melendez-Howell LM , s2cid=41451034 , title=''Amanita muscaria'': chemistry, biology, toxicology, and ethnomycology , journal=Mycological Research , volume=107 , issue=Pt 2 , pages=131–46 , date=February 2003 , pmid=12747324 , doi=10.1017/S0953756203007305{{cite book , veditors=Mitzka W , year=1960 , title=Etymologisches Wörterbuch der deutschen Sprache , trans-title=Etymological dictionary of the German language , location=Berlin , publisher=Walter de Gruyter , language=de{{cite journal , vauthors=Molina L, Kahmann R , title=An ''Ustilago maydis'' gene involved in H2O2 detoxification is required for virulence , journal=The Plant Cell , volume=19 , issue=7 , pages=2293–309 , date=July 2007 , pmid=17616735 , pmc=1955693 , doi=10.1105/tpc.107.052332{{cite book , vauthors=Money NP , chapter=Mechanics of invasive fungal growth and the significance of turgor in plant infection , title=Molecular Genetics of Host-Specific Toxins in Plant Disease: Proceedings of the 3rd Tottori International Symposium on Host-Specific Toxins, Daisen, Tottori, Japan, August 24–29, 1997 , publisher=Kluwer Academic Publishers , year=1998 , location=Netherlands , pages=261–271 , isbn=978-0-7923-4981-5{{cite journal , vauthors=Moore RT , year=1980 , title=Taxonomic proposals for the classification of marine yeasts and other yeast-like fungi including the smuts , journal=Botanica Marina , volume=23 , pages=361–373{{cite journal , vauthors=Money NP , title=Mushroom stem cells , journal=BioEssays , volume=24 , issue=10 , pages=949–52 , date=October 2002 , pmid=12325127 , doi=10.1002/bies.10160{{cite book , vauthors=Moss ST , title=The Biology of Marine Fungi , publisher=Cambridge University Press , location=Cambridge, UK , year=1986 , page=76 , isbn=978-0-521-30899-1{{cite journal , vauthors=Mueller GM, Schmit JP , s2cid=23827807 , year=2006 , title=Fungal biodiversity: what do we know? What can we predict? , journal=Biodiversity and Conservation , volume=16 , pages=1–5 , doi=10.1007/s10531-006-9117-7{{cite book , vauthors=Esser K , editor1-first=David J , editor1-last=McLaughlin , editor2-first=Joseph W , editor2-last=Spatafora , title=The Mycota VII A: Systematics and Evolution (2nd ed.) , series=The Mycota , publisher=Springer , year=2014 , page=461 , doi=10.1007/978-3-642-55318-9 , isbn=978-3-642-55317-2 , s2cid=46141350 , url=https://www.springer.com/gp/book/9783642553172{{cite journal , last1=Naranjo‐Ortiz , first1=Miguel A. , last2=Gabaldón , first2=Toni , title=Fungal evolution: Diversity, taxonomy and phylogeny of the Fungi , journal=Biological Reviews , volume=94 , issue=6 , year=2019 , pages=2101–2137 , doi=10.1111/brv.12550 , pmid=31659870 , pmc=6899921 , doi-access=free{{cite journal , vauthors=Nikoh N, Fukatsu T , title=Interkingdom host jumping underground: phylogenetic analysis of entomoparasitic fungi of the genus ''Cordyceps'' , journal=Molecular Biology and Evolution , volume=17 , issue=4 , pages=629–38 , date=April 2000 , pmid=10742053 , doi=10.1093/oxfordjournals.molbev.a026341 , doi-access=free{{Cite book , vauthors=Nielsen K, Heitman J , volume=57 , pages=143–73 , year=2007 , pmid=17352904 , doi=10.1016/S0065-2660(06)57004-X , isbn=978-0-12-017657-1 , series=Advances in Genetics , title=Fungal Genomics , chapter=Sex and Virulence of Human Pathogenic Fungi{{cite journal , vauthors=Nguyen NH, Suh SO, Blackwell M , title=Five novel Candida species in insect-associated yeast clades isolated from Neuroptera and other insects , journal=Mycologia , volume=99 , issue=6 , pages=842–58 , year=2007 , pmid=18333508 , doi=10.3852/mycologia.99.6.842 , url=http://www.cybertruffle.org.uk/cyberliber/59350/0099/006/0842.htm , access-date=5 July 2011 , archive-url=https://web.archive.org/web/20170507072249/http://www.cybertruffle.org.uk/cyberliber/59350/0099/006/0842.htm , archive-date=7 May 2017 , url-status=live{{cite web , url=http://www.biocontrol.entomology.cornell.edu/pathogens/trichoderma.html , title=''Trichoderma'' spp., including ''T. harzianum'', ''T. viride'', ''T. koningii'', ''T. hamatum'' and other spp. Deuteromycetes, Moniliales (asexual classification system) , access-date=10 July 2007 , work=Biological Control: A Guide to Natural Enemies in North America , url-status=dead , archive-url=https://web.archive.org/web/20110414111846/http://www.biocontrol.entomology.cornell.edu/pathogens/trichoderma.html , archive-date=14 April 2011{{cite journal , vauthors=O'Donnell K, Cigelnik E, Casper HH , title=Molecular phylogenetic, morphological, and mycotoxin data support reidentification of the Quorn mycoprotein fungus as ''Fusarium venenatum'' , journal=Fungal Genetics and Biology , volume=23 , issue=1 , pages=57–67 , date=February 1998 , pmid=9501477 , doi=10.1006/fgbi.1997.1018 , s2cid=23049409{{cite journal , last1=Olatunji , first1=Opeyemi Joshua , last2=Tang , first2=Jian , last3=Tola , first3=Adesola , last4=Auberon , first4=Florence , last5=Oluwaniyi , first5=Omolara , last6=Ouyang , first6=Zhen , title=The genus ''Cordyceps'': An extensive review of its traditional uses, phytochemistry and pharmacology , journal=Fitoterapia , volume=129 , year=2018, pages=293–316 , doi=10.1016/j.fitote.2018.05.010 , pmid=29775778 , s2cid=21741034{{cite journal , vauthors=Olempska-Beer ZS, Merker RI, Ditto MD, DiNovi MJ , title=Food-processing enzymes from recombinant microorganisms--a review , journal=Regulatory Toxicology and Pharmacology , volume=45 , issue=2 , pages=144–158 , date=July 2006 , pmid=16769167 , doi=10.1016/j.yrtph.2006.05.001 , url=https://zenodo.org/record/1259499 , access-date=3 July 2019 , archive-url=https://web.archive.org/web/20190703164318/https://zenodo.org/record/1259499 , archive-date=3 July 2019 , url-status=live{{cite journal , last1=Olicón-Hernández , first1=Dario R. , last2=Araiza-Villanueva , first2=Minerva G. , last3=Pardo , first3=Juan P. , last4=Aranda , first4=Elisabet , last5=Guerra-Sánchez , first5=Guadalupe , title=New insights of ''Ustilago maydis'' as yeast model for genetic and biotechnological research: A review , journal=Current Microbiology , volume=76 , issue=8 , year=2019 , pages=917–926 , doi=10.1007/s00284-019-01629-4 , pmid=30689003 , s2cid=59307118{{cite web , url=http://oxforddictionaries.com/definition/english/fungus , title=Fungus , work=Oxford Dictionaries , access-date=26 February 2011 , archive-url=https://web.archive.org/web/20120728023308/http://oxforddictionaries.com/definition/english/fungus , archive-date=28 July 2012 , url-status=dead{{cite book , vauthors=Orr DB, Orr RT , title=Mushrooms of Western North America , publisher=University of California Press , location=Berkeley, California , year=1979 , page=17 , isbn=978-0-520-03656-7{{cite journal , vauthors=Pan A, Lorenzotti S, Zoncada A , title=Registered and investigational drugs for the treatment of methicillin-resistant ''Staphylococcus aureus'' infection , journal=Recent Patents on Anti-Infective Drug Discovery , volume=3 , issue=1 , pages=10–33 , date=January 2008 , pmid=18221183 , doi=10.2174/157489108783413173{{cite journal , vauthors=Parish JA, McCann MA, Watson RH, Hoveland CS, Hawkins LL, Hill NS, Bouton JH , title=Use of nonergot alkaloid-producing endophytes for alleviating tall fescue toxicosis in sheep , journal=Journal of Animal Science , volume=81 , issue=5 , pages=1316–22 , date=May 2003 , pmid=12772860 , doi=10.2527/2003.8151316x{{cite journal , vauthors=Parniske M , s2cid=5432120 , title=Arbuscular mycorrhiza: the mother of plant root endosymbioses , journal=Nature Reviews. Microbiology , volume=6 , issue=10 , pages=763–75 , date=October 2008 , pmid=18794914 , doi=10.1038/nrmicro1987{{cite journal , vauthors=Paszkowski U , title=Mutualism and parasitism: the yin and yang of plant symbioses , journal=Current Opinion in Plant Biology , volume=9 , issue=4 , pages=364–70 , date=August 2006 , pmid=16713732 , doi=10.1016/j.pbi.2006.05.008{{cite journal , vauthors=Peñalva MA, Arst HN , title=Regulation of gene expression by ambient pH in filamentous fungi and yeasts , journal=Microbiology and Molecular Biology Reviews , volume=66 , issue=3 , pages=426–46, table of contents , date=September 2002 , pmid=12208998 , pmc=120796 , doi=10.1128/MMBR.66.3.426-446.2002{{cite journal , vauthors=Peintner U, Pöder R, Pümpel T , title=The Iceman's fungi , journal=Mycological Research , volume=102 , issue=10 , pages=1153–1162 , year=1998 , doi=10.1017/S0953756298006546{{cite journal , vauthors=Pereira JL, Noronha EF, Miller RN, Franco OL , title=Novel insights in the use of hydrolytic enzymes secreted by fungi with biotechnological potential , journal=Letters in Applied Microbiology , volume=44 , issue=6 , pages=573–81 , date=June 2007 , pmid=17576216 , doi=10.1111/j.1472-765X.2007.02151.x , doi-access=free{{cite journal , vauthors=Perotto S, Bonfante P , title=Bacterial associations with mycorrhizal fungi: close and distant friends in the rhizosphere , journal=Trends in Microbiology , volume=5 , issue=12 , pages=496–501 , date=December 1997 , pmid=9447662 , doi=10.1016/S0966-842X(97)01154-2{{cite journal , vauthors=Perfect JR , title=''Cryptococcus neoformans'': the yeast that likes it hot , journal=FEMS Yeast Research , volume=6 , issue=4 , pages=463–8 , date=June 2006 , pmid=16696642 , doi=10.1111/j.1567-1364.2006.00051.x , doi-access=free{{cite journal , vauthors=Piskur J, Rozpedowska E, Polakova S, Merico A, Compagno C , title=How did ''Saccharomyces'' evolve to become a good brewer? , journal=Trends in Genetics , volume=22 , issue=4 , pages=183–6 , date=April 2006 , pmid=16499989 , doi=10.1016/j.tig.2006.02.002{{cite journal , vauthors=Pringle A, Patek SN, Fischer M, Stolze J, Money NP , title=The captured launch of a ballistospore , journal=Mycologia , volume=97 , issue=4 , pages=866–71 , year=2005 , pmid=16457355 , doi=10.3852/mycologia.97.4.866 , url=http://www.cybertruffle.org.uk/cyberliber/59350/0097/004/0866.htm , access-date=5 July 2011 , archive-url=https://web.archive.org/web/20160412145956/http://www.cybertruffle.org.uk/cyberliber/59350/0097/004/0866.htm , archive-date=12 April 2016 , url-status=live{{cite journal , vauthors=Polizeli ML, Rizzatti AC, Monti R, Terenzi HF, Jorge JA, Amorim DS , s2cid=22956 , title=Xylanases from fungi: properties and industrial applications , journal=Applied Microbiology and Biotechnology , volume=67 , issue=5 , pages=577–91 , date=June 2005 , pmid=15944805 , doi=10.1007/s00253-005-1904-7{{cite book , vauthors=Purvis W , title=Lichens , publisher=Smithsonian Institution Press in association with the Natural History Museum, London , location=Washington, D.C. , year=2000 , page 49–75 , isbn=978-1-56098-879-3 , url-access=registration , url=https://archive.org/details/lichens00purv/page/49{{cite journal , vauthors=Raghukumar C, Raghukumar S , title=Barotolerance of fungi isolated from deep-sea sediments of the Indian Ocean , journal=Aquatic Microbial Ecology , volume=15 , issue=2 , pages=153–163 , year=1998 , doi=10.3354/ame015153 , doi-access=free{{cite book , vauthors=Raven PH, Evert RF, Eichhorn, SE , title=Biology of Plants , chapter-url=https://archive.org/details/biologyofplants00rave_0 , chapter-url-access=registration , edition=7 , publisher=W. H. Freeman , year=2005 , pag 290 , chapter=14—Fungi , isbn=978-0-7167-1007-3{{cite journal , vauthors=Redecker D, Raab P , title=Phylogeny of the glomeromycota (arbuscular mycorrhizal fungi): recent developments and new gene markers , journal=Mycologia , volume=98 , issue=6 , pages=885–95 , year=2006 , pmid=17486965 , doi=10.3852/mycologia.98.6.885 , url=http://www.cybertruffle.org.uk/cyberliber/59350/0098/006/0885.htm , access-date=5 July 2011 , archive-url=https://web.archive.org/web/20150923230805/http://www.cybertruffle.org.uk/cyberliber/59350/0098/006/0885.htm , archive-date=23 September 2015 , url-status=live{{cite journal , vauthors=Redecker D, Kodner R, Graham LE , s2cid=43553633 , title=Glomalean fungi from the Ordovician , journal=Science (journal), Science , volume=289 , issue=5486 , pages=1920–1 , date=September 2000 , pmid=10988069 , doi=10.1126/science.289.5486.1920 , bibcode=2000Sci...289.1920R{{cite journal , last1=Redhead , first1=Scott , last2=Norvell , first2=Lorelei , year=2013 , title=MycoBank, Index Fungorum, and Fungal Names recommended as official nomenclatural repositories for 2013 , journal=IMA Fungus , volume=3 , issue=2 , pages=44–45 , url=https://www.researchgate.net/publication/255719633{{cite journal , vauthors=Remy W, Taylor TN, Hass H, Kerp H , title=Four hundred-million-year-old vesicular arbuscular mycorrhizae , journal=Proceedings of the National Academy of Sciences of the United States of America , volume=91 , issue=25 , pages=11841–3 , date=December 1994 , pmid=11607500 , pmc=45331 , doi=10.1073/pnas.91.25.11841 , bibcode=1994PNAS...9111841R , doi-access=free{{cite journal , last1=Rhimi , first1=Wafa , last2=Theelen , first2=Bart , last3=Boekhout , first3=Teun , last4=Otranto , first4=Domenico , last5=Cafarchia , first5=Claudia , title=''Malassezia'' spp. yeasts of emerging concern in fungemia , journal=Frontiers in Cellular and Infection Microbiology , volume=10 , year=2020 , page=370 , doi=10.3389/fcimb.2020.00370 , pmid=32850475 , pmc=7399178, doi-access=free{{cite journal , last1=Rigling , first1=Daniel , last2=Prospero , first2=Simone , title=''Cryphonectria parasitica'', the causal agent of chestnut blight: invasion history, population biology and disease control , journal=Molecular Plant Pathology , volume=19 , issue=1 , year=2018 , pages=7–20 , doi=10.1111/mpp.12542 , pmid=28142223 , pmc=6638123{{cite journal , vauthors=Rohlfs M, Albert M, Keller NP, Kempken F , title=Secondary chemicals protect mould from fungivory , journal=Biology Letters , volume=3 , issue=5 , pages=523–5 , date=October 2007 , pmid=17686752 , pmc=2391202 , doi=10.1098/rsbl.2007.0338{{cite journal , last1=Rossman , first1=Amy Y. , title=Lessons learned from moving to one scientific name for fungi , journal=IMA Fungus , volume=5 , issue=1 , year=2014 , pages=81–89 , pmc=4107901 , pmid=25083410 , doi=10.5598/imafungus.2014.05.01.10For an example, see {{cite journal , vauthors=Samuels GJ , title=''Trichoderma'': systematics, the sexual state, and ecology , journal=Phytopathology , volume=96 , issue=2 , pages=195–206 , date=February 2006 , pmid=18943925 , doi=10.1094/PHYTO-96-0195 , url=https://zenodo.org/record/1235933 , doi-access=free{{cite journal , vauthors=Sancho LG, de la Torre R, Horneck G, Ascaso C, de Los Rios A, Pintado A, Wierzchos J, Schuster M , s2cid=4121180 , title=Lichens survive in space: results from the 2005 LICHENS experiment , journal=Astrobiology , volume=7 , issue=3 , pages=443–54 , date=June 2007 , pmid=17630840 , doi=10.1089/ast.2006.0046 , bibcode=2007AsBio...7..443S{{cite journal , last1=Santini , first1=Alberto , last2=Battisti , first2=Andrea , title=Complex insect–pathogen interactions in tree pandemics , journal=Frontiers in Physiology , volume=10 , year=2019 , page=550 , doi=10.3389/fphys.2019.00550 , pmc=6517489 , pmid=31133880 , doi-access=free{{cite journal , vauthors=Schaller M, Borelli C, Korting HC, Hube B , title=Hydrolytic enzymes as virulence factors of ''Candida albicans'' , journal=Mycoses , volume=48 , issue=6 , pages=365–77 , date=November 2005 , pmid=16262871 , doi=10.1111/j.1439-0507.2005.01165.x , s2cid=1356254{{cite journal , vauthors=Schardl CL, Craven KD , title=Interspecific hybridization in plant-associated fungi and oomycetes: a review , journal=Molecular Ecology , volume=12 , issue=11 , pages=2861–73 , date=November 2003 , pmid=14629368 , doi=10.1046/j.1365-294X.2003.01965.x , s2cid=25879264{{cite book , vauthors=Schlegel HG , title=General Microbiology , publisher=Cambridge University Press , location=Cambridge, UK , year=1993 , page=360 , isbn=978-0-521-43980-0{{Cite book , vauthors=Schardl CL, Panaccione DG, Tudzynski P , title=Ergot alkaloids--biology and molecular biology , volume=63 , pages=45–86 , year=2006 , pmid=17133714 , doi=10.1016/S1099-4831(06)63002-2 , isbn=978-0-12-469563-4 , series=The Alkaloids: Chemistry and Biology{{cite journal , vauthors=Schoch CL, Sung GH, López-Giráldez F, Townsend JP, Miadlikowska J, Hofstetter V, Robbertse B, Matheny PB, Kauff F, Wang Z, Gueidan C, Andrie RM, Trippe K, Ciufetti LM, Wynns A, Fraker E, Hodkinson BP, Bonito G, Groenewald JZ, Arzanlou M, de Hoog GS, Crous PW, Hewitt D, Pfister DH, Peterson K, Gryzenhout M, Wingfield MJ, Aptroot A, Suh SO, Blackwell M, Hillis DM, Griffith GW, Castlebury LA, Rossman AY, Lumbsch HT, Lücking R, Büdel B, Rauhut A, Diederich P, Ertz D, Geiser DM, Hosaka K, Inderbitzin P, Kohlmeyer J, Volkmann-Kohlmeyer B, Mostert L, O'Donnell K, Sipman H, Rogers JD, Shoemaker RA, Sugiyama J, Summerbell RC, Untereiner W, Johnston PR, Stenroos S, Zuccaro A, Dyer PS, Crittenden PD, Cole MS, Hansen K, Trappe JM, Yahr R, Lutzoni F, Spatafora JW , display-authors=6 , title=The Ascomycota tree of life: a phylum-wide phylogeny clarifies the origin and evolution of fundamental reproductive and ecological traits , journal=Systematic Biology , volume=58 , issue=2 , pages=224–39 , date=April 2009 , pmid=20525580 , doi=10.1093/sysbio/syp020 , doi-access=free{{cite journal , vauthors=Schüssler A, Schwarzott D, Walker C , s2cid=82128210 , year=2001 , title=A new fungal phylum, the Glomeromycota: phylogeny and evolution , journal=Mycological Research , volume=105 , issue=12 , pages=1413–1421 , doi=10.1017/S0953756201005196{{cite web , url=http://www.sci-news.com/biology/science-brazilian-stingless-bee-monascus-fungus-03372.html , title=Entomologists: Brazilian Stingless Bee Must Cultivate Special Type of Fungus to Survive , date=23 October 2015 , website=Sci-News.com , access-date=25 October 2015 , archive-url=https://web.archive.org/web/20151025012743/http://www.sci-news.com/biology/science-brazilian-stingless-bee-monascus-fungus-03372.html , archive-date=25 October 2015 , url-status=live{{cite journal , vauthors=Selosse MA, Richard F, He X, Simard SW , title=Mycorrhizal networks: des liaisons dangereuses? , journal=Trends in Ecology & Evolution , volume=21 , issue=11 , pages=621–8 , date=November 2006 , pmid=16843567 , doi=10.1016/j.tree.2006.07.003{{cite journal , vauthors=Schulz B, Boyle C , s2cid=23182632 , title=The endophytic continuum , journal=Mycological Research , volume=109 , issue=Pt 6 , pages=661–86 , date=June 2005 , pmid=16080390 , doi=10.1017/S095375620500273X{{cite journal , vauthors=Shoji JY, Arioka M, Kitamoto K , title=Possible involvement of pleiomorphic vacuolar networks in nutrient recycling in filamentous fungi , journal=Autophagy , volume=2 , issue=3 , pages=226–7 , year=2006 , pmid=16874107 , doi=10.4161/auto.2695 , doi-access=free{{cite journal , vauthors=Shalchian-Tabrizi K, Minge MA, Espelund M, Orr R, Ruden T, Jakobsen KS, Cavalier-Smith T , title=Multigene phylogeny of choanozoa and the origin of animals , journal=PLOS ONE , volume=3 , issue=5 , pages=e2098 , year=2008 , pmid=18461162 , pmc=2346548 , doi=10.1371/journal.pone.0002098 , doi-access=free , bibcode=2008PLoSO...3.2098S{{Cite book , vauthors=Silar P , title=Protistes Eucaryotes: Origine, Evolution et Biologie des Microbes Eucaryotes , publisher=HAL , year=2016 , page=462 , isbn=978-2-9555841-0-1 , url=https://hal.archives-ouvertes.fr/hal-01263138/document , access-date=7 April 2016 , archive-url=https://web.archive.org/web/20170925132023/https://hal.archives-ouvertes.fr/hal-01263138/document , archive-date=25 September 2017 , url-status=live{{cite book , vauthors=Simpson DP , title=Cassell's Latin Dictionary , publisher=Cassell Ltd , year=1979 , edition=5 , location=London, UK , page=883 , isbn=978-0-304-52257-6{{cite journal , vauthors=Simon-Nobbe B, Denk U, Pöll V, Rid R, Breitenbach M , title=The spectrum of fungal allergy , journal=International Archives of Allergy and Immunology , volume=145 , issue=1 , pages=58–86 , year=2008 , pmid=17709917 , doi=10.1159/000107578 , doi-access=free{{cite journal , vauthors=Steenkamp ET, Wright J, Baldauf SL , title=The protistan origins of animals and fungi , journal=Molecular Biology and Evolution , volume=23 , issue=1 , pages=93–106 , date=January 2006 , pmid=16151185 , doi=10.1093/molbev/msj011 , doi-access=free{{cite book , vauthors=Stamets P , title=Growing Gourmet and Medicinal Mushrooms , trans-title=Shokuyō oyobi yakuyō kinoko no saibai , publisher=Ten Speed Press , location=Berkeley, California , year=2000 , pages=233–248 , isbn=978-1-58008-175-7According to one 2001 estimate, some 10,000 fungal diseases are known. {{cite book , vauthors=Struck C , veditors=Cooke BM, Jones DG, Kaye B , title=The Epidemiology of Plant Diseases , publisher=Springer , location=Berlin, Germany , year=2006 , page=117 , isbn=978-1-4020-4580-6 , chapter=Infection strategies of plant parasitic fungi{{cite journal , vauthors=Sullivan R, Smith JE, Rowan NJ , s2cid=29723996 , title=Medicinal mushrooms and cancer therapy: translating a traditional practice into Western medicine , journal=Perspectives in Biology and Medicine , volume=49 , issue=2 , pages=159–70 , year=2006 , pmid=16702701 , doi=10.1353/pbm.2006.0034{{cite journal , vauthors=Taylor TN, Taylor EL , year=1996 , title=The distribution and interactions of some Paleozoic fungi , journal=Review of Palaeobotany and Palynology , volume=95 , issue=1–4 , pages=83–94 , doi=10.1016/S0034-6667(96)00029-2{{cite journal , vauthors=Taylor JW, Jacobson DJ, Kroken S, Kasuga T, Geiser DM, Hibbett DS, Fisher MC , s2cid=2551424 , title=Phylogenetic species recognition and species concepts in fungi , journal=Fungal Genetics and Biology , volume=31 , issue=1 , pages=21–32 , date=October 2000 , pmid=11118132 , doi=10.1006/fgbi.2000.1228{{cite journal , vauthors=Taylor TN, Hass H, Kerp H, Krings M, Hanlin RT , title=Perithecial ascomycetes from the 400 million year old Rhynie chert: an example of ancestral polymorphism , journal=Mycologia , volume=97 , issue=1 , pages=269–85 , year=2005 , pmid=16389979 , doi=10.3852/mycologia.97.1.269 , url=http://www.cybertruffle.org.uk/cyberliber/59350/0097/001/0269.htm , hdl=1808/16786 , access-date=5 July 2011 , archive-url=https://web.archive.org/web/20160412150211/http://www.cybertruffle.org.uk/cyberliber/59350/0097/001/0269.htm , archive-date=12 April 2016 , url-status=live , hdl-access=free{{cite journal , vauthors=Taylor JW, Berbee ML , title=Dating divergences in the Fungal Tree of Life: review and new analyses , journal=Mycologia , volume=98 , issue=6 , pages=838–49 , year=2006 , pmid=17486961 , doi=10.3852/mycologia.98.6.838 , url=http://www.cybertruffle.org.uk/cyberliber/59350/0098/006/0838.htm , access-date=5 July 2011 , archive-url=https://web.archive.org/web/20160412150130/http://www.cybertruffle.org.uk/cyberliber/59350/0098/006/0838.htm , archive-date=12 April 2016 , url-status=live{{cite journal , last1=Thambugala , first1=Kasun M. , last2=Daranagama , first2=Dinushani A. , last3=Phillips , first3=Alan J. L. , last4=Kannangara , first4=Sagarika D. , last5=Promputtha , first5=Itthayakorn , title=Fungi vs. fungi in biocontrol: An overview of fungal antagonists applied against fungal plant pathogens , journal=Frontiers in Cellular and Infection Microbiology , volume=10 , year=2020 , page=604923 , doi=10.3389/fcimb.2020.604923 , pmid=33330142 , pmc=7734056 , doi-access=free{{cite journal , vauthors=Thomas MB, Read AF , s2cid=14460348 , title=Can fungal biopesticides control malaria? , journal=Nature Reviews. Microbiology , volume=5 , issue=5 , pages=377–83 , date=May 2007 , pmid=17426726 , doi=10.1038/nrmicro1638 , hdl=1842/2089 , hdl-access=free{{cite journal , vauthors=Tudzynski B , s2cid=11191347 , title=Gibberellin biosynthesis in fungi: genes, enzymes, evolution, and impact on biotechnology , journal=Applied Microbiology and Biotechnology , volume=66 , issue=6 , pages=597–611 , date=March 2005 , pmid=15578178 , doi=10.1007/s00253-004-1805-1{{cite journal , last1=Tudzynski , first1=Bettina , year=2014 , title=Nitrogen regulation of fungal secondary metabolism in fungi , journal=Frontiers in Microbiology , volume=5 , page=656 , pmid=25506342 , pmc=4246892 , doi=10.3389/fmicb.2014.00656 , doi-access=free{{cite journal , vauthors=Trail F , title=Fungal cannons: explosive spore discharge in the Ascomycota , journal=FEMS Microbiology Letters , volume=276 , issue=1 , pages=12–8 , date=November 2007 , pmid=17784861 , doi=10.1111/j.1574-6968.2007.00900.x , doi-access=free{{cite book , author1=Ulloa, Miguel , author2=Halin, Richard T. , title=Illustrated Dictionary of Mycology , edition=2nd , year=2012 , publisher=The American Phytopathological Society , location=St. Paul, Minnesota , isbn=978-0-89054-400-6 , page=156{{cite web , url=http://tolweb.org/Fungi/2377 , title=Fungi. Eumycota: mushrooms, sac fungi, yeast, molds, rusts, smuts, etc. , vauthors=Blackwell M, Vilgalys R, James TY, Taylor JW , publisher=Tree of Life Web Project , year=2009 , access-date=25 April 2009 , archive-url=https://web.archive.org/web/20090413045121/http://tolweb.org/Fungi/2377 , archive-date=13 April 2009 , url-status=live{{cite web , url=http://www.ars.usda.gov/is/AR/archive/jul98/fung0798.htm , vauthors=Becker H , title=Setting the Stage To Screen Biocontrol Fungi , publisher=United States Department of Agriculture, Agricultural Research Service , year=1998 , access-date=23 February 2009 , archive-url=https://web.archive.org/web/20090116041447/http://www.ars.usda.gov/is/AR/archive/jul98/fung0798.htm , archive-date=16 January 2009 , url-status=live{{cite magazine , last1=Vargas-Gastélum , first1=Lluvia , last2=Riquelme , first2=Meritxell , title=The mycobiota of the deep sea: What omics can offer , magazine=Life (magazine), Life , volume=10 , issue=11 , year=2020 , pages=292 , doi=10.3390/life10110292 , pmid=33228036 , pmc=7699357 , doi-access=free{{cite journal , vauthors=Yang Y, Yang E, An Z, Liu X , title=Evolution of nematode-trapping cells of predatory fungi of the Orbiliaceae based on evidence from rRNA-encoding DNA and multiprotein sequences , journal=Proceedings of the National Academy of Sciences of the United States of America , volume=104 , issue=20 , pages=8379–84 , date=May 2007 , pmid=17494736 , pmc=1895958 , doi=10.1073/pnas.0702770104 , bibcode=2007PNAS..104.8379Y , doi-access=free{{cite journal , vauthors=Vetter J , title=Toxins of ''Amanita phalloides'' , journal=Toxicon , volume=36 , issue=1 , pages=13–24 , date=January 1998 , pmid=9604278 , doi=10.1016/S0041-0101(97)00074-3{{cite journal , vauthors=Vaupotic T, Veranic P, Jenoe P, Plemenitas A , title=Mitochondrial mediation of environmental osmolytes discrimination during osmoadaptation in the extremely halotolerant black yeast Hortaea werneckii , journal=Fungal Genetics and Biology , volume=45 , issue=6 , pages=994–1007 , date=June 2008 , pmid=18343697 , doi=10.1016/j.fgb.2008.01.006{{cite journal , last1=Walther , first1=Grit , last2=Wagner , first2=Lysett , last3=Kurzai , first3=Oliver , year=2019 , title=Updates on the taxonomy of Mucorales with an emphasis on clinically important taxa , journal=Journal of Fungi , volume=5 , issue=4 , page=106 , doi=10.3390/jof5040106 , pmc=6958464 , pmid=31739583, doi-access=free{{cite journal , vauthors=Wang ZY, Jenkinson JM, Holcombe LJ, Soanes DM, Veneault-Fourrey C, Bhambra GK, Talbot NJ , s2cid=7111935 , title=The molecular biology of appressorium turgor generation by the rice blast fungus ''Magnaporthe grisea'' , journal=Biochemical Society Transactions , volume=33 , issue=Pt 2 , pages=384–8 , date=April 2005 , pmid=15787612 , doi=10.1042/BST0330384{{cite journal , last1=Wang , first1=Ke , last2=Cai , first2=Lei , last3=Yao , first3=Yijian , title=Overview of nomenclature novelties of fungi in the world and China (2020) , journal=Biodiversity Science , volume=29 , issue=8 , year=2021 , doi=10.17520/biods.2021202 , pages=1064–1072 , s2cid=240568551{{cite journal , vauthors=Willensdorfer M , s2cid=39155292 , title=On the evolution of differentiated multicellularity , journal=Evolution; International Journal of Organic Evolution , volume=63 , issue=2 , pages=306–23 , date=February 2009 , pmid=19154376 , doi=10.1111/j.1558-5646.2008.00541.x , arxiv=0801.2610{{cite journal , vauthors=Ward PD, Botha J, Buick R, De Kock MO, Erwin DH, Garrison GH, Kirschvink JL, Smith R , s2cid=46198018 , title=Abrupt and gradual extinction among Late Permian land vertebrates in the Karoo basin, South Africa , journal=Science (journal), Science , volume=307 , issue=5710 , pages=709–14 , date=February 2005 , pmid=15661973 , doi=10.1126/science.1107068 , bibcode=2005Sci...307..709W , citeseerx=10.1.1.503.2065{{cite journal , vauthors=Xu H, Andi B, Qian J, West AH, Cook PF , s2cid=22370361 , title=The alpha-aminoadipate pathway for lysine biosynthesis in fungi , journal=Cell Biochemistry and Biophysics , volume=46 , issue=1 , pages=43–64 , year=2006 , pmid=16943623 , doi=10.1385/CBB:46:1:43{{cite journal , vauthors=Wu S, Schalk M, Clark A, Miles RB, Coates R, Chappell J , s2cid=23358348 , title=Redirection of cytosolic or plastidic isoprenoid precursors elevates terpene production in plants , journal=Nature Biotechnology , volume=24 , issue=11 , pages=1441–7 , date=November 2006 , pmid=17057703 , doi=10.1038/nbt1251{{cite journal , vauthors=Zabriskie TM, Jackson MD , title=Lysine biosynthesis and metabolism in fungi , journal=Natural Product Reports , volume=17 , issue=1 , pages=85–97 , date=February 2000 , pmid=10714900 , doi=10.1039/a801345d{{cite journal , last1=Zhuo , first1=Rui , last2=Fan , first2=Fangfang , title=A comprehensive insight into the application of white rot fungi and their lignocellulolytic enzymes in the removal of organic pollutants , journal=Science of the Total Environment , volume=778 , year=2021 , pages=146132 , doi=10.1016/j.scitotenv.2021.146132 , pmid=33714829 , bibcode=2021ScTEn.778n6132Z , s2cid=232230208{{cite journal , vauthors=Li Y, Steenwyk JL, Chang Y, Wang Y, James TY, Stajich JE, Spatafora JW, Groenewald M, Dunn CW, Hittinger CT, Shen X, Rokas, A , date=2021 , title=A genome-scale phylogeny of the kingdom Fungi , journal=Current Biology , volume=31 , issue=8 , pages=1653–1665 , doi=10.1016/j.cub.2021.01.074 , pmid=33607033 , pmc=8347878
Cited literature
{{refbegin
* {{cite book , vauthors=Ainsworth GC , title=Introduction to the History of Mycology , publisher=Cambridge University Press , location=Cambridge, UK , year=1976 , isbn=978-0-521-11295-6
* {{cite book , vauthors=Alexopoulos CJ, Mims CW, Blackwell M , title=Introductory Mycology , year=1996 , publisher=John Wiley & Sons , isbn=978-0-471-52229-4
* {{cite book , vauthors=Deacon J , title=Fungal Biology , publisher=Blackwell Publishers , location=Cambridge, Massachusetts , year=2005 , isbn=978-1-4051-3066-0
* {{cite book , vauthors=Hall IR , title=Edible and Poisonous Mushrooms of the World , publisher=Timber Press , location=Portland, Oregon , year=2003 , isbn=978-0-88192-586-9
* {{cite book , vauthors=Hanson JR , title=The Chemistry of Fungi , year=2008 , publisher=Royal Society of Chemistry , isbn=978-0-85404-136-7
* {{cite book , vauthors=Jennings DH, Lysek G , title=Fungal Biology: Understanding the Fungal Lifestyle , year=1996 , publisher=Bios Scientific Publishers Ltd , location=Guildford, UK , isbn=978-1-85996-150-6
* {{cite book , vauthors=Kirk PM, Cannon PF, Minter DW, Stalpers JA , title=Dictionary of the Fungi , edition=10th , publisher=CAB International , location=Wallingford, UK , year=2008 , isbn=978-0-85199-826-8
* {{cite book , vauthors=Taylor EL, Taylor TN , title=The Biology and Evolution of Fossil Plants , publisher=Prentice Hall , location=Englewood Cliffs, New Jersey , year=1993 , isbn=978-0-13-651589-0
{{refend
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