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A fungus (
plural The plural (sometimes abbreviated An abbreviation (from Latin ''brevis'', meaning ''short'') is a shortened form of a word or phrase, by any method. It may consist of a group of letters, or words taken from the full version of the word or ph ...

plural
: fungi or funguses) is any member of the group of
eukaryotic Eukaryotes () are organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy (biology), tax ...

eukaryotic
organisms that includes microorganisms such as
yeast Yeasts are eukaryotic Eukaryotes () are organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classifie ...

yeast
s and
mold A mold () or mould () is a fungus A fungus (plural: fungi or funguses) is any member of the group of Eukaryote, eukaryotic organisms that includes microorganisms such as yeasts and Mold (fungus), molds, as well as the more familiar mush ...
s, as well as the more familiar
mushroom A mushroom or toadstool is the fleshy, spore )'', growing on a thinning, thinned hybrid black poplar ''(populus, Populus x canadensis)''. The last stage of the moss#Life cycle, moss lifecycle is shown, where the sporophytes are visible befor ...

mushroom
s. These organisms are classified as a
kingdom Kingdom may refer to: Monarchy * A type of monarchy * A realm ruled by: **A king, during the reign of a male monarch **A queen regnant, during the reign of a female monarch Taxonomy * Kingdom (biology), a category in biological taxonomy Arts an ...
, separately from the other eukaryotic kingdoms, which by one traditional classification include
Plant Plants are predominantly photosynthetic Photosynthesis is a process used by plants and other organisms to Energy transformation, convert light energy into chemical energy that, through cellular respiration, can later be released to fuel ...

Plant
ae,
Animal Animals (also called Metazoa) are organisms that form the Animalia. With few exceptions, animals , , are , can , and grow from a hollow sphere of , the , during . Over 1.5 million animal have been —of which around 1 million are —b ...

Animal
ia,
Protozoa Protozoa (singular protozoon or protozoan, plural protozoa or protozoans) is an informal term for a group of Unicellular organism, single-celled eukaryotes, either free-living or Parasitism, parasitic, that feed on organic matter such as other mi ...

Protozoa
, and
Chromista Chromista is a biological kingdom consisting of single-celled and multicellular eukaryotic Eukaryotes () are organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual conti ...
. A characteristic that places fungi in a different kingdom from plants, bacteria, and some protists is
chitin units that repeat to form long chains in β-(1→4)-linkage. of the chitin molecule. Chitin ( carbon, C8H13O5N)n ( ) is a long-chain polymer A polymer (; Greek ''wikt:poly-, poly-'', "many" + ''wikt:-mer, -mer'', "part") is a Chemical su ...

chitin
in their
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane cell membrane vs. Prokaryotes The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to a ...
s. Fungi, like animals, are
heterotroph A heterotroph (; from Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following peri ...
s; they acquire their food by absorbing dissolved molecules, typically by secreting
digestive enzyme Digestive may refer to: Biology *Digestion Digestion is the breakdown of large insoluble food Food is any substance consumed to provide nutritional support for an organism In biology, an organism (from Ancient Greek, Greek: ὀργ ...
s into their environment. Fungi do not
photosynthesize upright=1.8, Overall equation for the type of photosynthesis that occurs in plants Photosynthesis is a process used by plants and other organisms to convert Conversion or convert may refer to: Arts, entertainment, and media * Conversion (Docto ...

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 contempo ...

mobility
, except for
spore )'', growing on a thinning, thinned hybrid black poplar ''(populus, Populus x canadensis)''. The last stage of the moss#Life cycle, moss lifecycle is shown, where the sporophytes are visible before dispersion of their spores: the calyptra (1) is ...
s (a few of which are
flagellated A flagellum (; plural: flagella) is a lash-like appendage that protrudes from the soma (biology), cell body of certain cells termed as flagellates. A flagellate can have one or several flagella. The primary function of a flagellum is that of ani ...
), which may travel through the air or water. Fungi are the principal
decomposer Decomposers are Organism, organisms that break down dead or decaying organisms; they carry out decomposition, a process possible by only certain kingdoms, such as fungi. Like herbivores and predators, decomposers are heterotrophic, meaning that th ...
s 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 Common descent is a concept in evolutionary biology Evolutionary biology is the subfield of biology that studies the evolution, evolutionary processes (natural selection, common descent, speciation) that produced the Biodiversity, diversity ...
(i.e. they form a ''
monophyletic 300px, A cladogram of the primates, showing a ''monophyletic'' taxon: ''the simians'' (in yellow); a ''paraphyletic'' taxon: ''the prosimians'' (in cyan, including the red patch); and a ''polyphyletic'' group: ''the night-active primates, i.e., ...

monophyletic
group''), an interpretation that is also strongly supported by
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram A diagram is a symb ...
. This fungal group is distinct from the structurally similar myxomycetes (slime molds) and
oomycete Oomycota or oomycetes () form a distinct phylogenetic In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, mole ...
s (water molds). The discipline of
biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanisms, Development ...

biology
devoted to the study of fungi is known as
mycology Mycology is the branch of biology concerned with the study of fungus, fungi, including their genetics, genetic and biochemistry, biochemical properties, their Taxonomy (biology), taxonomy and ethnomycology, their use to humans as a source for ti ...
(from the
Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approximately 10.7 million as of ...
μύκης ', 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 wo ...

botany
, although it is now known fungi are genetically more closely related to animals than to plants. Abundant worldwide, most fungi are inconspicuous because of the small size of their structures, and their cryptic lifestyles in soil or on dead matter. Fungi include
symbionts Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction In ecology Ecology (from el, οἶκος, "house" and el, -λογία, label=none, ...

symbionts
of plants, animals, or other fungi and also
parasites Parasitism is a symbiotic Symbiosis (from Ancient Greek, Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different Organism, biological or ...
. 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 called bicycling or biking, is the use of bicycles for transport, recreation, Physical exercise, exercise or sport. People engaged in cycling are referred to as "cyclists", "bicyclists", or "bikers". Apart from two-wheeled bicy ...
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 (fungus), Tuber''. In addition to ''Tuber'', many other genera of fungi are classified as truffles including ''Geopor ...
; as a leavening agent for bread; and in the
fermentation Fermentation is a metabolism, metabolic process that produces chemical changes in organic Substrate (chemistry), substrates through the action of enzymes. In biochemistry, it is narrowly defined as the extraction of energy from carbohydrates in ...
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 ...

wine
,
beer Beer is one of the oldest and most widely consumed alcoholic drink An alcoholic drink is a drink A drink (or beverage) is a liquid A liquid is a nearly incompressible fluid In physics, a fluid is a substance that continual ...

beer
, and
soy sauce Soy sauce (also called simply soy in American English American English (AmE, AE, AmEng, USEng, en-US), sometimes called United States English or U.S. English, is the set of varieties of the English language native to the United States. C ...

soy sauce
. 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 pathogenic bacteria, bacterial infections, and antibiotic medications are widely used in the therapy, ...
s, and, more recently, various
enzyme Enzymes () are protein Proteins are large s and s that comprise one or more long chains of . Proteins perform a vast array of functions within organisms, including , , , providing and , and from one location to another. Proteins diff ...

enzyme
s produced by fungi are used industrially and in detergents. Fungi are also used as
biological pesticideBiopesticides, a contraction of 'biological pesticides', include several types of pest management intervention: through predatory, parasitic, or chemical relationships. The term has been associated historically with biological pest control – and by ...
s to control weeds, plant diseases and insect pests. Many species produce bioactive compounds called
mycotoxin A mycotoxin (from the Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is appr ...
s, such as
alkaloid Alkaloids are a class of base (chemistry), basic, natural product, 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. Som ...
s and
polyketidePolyketides are a large group of secondary metabolites which either contain alternating carbonyl groups and methylene groups (-CO-CH2-), or are derived from precursors which contain such alternating groups. Many polyketides are medicinal or exhibit a ...
s, that are toxic to animals including humans. The fruiting structures of a few species contain
psychotropic A psychoactive drug, psychopharmaceutical, or psychotropic drug is a chemical substance that changes nervous system In Biology, biology, the nervous system is a Complex system, highly complex part of an animal that coordinates its Behavior, ...
compounds and are consumed recreationally or in traditional spiritual ceremonies. Fungi can break down manufactured materials and buildings, and become significant
pathogens In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanisms, ...
of humans and other animals. Losses of crops due to fungal diseases (e.g., rice blast disease) or food spoilage can have a large impact on human food supplies and local economies. The fungus kingdom encompasses an enormous diversity of
taxa In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanism ...
with varied ecologies,
life cycle Life cycle, life-cycle, or lifecycle may refer to: Science and academia *Biological life cycle, the sequence of life stages that an organism undergoes from birth to reproduction ending with the production of the offspring *Life-cycle hypothesis, ...
strategies, and morphologies ranging from unicellular aquatic
chytrid Chytridiomycota are a division of zoosporic organisms in the kingdom (biology), kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoos ...
s to large mushrooms. However, little is known of the true
biodiversity Biodiversity is the biological variety and of . Biodiversity is a measure of variation at the , , and level. Terrestrial biodiversity is usually greater near the , which is the result of the warm and high . Biodiversity is not distributed ev ...

biodiversity
of Kingdom Fungi, 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 Nobility#Ennoblement, ennoblement as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomi ...

Carl Linnaeus
,
Christiaan Hendrik Persoon Christiaan Hendrik Persoon (1 February 1761 – 16 November 1836) was a German mycologist who made additions to Carl Linnaeus, Linnaeus' mushroom Taxonomy (biology), taxonomy. Early life Persoon was born in South Africa at the Cape of Good ...

Christiaan Hendrik Persoon
, and
Elias Magnus Fries Elias Magnus Fries (15 August 1794 – 8 February 1878) was a Swedish mycologist Mycology is the branch of biology concerned with the study of fungus, fungi, including their genetics, genetic and biochemistry, biochemical properties, th ...

Elias Magnus Fries
, fungi have been classified according to their morphology (e.g., characteristics such as spore color or microscopic features) or
physiology Physiology (; ) is the scientific Science () is a systematic enterprise that builds and organizes knowledge Knowledge is a familiarity or awareness, of someone or something, such as facts A fact is an occurrence in the real world. ...
. 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 The structure of part of a DNA double helix Deoxyribonucleic acid (; DNA) is a molecule File:Pentacene on Ni(111) STM.jpg, A scanning tunneling microscopy i ...

DNA analysis
to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits.
Phylogenetic In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanisms, ...

Phylogenetic
studies published in the first decade of the 21st century have helped reshape the classification within Kingdom Fungi, which is divided into one
subkingdom In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanisms, ...
, seven phyla, and ten subphyla.


Etymology

The English word ''fungus'' is directly adopted from the
Latin Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communication Communication (from Latin ''communicare'', meaning "to share" or "to be in relation with") is "an appa ...

Latin
''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 Roman or Romans usually refers to: *Rome, the capital city of Italy *Ancient Rome, Roman civilization from ...

Horace
and
Pliny Pliny may refer to: People from antiquity * Pliny the Elder (AD 23–79), ancient Roman nobleman, scientist, historian, and author of ''Naturalis Historia'' (''Pliny's Natural History'') * Pliny the Younger (died 113), ancient Roman statesman, ...

Pliny
. This in turn is derived from the
Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approximately 10.7 million as of ...
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 The microscopic scale (from , ''mikrós'', "sm ...
structures and morphology of mushrooms and molds;Ainsworth, p. 2. the
root In vascular plant Vascular plants (from Latin ''vasculum'': duct), also known as Tracheophyta (the tracheophytes , from Greek τραχεῖα ἀρτηρία ''trācheia artēria'' 'windpipe' + φυτά ''phutá'' 'plants'), form a large group ...
is also used in other languages, such as the German '''' ('sponge') and '''' ('mold'). The word ''
mycology Mycology is the branch of biology concerned with the study of fungus, fungi, including their genetics, genetic and biochemistry, biochemical properties, their Taxonomy (biology), taxonomy and ethnomycology, their use to humans as a source for ti ...
'' 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 Carl Linnaeus, Linnaeus' mushroom Taxonomy (biology), taxonomy. Early life Persoon was born in South Africa at the Cape of Good ...

Christiaan Hendrik Persoon
. The word appeared in English as early as 1824 in a book by . 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 Benefield#Biggin Hall, Biggin Hall, Benefield, Northamptonshir ...

Miles Joseph Berkeley
'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'' (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'' has been proposed as a less ambiguous term similar in use to
fauna Fauna is all of the animal Animals (also called Metazoa) are multicellular eukaryotic organisms that form the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular r ...

fauna
and
flora Flora is all the plant life present in a particular region or time, generally the naturally occurring (indigenous (ecology), indigenous) native plant, native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms ' ...

flora
. The Species Survival Commission (SSC) of the
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 working in the field of and of s. It is involved in data gathering and analysis, research, field p ...
(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 for phylogenetic analysis,
taxonomists In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mecha ...
considered fungi to be members of the
plant kingdom Plants are mainly multicellular Multicellular organisms are organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual contiguous system that embodies the Life#Biology, prope ...

plant kingdom
because of similarities in lifestyle: both fungi and plants are mainly immobile, and have similarities in general morphology and growth habitat. Like plants, fungi often grow in soil and, in the case of
mushroom A mushroom or toadstool is the fleshy, spore )'', growing on a thinning, thinned hybrid black poplar ''(populus, Populus x canadensis)''. The last stage of the moss#Life cycle, moss lifecycle is shown, where the sporophytes are visible befor ...

mushroom
s, form conspicuous
fruit bodies In fungi A fungus (plural The plural (sometimes list of glossing abbreviations, abbreviated ), in many languages, is one of the values of the grammatical number, grammatical category of number. The plural of a noun typically denotes a qua ...
, which sometimes resemble plants such as
mosses Mosses are small, non-vascular flower A flower, sometimes known as a bloom or blossom, is the reproduction, reproductive structure found in flowering plants (plants of the division Magnoliophyta, also called angiosperms). The biological func ...

mosses
. The fungi are now considered a separate kingdom, distinct from both plants and animals, from which they appear to have diverged around one billion years ago (around the start of the
Neoproterozoic The Neoproterozoic Era is the unit of geologic time The geologic time scale (GTS) is a system of chronological datingChronological dating, or simply dating, is the process of attributing to an object or event a date in the past, allowing such ob ...
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 organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy (biology), tax ...

eukaryote
s: Fungal cells contain
membrane-bound A biological membrane, biomembrane or cell membrane is a selectively permeable membrane up150px, Schematic of size-based membrane exclusion A membrane is a selective barrier; it allows some things to pass through but stops others. Such things ...
nuclei ''Nucleus'' (plural nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA ...

nuclei
with
chromosomes A chromosome is a long DNA molecule with part or all of the genome, genetic material of an organism. Most eukaryotic chromosomes include packaging proteins called histones which, aided by Chaperone (protein), chaperone proteins, bind to and D ...
that contain
DNA Deoxyribonucleic acid (; DNA) is a molecule File:Pentacene on Ni(111) STM.jpg, A scanning tunneling microscopy image of pentacene molecules, which consist of linear chains of five carbon rings. A molecule is an electrically neutral gro ...

DNA
with noncoding regions called
intron An intron (for ''intragenic region'') is any nucleotide sequence A nucleic acid sequence is a succession of bases signified by a series of a set of five different letters that indicate the order of nucleotides Nucleotides are organic molecul ...

intron
s and coding regions called
exons An exon is any part of a gene In biology, a gene (from ''genos'' "...Wilhelm Johannsen coined the word gene to describe the Mendelian_inheritance#History, Mendelian units of heredity..." (Greek language, Greek) meaning ''generation'' or ...
. Fungi have membrane-bound cytoplasmic
organelles In cell biology Cell biology (also cellular biology or cytology) is a branch of biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, ...
such as
mitochondria A mitochondrion (; ) is a double-membrane Image:Schematic size.jpg, up150px, Schematic of size-based membrane exclusion A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, i ...

mitochondria
,
sterol Sterol is an organic compound In , organic compounds are generally any s that contain - . Due to carbon's ability to (form chains with other carbon s), millions of organic compounds are known. The study of the properties, reactions, and synth ...

sterol
-containing membranes, and
ribosomes Ribosomes () are molecular machine, macromolecular machines, found within all living cell (biology), cells, that perform Translation (biology), biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order spec ...

ribosomes
of the
80S Ribosome Ribosomes () are macromolecular machines, found within all living cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids Amino acids are organic compounds that contain amino (–NH2) and ...
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 compound , CH4; is among the simplest organic compounds. In chemistry Chemistry is the scientific discipline involved with Chemical element, e ...
s (e.g.,
mannitol Mannitol is a type of sugar alcohol Sugar alcohols (also called polyhydric alcohols, polyalcohols, alditols or glycitols) are organic compound , CH4; is among the simplest organic compounds. In chemistry Chemistry is the scientific dis ...
),
disaccharide A disaccharide (also called a double sugar or ''biose'') is the sugar Sugar is the generic name for sweet-tasting, soluble carbohydrate is a disaccharide found in animal milk. It consists of a molecule of D-galactose and a molecule of ...
s, (e.g.,
trehalose Trehalose (from Turkish 'trehala' – a sugar derived from insect cocoons + -ose) is a sugar Sugar is the generic name for Sweetness, sweet-tasting, soluble carbohydrates, many of which are used in food. Table sugar, granulated sugar, or regu ...

trehalose
), and
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant found in . They are long chain carbohydrates composed of units bound together by . This carbohydrate can react with water () using as catalyst, which produces constituent sugars ...
s (e.g.,
glycogen Glycogen is a multibranched of that serves as a form of energy storage in s, , and bacteria. The polysaccharide structure represents the main storage form of glucose in the body. Glycogen functions as one of two forms of energy reserves, g ...

glycogen
, which is also found in animalsDeacon, pp. 128–129.). * With animals: Fungi lack
chloroplast A chloroplast is a type of membrane-bound organelle In cell biology, an organelle is a specialized subunit, usually within a cell (biology), cell, that has a specific function. The name ''organelle'' comes from the idea that these structure ...

chloroplast
s and are
heterotroph A heterotroph (; from Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following peri ...
ic organisms and so require preformed
organic compound In chemistry Chemistry is the study of the properties and behavior of . It is a that covers the that make up matter to the composed of s, s and s: their composition, structure, properties, behavior and the changes they undergo during ...
s as energy sources. * With plants: Fungi have a cell wall and
vacuole A vacuole () is a membrane-bound organelle In cell biology, an organelle is a specialized subunit, usually within a cell (biology), cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts o ...
s. They reproduce by both sexual and asexual means, and like
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is neces ...
plant groups (such as
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plant Vascular plants (from Latin ''vasculum'': duct), also known as Tracheophyta (the tracheophytes , from Greek τραχεῖα ἀρτηρία ''trācheia art ...

fern
s and
moss Mosses are small, non-vascular plant, non-vascular flowerless plants in the taxonomic phylum, division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Wilhelm Philippe Schimper, Schimp. 1879) may also refer to the parent group bryo ...

moss
es) produce
spore )'', growing on a thinning, thinned hybrid black poplar ''(populus, Populus x canadensis)''. The last stage of the moss#Life cycle, moss lifecycle is shown, where the sporophytes are visible before dispersion of their spores: the calyptra (1) is ...
s. Similar to mosses and algae, fungi typically have
haploid Ploidy () is the number of complete sets of chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. Most eukaryotic chromosomes include packaging proteins called histones which, aided by ...
nuclei.Deacon, p. 58. * With
euglenoid Euglenids (euglenoids, or euglenophytes, formally Euglenida/Euglenoida, International Code of Zoological Nomenclature, ICZN, or Euglenophyceae, ICBN) are one of the best-known groups of flagellates, which are Excavata, excavate eukaryotes of the ph ...
s and bacteria: Higher fungi, euglenoids, and some bacteria produce the
amino acid Amino acids are organic compound In , organic compounds are generally any s that contain - . Due to carbon's ability to (form chains with other carbon s), millions of organic compounds are known. The study of the properties, reactions, a ...

amino acid
L-lysine in specific
biosynthesis Biosynthesis is a multi-step, enzyme-Catalysis, catalyzed process where substrate (chemistry), substrates are converted into more complex Product (chemistry), products in living organisms. In biosynthesis, simple Chemical compound, compounds are mod ...

biosynthesis
steps, called the α-aminoadipate pathway. * The cells of most fungi grow as tubular, elongated, and thread-like (filamentous) structures called
hypha Hyphae of '' Penicillium'' File:Aspergillus niger 01.jpg, ''Aspergillus niger'' 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 ...
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 of ...
—cellular structures consisting of
protein Proteins are large biomolecule , showing alpha helices, represented by ribbons. This poten was the first to have its suckture solved by X-ray crystallography by Max Perutz and Sir John Cowdery Kendrew in 1958, for which they received a No ...

protein
s,
lipid In and , a lipid is a macro that is soluble in solvents. are typically s used to dissolve other naturally occurring hydrocarbon lipid s that do not (or do not easily) dissolve in water, including s, es, s, fat-soluble s (such as vitamins A, ...
s, and other organic molecules—called the
Spitzenkörper The Spitzenkörper (German German(s) may refer to: Common uses * of or related to Germany * Germans, Germanic ethnic group, citizens of Germany or people of German ancestry * For citizens of Germany, see also German nationality law * German l ...
. Both fungi and
oomycete Oomycota or oomycetes () form a distinct phylogenetic In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, mole ...
s grow as filamentous hyphal cells. In contrast, similar-looking organisms, such as filamentous
green algae The green algae (singular: green alga) are a large, informal grouping of algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic Photosynthesis is a process used by plants and other organisms to ...

green algae
, grow by repeated cell division within a chain of cells. There are also single-celled fungi (
yeast Yeasts are eukaryotic Eukaryotes () are organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classifie ...

yeast
s) that do not form hyphae, and some fungi have both hyphal and yeast forms.Alexopoulos ''et al''., p. 30. * In common with some plant and animal species, more than 70 fungal species display bioluminescence. Unique features: * Some species grow as unicellular yeasts that reproduce by budding or binary fission, fission. Dimorphic fungi can switch between a yeast phase and a hyphal phase in response to environmental conditions. * The fungal cell wall is made of glucans and
chitin units that repeat to form long chains in β-(1→4)-linkage. of the chitin molecule. Chitin ( carbon, C8H13O5N)n ( ) is a long-chain polymer A polymer (; Greek ''wikt:poly-, poly-'', "many" + ''wikt:-mer, -mer'', "part") is a Chemical su ...

chitin
; while glucans are also found in plants and chitin in the exoskeleton of arthropods, fungi are the only organisms that combine these two structural molecules in their cell wall. Unlike those of plants and oomycetes, fungal cell walls do not contain cellulose. Most fungi lack an efficient system for the long-distance transport of water and nutrients, such as the xylem and phloem in many plants. To overcome this limitation, some fungi, such as ''Armillaria'', form Mycelial cord, rhizomorphs, which resemble and perform functions similar to the roots of plants. As eukaryotes, fungi possess a Biochemical pathway, biosynthetic pathway for producing terpenes that uses mevalonic acid and pyrophosphate as Precursor (chemistry), 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 metabolites that are similar or identical in structure to those made by plants. Many of the plant and fungal enzymes that make these compounds differ from each other in peptide sequence, sequence and other characteristics, which indicates separate origins and convergent evolution 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 desert fungi, deserts or areas with high salt concentrations or ionizing radiation, as well as in deep sea sediments. Some can survive the intense ultraviolet radiation, UV and cosmic radiation encountered during space travel. Most grow in terrestrial environments, though several species live partly or solely in aquatic habitats, such as the
chytrid Chytridiomycota are a division of zoosporic organisms in the kingdom (biology), kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoos ...
fungi ''Batrachochytrium dendrobatidis'' and ''Batrachochytrium salamandrivorans, B. salamandrivorans'', parasites that have been responsible for a worldwide decline in amphibian 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 areas of the ocean. As of 2020, around 148,000 species of fungi have been species description, described by
taxonomists In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mecha ...
, 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. In mycology, species have historically been distinguished by a variety of methods and concepts. Classification based on morphology (biology), 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 Biochemistry, biochemical and physiology, physiological characteristics, such as their ability to metabolize certain biochemicals, or their reaction to Chemical tests in mushroom identification, chemical tests. The Species#The isolation species concept in more detail, biological species concept discriminates species based on their ability to mating in fungi, mate. The application of Molecular biology, molecular tools, such as DNA sequencing and phylogenetic analysis, to study diversity has greatly enhanced the resolution and added robustness to estimates of genetic diversity within various taxonomic groups.


Mycology

Mycology is the branch of
biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanisms, Development ...

biology
concerned with the systematic study of fungi, including their genetic and biochemical properties, their taxonomy, and their use to humans as a source of medicine, food, and entheogen, psychotropic substances consumed for religious purposes, as well as their dangers, such as poisoning or infection. The field of phytopathology, the study of plant diseases, is closely related because many plant pathogens are fungi. The use of fungi by humans dates back to prehistory; Ötzi the Iceman, a well-preserved mummy of a 5,300-year-old Neolithic man found frozen in the Austrian Alps, carried two species of polypore mushrooms that may have been used as tinder (''Fomes fomentarius''), or for medicinal purposes (''Piptoporus betulinus''). Ancient peoples have used fungi as food sources—often unknowingly—for millennia, in the preparation of leavened bread and fermented juices. Some of the oldest written records contain references to the destruction of crops that were probably caused by pathogenic fungi.


History

Mycology is a relatively new science that became systematic after the development of the microscope in the 17th century. Although fungal spores were first observed by Giambattista della Porta in 1588, the seminal work in the development of mycology is considered to be the publication of Pier Antonio Micheli's 1729 work ''Nova plantarum genera''. 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 nomenclature, binomial system of nomenclature introduced by
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomi ...

Carl Linnaeus
in his ''Species plantarum'' (1753), the Dutch
Christiaan Hendrik Persoon Christiaan Hendrik Persoon (1 February 1761 – 16 November 1836) was a German mycologist who made additions to Carl Linnaeus, Linnaeus' mushroom Taxonomy (biology), taxonomy. Early life Persoon was born in South Africa at the Cape of Good ...

Christiaan Hendrik Persoon
(1761–1836) established the first classification of mushrooms with such skill as to be considered a founder of modern mycology. Later,
Elias Magnus Fries Elias Magnus Fries (15 August 1794 – 8 February 1878) was a Swedish mycologist Mycology is the branch of biology concerned with the study of fungus, fungi, including their genetics, genetic and biochemistry, biochemical properties, th ...

Elias Magnus Fries
(1794–1878) further elaborated the Biological classification, classification of fungi, using spore color and microscopic characteristics, methods still used by taxonomists today. Other notable early contributors to mycology in the 17th–19th and early 20th centuries include
Miles Joseph Berkeley 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 Benefield#Biggin Hall, Biggin Hall, Benefield, Northamptonshir ...

Miles Joseph Berkeley
, August Carl Joseph Corda, Anton de Bary, the brothers Louis René Tulasne, Louis René and Charles Tulasne, Arthur Henry Reginald Buller, Arthur H. R. Buller, Curtis Gates Lloyd, Curtis G. Lloyd, and Pier Andrea Saccardo. In the 20th and 21st centuries, advances in biochemistry, genetics, molecular biology, biotechnology, DNA sequencing and phylogenetic analysis has provided new insights into fungal relationships and
biodiversity Biodiversity is the biological variety and of . Biodiversity is a measure of variation at the , , and level. Terrestrial biodiversity is usually greater near the , which is the result of the warm and high . Biodiversity is not distributed ev ...

biodiversity
, and has challenged traditional morphology-based groupings in fungal Taxonomy (biology), taxonomy.


Morphology


Microscopic structures

Most fungi grow as
hypha Hyphae of '' Penicillium'' File:Aspergillus niger 01.jpg, ''Aspergillus niger'' 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 ...
e, which are cylindrical, thread-like structures 2–10micrometres, µm in diameter and up to several centimeters in length. Hyphae grow at their tips (apices); new hyphae are typically formed by emergence of new tips along existing hyphae by a process called ''branching'', or occasionally growing hyphal tips fork, giving rise to two parallel-growing hyphae. Hyphae also sometimes fuse when they come into contact, a process called hyphal fusion (or anastomosis). These growth processes lead to the development of a mycelium, an interconnected network of hyphae. Hyphae can be either septum, septate or coenocytic. Septate hyphae are divided into compartments separated by cross walls (internal cell walls, called septa, that are formed at right angles to the cell wall giving the hypha its shape), with each compartment containing one or more nuclei; coenocytic hyphae are not compartmentalized. Septa have Pit connection#Characteristics, pores that allow cytoplasm, organelles, and sometimes nuclei to pass through; an example is the dolipore septum in fungi of the phylum Basidiomycota. Coenocytic hyphae are in essence multinucleate supercells. Many species have developed specialized hyphal structures for nutrient uptake from living hosts; examples include haustoria in plant-parasitic species of most fungal phyla, and arbuscular mycorrhiza, arbuscules of several mycorrhizal fungi, which penetrate into the host cells to consume nutrients. Although fungi are opisthokonts—a grouping of evolutionarily related organisms broadly characterized by a single posterior flagellum—all phyla except for the chytrids have lost their posterior flagella. Fungi are unusual among the eukaryotes in having a cell wall that, in addition to glucans (e.g., Beta-glucan, β-1,3-glucan) and other typical components, also contains the biopolymer chitin.


Macroscopic structures

Fungal mycelia can become visible to the naked eye, for example, on various surfaces and substrate (biology), substrates, such as damp walls and spoiled food, where they are commonly called
mold A mold () or mould () is a fungus A fungus (plural: fungi or funguses) is any member of the group of Eukaryote, eukaryotic organisms that includes microorganisms such as yeasts and Mold (fungus), molds, as well as the more familiar mush ...
s. Mycelia grown on solid agar media in laboratory petri dishes are usually referred to as colony (biology), colonies. These colonies can exhibit growth shapes and colors (due to spores or biological pigment, 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 Clone (cell biology), clonal colony of ''Armillaria solidipes'', which extends over an area of more than 900hectare, ha (3.5 square miles), with an estimated age of nearly 9,000years. The apothecium—a specialized structure important in sexual reproduction in the ascomycetes—is a cup-shaped fruit body that is often macroscopic and holds the hymenium, a layer of tissue containing the spore-bearing cells. The fruit bodies of the basidiomycetes (basidiocarps) and some ascomycetes can sometimes grow very large, and many are well known as
mushroom A mushroom or toadstool is the fleshy, spore )'', growing on a thinning, thinned hybrid black poplar ''(populus, Populus x canadensis)''. The last stage of the moss#Life cycle, moss lifecycle is shown, where the sporophytes are visible befor ...

mushroom
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 substrate (biology), substrates and tissues. They can exert large penetrative mechanical forces; for example, many plant pathogens, including ''Magnaporthe grisea'', form a structure called an appressorium that evolved to puncture plant tissues. The pressure generated by the appressorium, directed against the plant Epidermis (botany), epidermis, can exceed . The filamentous fungus ''Paecilomyces lilacinus'' uses a similar structure to penetrate the eggs of nematodes. The mechanical pressure exerted by the appressorium is generated from physiological processes that increase intracellular turgor by producing osmolytes such as glycerol. Adaptations such as these are complemented by cellulase, hydrolytic enzymes secreted into the environment to digest large organic molecules—such as
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant found in . They are long chain carbohydrates composed of units bound together by . This carbohydrate can react with water () using as catalyst, which produces constituent sugars ...
s,
protein Proteins are large biomolecule , showing alpha helices, represented by ribbons. This poten was the first to have its suckture solved by X-ray crystallography by Max Perutz and Sir John Cowdery Kendrew in 1958, for which they received a No ...

protein
s, and
lipid In and , a lipid is a macro that is soluble in solvents. are typically s used to dissolve other naturally occurring hydrocarbon lipid s that do not (or do not easily) dissolve in water, including s, es, s, fat-soluble s (such as vitamins A, ...
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 Endophyte, endophytic fungi, or growth by volume expansion during the development of mushroom stipe (mycology), stipes and other large organs. Growth of fungi as Multicellularity, multicellular structures consisting of Somatic (biology), somatic and reproductive cells—a feature independently evolved in animals and plants—has several functions, including the development of fruit bodies for dissemination of sexual spores (see above) and biofilms for substrate colonization and intercellular communication. The fungi are traditionally considered
heterotroph A heterotroph (; from Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following peri ...
s, organisms that rely solely on carbon fixation, carbon fixed by other organisms for metabolism. Fungi have evolution, evolved a high degree of metabolic versatility that allows them to use a diverse range of organic substrates for growth, including simple compounds such as nitrate, ammonia, acetate, or ethanol. In some species the pigment melanin may play a role in extracting energy from ionizing radiation, such as gamma rays, gamma radiation. This form of radiotrophic fungus, "radiotrophic" growth has been described for only a few species, the effects on growth rates are small, and the underlying Biophysics, biophysical and biochemical processes are not well known. This process might bear similarity to carbon fixation, CO2 fixation via Visible spectrum, visible light, 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 Biological life cycle, life cycle of a species, the teleomorph (sexual reproduction) and the anamorph (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 occurs via vegetative spores (conidium, conidia) or through Fragmentation (reproduction), 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 clone (genetics), clonal populations adapted to a specific Ecological niche, niche, 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 (alternately 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 has been directly observed in all fungal phyla except Glomeromycota (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, ascus, asci and basidium, basidia, can be used in the identification of ascomycetes and basidiomycetes, respectively. Fungi employ two mating systems: heterothallic species allow mating only between individuals of the opposite mating type, 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 chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. Most eukaryotic chromosomes include packaging proteins called histones which, aided by ...
and a diploid stage in their life cycles. In sexually reproducing fungi, compatible individuals may combine by fusing their hyphae together into an interconnected network; this process, anastomosis, is required for the initiation of the sexual cycle. Many ascomycetes and basidiomycetes go through a dikaryotic stage, in which the nuclei inherited from the two parents do not combine immediately after cell fusion, but remain separate in the hyphal cells (see heterokaryosis). In ascomycetes, dikaryotic hyphae of the hymenium (the spore-bearing tissue layer) form a characteristic ''hook'' (crozier) at the hyphal septum. During cell division, 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, or fruiting body. Karyogamy in the asci is followed immediately by meiosis and the production of ascospores. After dispersal, the ascospores may germinate and form a new haploid mycelium. Sexual reproduction in basidiomycetes is similar to that of the ascomycetes. Compatible haploid hyphae fuse to produce a dikaryotic mycelium. However, the dikaryotic phase is more extensive in the basidiomycetes, often also present in the vegetatively growing mycelium. A specialized anatomical structure, called a clamp connection, is formed at each hyphal septum. As with the structurally similar hook in the ascomycetes, the clamp connection in the basidiomycetes is required for controlled transfer of nuclei during cell division, to maintain the dikaryotic stage with two genetically different nuclei in each hyphal compartment. A basidiocarp is formed in which club-like structures known as basidia generate haploid basidiospores after karyogamy and meiosis. The most commonly known basidiocarps are mushrooms, but they may also take other forms (see #Morphology, Morphology section). In fungi formerly classified as Zygomycota, haploid hyphae of two individuals fuse, forming a gametangium, a specialized cell structure that becomes a fertile gamete-producing cell. The gametangium develops into a zygospore, a thick-walled spore formed by the union of gametes. When the zygospore germinates, it undergoes meiosis, generating new haploid hyphae, which may then form asexual sporangiospores. These sporangiospores allow the fungus to rapidly disperse and germinate into new genetically identical haploid fungal mycelia.


Spore dispersal

The spores of most of the researched species of fungi are transported by wind. Such species often produce dry or Hydrophobe, hydrophobic 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 ascus, spore-bearing cells in some ascomycete species is such that the buildup of osmolyte, 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 10,000G-force, g; the net result is that the spore is ejected 0.01–0.02cm, sufficient distance for it to fall through the Agaricales, gills or polypore, pores into the air below. Other fungi, like the puffballs, 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 Nidulariaceae, 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.


Homothallism

In homothallism, homothallic sexual reproduction, two ploidy, haploid nuclei derived from the same individual fuse to form a zygote that can then undergo meiosis. Homothallic fungi include species with an ''Aspergillus''-like asexual stage (anamorphs) occurring in numerous different genera, several species of the Ascomycota, ascomycete genus ''Cochliobolus'', and the ascomycete ''Pneumocystis jiroveccii''. The earliest mode of sexual reproduction among eukaryotes was likely homothallism, that is, selfing, self-fertile unisexual reproduction.


Other sexual processes

Besides regular sexual reproduction with meiosis, certain fungi, such as those in the genera ''Penicillium'' and ''Aspergillus'', may exchange genetic material via parasexuality, 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 Evolutionary history of plants, plants and Evolutionary history of life, animals, the early fossil record of the fungi is meager. Factors that likely contribute to the under-representation of fungal species among fossils include the nature of fungal sporocarp (fungi), 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 taxon, extant fungi. Often recovered from a Permineralization, permineralized plant or animal host, these samples are typically studied by making thin-section preparations that can be examined with optical microscope, light microscopy or transmission electron microscopy. Researchers study compression fossils by dissolving the surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details. The earliest fossils possessing features typical of fungi date to the Paleoproterozoic era, some (annum, Ma); these multicellular benthic organisms had filamentous structures capable of anastomosis. 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 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 (biology), kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoos ...
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, Saphrotrophic nutrition, saprobism, and the development of Mutualism (biology), mutualistic relationships such as mycorrhiza and lichenization. Studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent lichenization events have occurred multiple times. In May 2019, scientists reported the discovery of a fossilized fungus, named ''Ourasphaira giraldae'', in the Northern Canada, Canadian Arctic, that may have grown on land a billion years ago, well before plants were living on land. Permineralization#Pyritization, Pyritized fungus-like microfossils 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 (542–488.3Ma), also long before land plants. Fossilized hyphae and spores recovered from the Ordovician of Wisconsin (460Ma) resemble modern-day Glomerales, and existed at a time when the land flora likely consisted of only non-vascular bryophyte-like plants. Prototaxites, which was probably a fungus or lichen, would have been the tallest organism of the late Silurian and early Devonian. Fungal fossils do not become common and uncontroversial until the early Devonian (416–359.2Ma), when they occur abundantly in the Rhynie chert, mostly as Zygomycota and Chytridiomycota. At about this same time, approximately 400Ma, the Ascomycota and Basidiomycota diverged, and all modern Class (biology), classes of fungi were present by the Late Carboniferous (Pennsylvanian (geology), 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 (fungi), sporocarp fossil, a ''Paleopyrenomycites'' species found in the Rhynie Chert. The oldest fossil with microscopic features resembling modern-day basidiomycetes is ''Palaeoancistrus'', found permineralized with a
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plant Vascular plants (from Latin ''vasculum'': duct), also known as Tracheophyta (the tracheophytes , from Greek τραχεῖα ἀρτηρία ''trācheia art ...

fern
from the Pennsylvanian. Rare in the fossil record are the Homobasidiomycetes (a taxon roughly equivalent to the mushroom-producing species of the Agaricomycetes). Two amber-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species ''Archaeomarasmius leggetti'') appeared during the late Cretaceous, 90Ma. Some time after the Permian–Triassic extinction event (251.4Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in sediments) formed, suggesting that fungi were the dominant life form at this time, representing nearly 100% of the available fossil record for this period. However, the relative proportion of fungal spores relative to spores formed by algae, algal 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 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 animals than to plants and are placed with the animals in the
monophyletic 300px, A cladogram of the primates, showing a ''monophyletic'' taxon: ''the simians'' (in yellow); a ''paraphyletic'' taxon: ''the prosimians'' (in cyan, including the red patch); and a ''polyphyletic'' group: ''the night-active primates, i.e., ...

monophyletic
group of opisthokonts. Analyses using
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram A diagram is a symb ...
support a monophyletic group, monophyletic origin of fungi. The Taxonomy (biology), taxonomy 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 matings. 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 Botanical nomenclature, nomenclature. Until relatively recent (2012) changes to the International Code of Nomenclature for algae, fungi and plants, 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 and MycoBank are officially recognized Nomenclature codes, nomenclatural repositories and list current names of fungal species (with cross-references to older synonym (taxonomy), synonyms). The 2007 classification of Kingdom Fungi is the result of a large-scale collaborative research effort involving dozens of mycologists and other scientists working on fungal taxonomy. It recognizes seven phyla, two of which—the Ascomycota and the Basidiomycota—are contained within a branch representing subkingdom Dikarya, the most species rich and familiar group, including all the mushrooms, most food-spoilage molds, most plant pathogenic fungi, and the beer, wine, and bread yeasts. The accompanying cladogram depicts the major fungal Taxon, taxa 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 distances.


Taxonomic groups

The major phyla (sometimes called divisions) of fungi have been classified mainly on the basis of characteristics of their sexual reproduction, reproductive structures. , nine major Lineage (evolution), lineages have been identified: Opisthosporidia, Chytridiomycota, Neocallimastigomycota, Blastocladiomycota, Zoopagomycota, Mucoromycota, Glomeromycota, Ascomycota and Basidiomycota. Phylogenetic analysis has demonstrated that the Microsporidia, unicellular parasites of animals and protists, are fairly recent and highly derived endobiotic fungi (living within the tissue of another species). Previously considered to be "primitive" protozoa, they are now thought to be either a Basal (phylogenetics), basal branch of the Fungi, or a sister group–each other's closest evolutionary relative. The Chytridiomycota are commonly known as chytrids. These fungi are distributed worldwide. Chytrids and their close relatives Neocallimastigomycota and Blastocladiomycota (below) are the only fungi with active motility, producing zoospores that are capable of active movement through aqueous phases with a single flagellum, leading early taxonomists to classify them as protists. Molecular phylogenetics, Molecular phylogenies, inferred from rRNA sequences in ribosomes, suggest that the Chytrids are a Basal (phylogenetics), basal group divergent from the other fungal phyla, consisting of four major clades with suggestive evidence for paraphyly or possibly polyphyly. The Blastocladiomycota were previously considered a taxonomic clade within the Chytridiomycota. Molecular data and ultrastructure, ultrastructural characteristics, however, place the Blastocladiomycota as a sister clade to the Zygomycota, Glomeromycota, and Dikarya (Ascomycota and Basidiomycota). The blastocladiomycetes are Saprotrophic nutrition, saprotrophs, feeding on decomposing organic matter, and they are parasites of all eukaryotic groups. Unlike their close relatives, the chytrids, most of which exhibit Biological life cycle#Haplontic life cycle, zygotic meiosis, the blastocladiomycetes undergo Biological life cycle#Haplodiplontic life cycle, sporic meiosis. The Neocallimastigomycota were earlier placed in the phylum Chytridomycota. Members of this small phylum are anaerobic organisms, living in the digestive system of larger herbivorous mammals and in other terrestrial and aquatic environments enriched in cellulose (e.g., domestic waste landfill sites). They lack
mitochondria A mitochondrion (; ) is a double-membrane Image:Schematic size.jpg, up150px, Schematic of size-based membrane exclusion A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, i ...

mitochondria
but contain hydrogenosomes of mitochondrial origin. As in the related chrytrids, neocallimastigomycetes form zoospores that are posteriorly uniflagellate or polyflagellate. Members of the Glomeromycota form arbuscular mycorrhizae, a form of mutualist symbiosis wherein fungal hyphae invade plant root cells and both species benefit from the resulting increased supply of nutrients. All known Glomeromycota species reproduce asexually. The symbiotic association between the Glomeromycota and plants is ancient, with evidence dating to 400 million years ago. Formerly part of the Zygomycota (commonly known as 'sugar' and 'pin' molds), the Glomeromycota were elevated to phylum status in 2001 and now replace the older phylum Zygomycota. Fungi that were placed in the Zygomycota are now being reassigned to the Glomeromycota, or the subphyla incertae sedis Mucoromycotina, Kickxellomycotina, the Zoopagomycotina and the Entomophthoromycotina. Some well-known examples of fungi formerly in the Zygomycota include black bread mold (''Rhizopus stolonifer''), and ''Pilobolus'' species, capable of ejecting
spore )'', growing on a thinning, thinned hybrid black poplar ''(populus, Populus x canadensis)''. The last stage of the moss#Life cycle, moss lifecycle is shown, where the sporophytes are visible before dispersion of their spores: the calyptra (1) is ...
s several meters through the air. Medically relevant genera include ''Mucor'', ''Rhizomucor'', and ''Rhizopus''. The Ascomycota, commonly known as sac fungi or ascomycetes, constitute the largest taxonomic group within the Eumycota. These fungi form meiotic spores called ascospores, which are enclosed in a special sac-like structure called an ascus. This phylum includes morels, a few
mushroom A mushroom or toadstool is the fleshy, spore )'', growing on a thinning, thinned hybrid black poplar ''(populus, Populus x canadensis)''. The last stage of the moss#Life cycle, moss lifecycle is shown, where the sporophytes are visible befor ...

mushroom
s and
truffles A truffle is the fruiting body of a subterranean ascomycete fungus, predominantly one of the many species of the genus ''Tuber (fungus), Tuber''. In addition to ''Tuber'', many other genera of fungi are classified as truffles including ''Geopor ...
, unicellular
yeast Yeasts are eukaryotic Eukaryotes () are organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classifie ...

yeast
s (e.g., of the genera ''Saccharomyces'', ''Kluyveromyces'', ''Pichia'', and ''Candida (genus), Candida''), and many filamentous fungi living as saprotrophs, parasites, and mutualistic symbionts (e.g. lichens). Prominent and important genera of filamentous ascomycetes include ''Aspergillus'', ''Penicillium'', ''Fusarium'', and ''Claviceps''. Many ascomycete species have only been observed undergoing asexual reproduction (called anamorphic species), but analysis of molecular data has often been able to identify their closest teleomorphs in the Ascomycota. Because the products of meiosis are retained within the sac-like ascus, ascomycetes have been used for elucidating principles of genetics and heredity (e.g., ''Neurospora crassa''). Members of the Basidiomycota, commonly known as the club fungi or basidiomycetes, produce meiospores called basidiospores on club-like stalks called basidium, basidia. Most common mushrooms belong to this group, as well as rust (fungus), rust and smut (fungus), smut fungi, which are major pathogens of grains. Other important basidiomycetes include the maize pathogen ''Ustilago maydis'', human commensalism, commensal species of the genus ''Malassezia'', and the opportunistic infection, opportunistic human pathogen, ''Cryptococcus neoformans''.


Fungus-like organisms

Because of similarities in morphology and lifestyle, the slime molds (mycetozoans, plasmodiophorids, acrasids, ''Fonticula'' and labyrinthulids, now in Amoebozoa, Rhizaria, Excavata, Opisthokonta and Stramenopiles, respectively), water molds (
oomycete Oomycota or oomycetes () form a distinct phylogenetic In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, mole ...
s) and hyphochytrids (both Stramenopiles) were formerly classified in the kingdom Fungi, in groups like Mastigomycotina, Gymnomycota and Phycomycetes. The slime molds were studied also as protozoans, leading to an ambiregnal, duplicated taxonomy. Unlike true fungi, the
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane cell membrane vs. Prokaryotes The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to a ...
s of oomycetes contain cellulose and lack
chitin units that repeat to form long chains in β-(1→4)-linkage. of the chitin molecule. Chitin ( carbon, C8H13O5N)n ( ) is a long-chain polymer A polymer (; Greek ''wikt:poly-, poly-'', "many" + ''wikt:-mer, -mer'', "part") is a Chemical su ...

chitin
. Hyphochytrids have both chitin and cellulose. Slime molds lack a cell wall during the assimilative phase (except labyrinthulids, which have a wall of scales), and ingest nutrients by ingestion (phagocytosis, except labyrinthulids) rather than absorption (osmotrophy, as fungi, labyrinthulids, oomycetes and hyphochytrids). Neither water molds nor slime molds are closely related to the true fungi, and, therefore, taxonomists no longer group them in the kingdom Fungi. Nonetheless, studies of the oomycetes and myxomycetes are still often included in
mycology Mycology is the branch of biology concerned with the study of fungus, fungi, including their genetics, genetic and biochemistry, biochemical properties, their Taxonomy (biology), taxonomy and ethnomycology, their use to humans as a source for ti ...
textbooks and primary research literature. The Eccrinales and Amoebidiales are opisthokont protists, previously thought to be zygomycete fungi. Other groups now in Opisthokonta (e.g., ''Corallochytrium'', Ichthyosporea) were also at given time classified as fungi. The genus ''Blastocystis'', now in Stramenopiles, was originally classified as a yeast. ''Ellobiopsis'', now in Alveolata, was considered a chytrid. The bacteria were also included in fungi in some classifications, as the group Schizomycetes. The Rozellida clade, including the "ex-chytrid" ''Rozella'', is a genetically disparate group known mostly from environmental DNA sequences that is a sister group to fungi. Members of the group that have been isolated lack the chitinous cell wall that is characteristic of fungi. Alternatively, Rozella can be classified as a basal fungal group. The nucleariids may be the next sister group to the eumycete clade, and as such could be included in an expanded fungal kingdom. Many Actinomycetales (Actinobacteria), a group with many filamentous bacteria, were also long believed to be fungi.


Ecology

Although often inconspicuous, fungi occur in every environment on Earth and play very important roles in most ecosystems. Along with bacteria, fungi are the major decomposers in most terrestrial (and some aquatic) ecosystems, and therefore play a critical role in biogeochemical cycles and in many food webs. As decomposers, they play an essential role in nutrient cycling, especially as saprotrophs and symbionts, degrading organic matter to inorganic molecules, which can then re-enter anabolic metabolic pathways in plants or other organisms.


Symbiosis

Many fungi have important symbiotic relationships with organisms from most if not all Kingdom (biology), kingdoms. These interactions can be Mutualism (biology), mutualistic or antagonistic in nature, or in the case of commensal fungi are of no apparent benefit or detriment to the host.


With plants

Mycorrhizal symbiosis between plants and fungi is one of the most well-known plant–fungus associations and is of significant importance for plant growth and persistence in many ecosystems; over 90% of all plant species engage in mycorrhizal relationships with fungi and are dependent upon this relationship for survival. The mycorrhizal symbiosis is ancient, dating back to at least 400 million years. It often increases the plant's uptake of inorganic compounds, such as nitrate and phosphate from soils having low concentrations of these key plant nutrients. The fungal partners may also mediate plant-to-plant transfer of carbohydrates and other nutrients. Such mycorrhizal communities are called "common mycorrhizal networks". A special case of mycorrhiza is myco-heterotrophy, whereby the plant parasitizes the fungus, obtaining all of its nutrients from its fungal symbiont. Some fungal species inhabit the tissues inside roots, stems, and leaves, in which case they are called endophytes. Similar to mycorrhiza, endophytic colonization by fungi may benefit both symbionts; for example, endophytes of grasses impart to their host increased resistance to herbivores and other environmental stresses and receive food and shelter from the plant in return.


With algae and cyanobacteria

Lichens are a symbiotic relationship between fungi and photosynthetic algae or cyanobacteria. The photosynthetic partner in the relationship is referred to in lichen terminology as a "photobiont". The fungal part of the relationship is composed mostly of various species of ascomycetes and a few basidiomycetes. Lichens occur in every ecosystem on all continents, play a key role in soil formation and the initiation of Ecological succession, biological succession, and are prominent in some extreme environments, including polar region, polar, Alpine climate, alpine, and semiarid climate, semiarid desert regions. They are able to grow on inhospitable surfaces, including bare soil, rocks, Bark (botany), tree bark, wood, shells, barnacles and leaves. As in mycorrhizas, the photobiont provides sugars and other carbohydrates via photosynthesis to the fungus, while the fungus provides minerals and water to the photobiont. The functions of both symbiotic organisms are so closely intertwined that they function almost as a single organism; in most cases the resulting organism differs greatly from the individual components. Lichenization is a common mode of nutrition for fungi; around 27% of known fungi—more than 19,400 species—are lichenized. Characteristics common to most lichens include obtaining organic carbon by photosynthesis, slow growth, small size, long life, long-lasting (seasonal) Vegetative reproduction, vegetative reproductive structures, mineral nutrition obtained largely from airborne sources, and greater tolerance of desiccation than most other photosynthetic organisms in the same habitat.


With insects

Many insects also engage in Ant-fungus mutualism, mutualistic relationships with fungi. Several groups of ants cultivate fungi in the order Chaetothyriales for several purposes: as a food source, as a structural component of their nests, and as a part of an ant/plant symbiosis in the domatium, domatia (tiny chambers in plants that house arthropods). Ambrosia beetles cultivate various species of fungi in the bark of trees that they infest. Likewise, females of several wood wasp species (genus ''Sirex'') inject their eggs together with spores of the wood-rotting fungus ''Amylostereum areolatum'' into the Wood#Heartwood and sapwood, sapwood of pine trees; the growth of the fungus provides ideal nutritional conditions for the development of the wasp larvae. At least one species of stingless bee has a relationship with a fungus in the genus ''Monascus'', where the larvae consume and depend on fungus transferred from old to new nests. Termites on the African savannah are also known to cultivate fungi, and yeasts of the genera ''Candida (genus), Candida'' and ''Lachancea'' inhabit the Gastrointestinal tract, gut of a wide range of insects, including neuropterans, beetles, and cockroaches; it is not known whether these fungi benefit their hosts. Fungi growing in Coarse woody debris, dead wood are essential for Xylophagy, xylophagous insects (e.g. woodboring beetles). They deliver nutrients needed by Xylophagy, xylophages to nutritionally scarce dead wood. Thanks to this nutritional enrichment the larvae of the woodboring insect is able to grow and develop to adulthood. The larvae of many families of fungicolous flies, particularly those within the superfamily Sciaroidea such as the Mycetophilidae and some Keroplatidae feed on fungal fruiting bodies and sterile mycorrhizae.


As pathogens and parasites

Many fungi are parasites on plants, animals (including humans), and other fungi. Serious pathogens of many cultivated plants causing extensive damage and losses to agriculture and forestry include the rice blast fungus ''Magnaporthe oryzae'', tree pathogens such as ''Ophiostoma ulmi'' and ''Ophiostoma novo-ulmi'' causing Dutch elm disease and ''Cryphonectria parasitica'' responsible for chestnut blight, and plant pathogens in the genera ''Fusarium'', ''Ustilago'', ''Alternaria'', and ''Cochliobolus''. Some carnivorous fungi, like ''Paecilomyces lilacinus'', are Nematophagous fungus, predators of nematodes, which they capture using an array of specialized structures such as constricting rings or adhesive nets. Many fungi that are plant pathogens, such as ''Magnaporthe oryzae'', can switch from being biotrophic (parasitic on living plants) to being necrotrophic (feeding on the dead tissues of plants they have killed). This same principle is applied to fungi-feeding parasites, including ''Asterotremella albida'', which feeds on the fruit bodies of other fungi both while they are living and after they are dead. Some fungi can cause serious diseases in humans, several of which may be fatal if untreated. These include aspergillosis, candidiasis, coccidioidomycosis, cryptococcosis, histoplasmosis, Eumycetoma, mycetomas, and paracoccidioidomycosis. Furthermore, persons with Immunodeficiency, immuno-deficiencies are particularly susceptible to disease by genera such as ''Aspergillus'', ''Candida (genus), Candida'', ''Cryptococcus neoformans, Cryptoccocus'', ''Histoplasma'', and ''Pneumocystis''. Other fungi can attack eyes, nails, hair, and especially skin, the so-called Dermatophyte, dermatophytic and keratinophilic fungi, and cause local infections such as ringworm and athlete's foot. Fungal spores are also a cause of allergies, and fungi from different taxonomic groups can evoke allergic reactions.


As targets of mycoparasites

Organisms that parasitize fungi are known as mycoparasitism, mycoparasitic organisms. About 300 species of fungi and fungus-like organisms, belonging to 13 classes and 113 genera, are used as biocontrol agents against plant fungal diseases. Fungi can also act as mycoparasites or antagonists of other fungi, such as ''Hypomyces chrysospermus'', which grows on bolete mushrooms. Fungi can also become the target of infection by mycoviruses.


Mycotoxins

Many fungi produce biological activity, biologically active compounds, several of which are toxin, toxic to animals or plants and are therefore called mycotoxins. Of particular relevance to humans are mycotoxins produced by molds causing food spoilage, and poisonous mushrooms (see above). Particularly infamous are the lethal amatoxins in some ''Amanita'' mushrooms, and Ergotamine, ergot alkaloids, which have a long history of causing serious epidemics of ergotism (St Anthony's Fire) in people consuming rye or related cereals contaminated with sclerotia of the ergot fungus, ''Claviceps purpurea''. Other notable mycotoxins include the aflatoxins, which are insidious Hepatotoxicity, liver toxins and highly carcinogenic metabolites produced by certain ''Aspergillus'' species often growing in or on grains and nuts consumed by humans, ochratoxins, patulin, and trichothecenes (e.g., T-2 mycotoxin) and fumonisins, which have significant impact on human food supplies or animal livestock. Mycotoxins are secondary metabolites (or natural products), and research has established the existence of biochemical pathways solely for the purpose of producing mycotoxins and other natural products in fungi. Mycotoxins may provide Fitness (biology), fitness benefits in terms of physiological adaptation, competition with other microbes and fungi, and protection from consumption (fungivore, fungivory). Many fungal secondary metabolites (or derivatives) are used medically, as described under Human Use below.


Pathogenic mechanisms

''Ustilago maydis'' is a pathogenic plant fungus that causes smut disease in maize and teosinte. Plants have evolved efficient defense systems against pathogenic microbes such as ''U. maydis''. A rapid defense reaction after pathogen attack is the oxidative burst where the plant produces reactive oxygen species at the site of the attempted invasion. ''U. maydis'' can respond to the oxidative burst with an oxidative stress response, regulated by the gene ''YAP1''. The response protects ''U. maydis'' from the host defense, and is necessary for the pathogen's virulence. Furthermore, ''U. maydis'' has a well-established recombinational DNA repair system which acts during mitosis and meiosis. The system may assist the pathogen in surviving DNA damage arising from the host plant's oxidative defensive response to infection. ''Cryptococcus neoformans'' is an encapsulated yeast that can live in both plants and animals. ''C.neoformans'' usually infects the lungs, where it is phagocytosed by alveolar macrophages. Some ''C.neoformans'' can survive intracellular, inside macrophages, which appears to be the basis for latency period, latency, disseminated disease, and resistance to antifungal agents. One mechanism by which ''C.neoformans'' survives the hostile macrophage environment is by up-regulating the expression of genes involved in the oxidative stress response. Another mechanism involves meiosis. The majority of ''C.neoformans'' are mating "type a". Filaments of mating "type a" ordinarily have haploid nuclei, but they can become diploid (perhaps by endoduplication or by stimulated nuclear fusion) to form blastospores. The diploid nuclei of blastospores can undergo meiosis, including recombination, to form haploid basidiospores that can be dispersed. This process is referred to as monokaryotic fruiting. This process requires a gene called ''DMC1'', which is a conserved homologue of genes ''recA'' in bacteria and ''RAD51'' in eukaryotes, that mediates homologous chromosome pairing during meiosis and repair of DNA double-strand breaks. Thus, ''C.neoformans'' can undergo a meiosis, monokaryotic fruiting, that promotes recombinational repair in the oxidative, DNA damaging environment of the host macrophage, and the repair capability may contribute to its virulence.


Human use

The human use of fungi for food preparation or preservation and other purposes is extensive and has a long history. Mushroom farming and mushroom gathering are large industries in many countries. The study of the historical uses and sociological impact of fungi is known as ethnomycology. Because of the capacity of this group to produce an enormous range of natural products with antimicrobial or other biological activities, many species have long been used or are being developed for industrial production of antibiotics, vitamins, and Taxol#Production, anti-cancer and Lovastatin, cholesterol-lowering drugs. Methods have been developed for genetic engineering of fungi, enabling metabolic engineering of fungal species. For example, genetic modification of yeast species—which are easy to grow at fast rates in large fermentation vessels—has opened up ways of pharmaceutical production that are potentially more efficient than production by the original source organisms.


Therapeutic uses


Modern chemotherapeutics

Many species produce metabolites that are major sources of pharmacology, pharmacologically active drugs. Particularly important are the antibiotics, including the penicillins, a structurally related group of β-lactam antibiotics that are synthesized from small peptides. Although naturally occurring penicillins such as penicillin G (produced by ''Penicillium chrysogenum'') have a relatively narrow spectrum of biological activity, a wide range of other penicillins can be produced by Chemical synthesis, chemical modification of the natural penicillins. Modern penicillins are semisynthesis, semisynthetic compounds, obtained initially from fermentation (biochemistry), fermentation cultures, but then structurally altered for specific desirable properties. Other antibiotics produced by fungi include: ciclosporin, commonly used as an immunosuppressant during organ transplant, transplant surgery; and fusidic acid, used to help control infection from Methicillin-resistant Staphylococcus aureus, methicillin-resistant ''Staphylococcus aureus'' bacteria. Widespread use of antibiotics for the treatment of bacterial diseases, such as tuberculosis, syphilis, leprosy, and others began in the early 20th century and continues to date. In nature, antibiotics of fungal or bacterial origin appear to play a dual role: at high concentrations they act as chemical defense against competition with other microorganisms in species-rich environments, such as the rhizosphere (ecology), rhizosphere, and at low concentrations as quorum sensing, quorum-sensing molecules for intra- or interspecies signaling. Other drugs produced by fungi include griseofulvin isolated from ''Penicillium griseofulvum'', used to treat fungal infections, and statins (HMG-CoA reductase inhibitors), used to inhibit cholesterol synthesis. Examples of statins found in fungi include mevastatin from ''Penicillium citrinum'' and lovastatin from ''Aspergillus terreus'' and the Pleurotus ostreatus, oyster mushroom. Fungi produce compounds that inhibit viruses and cancer cells. Specific metabolites, such as polysaccharide-K, ergotamine, and Beta-lactam antibiotic, β-lactam antibiotics, are routinely used in clinical medicine. The shiitake mushroom is a source of lentinan, a clinical drug approved for use in cancer treatments in several countries, including Japan. In Europe and Japan, polysaccharide-K (brand name Krestin), a chemical derived from ''Trametes versicolor'', is an approved adjuvant for cancer therapy.


Traditional medicine

Certain mushrooms are used as supposed therapeutics in traditional medicine, folk medicine practices, such as traditional Chinese medicine. Mushrooms with a history of such use include ''Agaricus subrufescens'', ''Ganoderma lucidum'', and ''Ophiocordyceps sinensis''.


Cultured foods

Baker's yeast or ''Saccharomyces cerevisiae'', a unicellular fungus, is used to make bread and other wheat-based products, such as pizza dough and dumplings. Yeast species of the genus ''Saccharomyces'' 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 American English (AmE, AE, AmEng, USEng, en-US), sometimes called United States English or U.S. English, is the set of varieties of the English language native to the United States. C ...

soy sauce
) 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, morels, chanterelles, truffles, 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 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 may be used to eliminate or suppress the growth of harmful plant pathogens, such as insects, mites, weeds, nematodes, and other fungi that cause diseases of important crop plants. This has generated strong interest in practical applications that use these fungi in the biological control of these agricultural pests. Entomopathogenic fungi can be used as biopesticides, as they actively kill insects. Examples that have been used as biological insecticides are ''Beauveria bassiana'', ''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, 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'' and ''Schizosaccharomyces pombe'', each of which with a long history of use to investigate issues in eukaryotic cell biology and genetics, such as cell cycle regulation, chromatin structure, and gene regulation. Other fungal models have emerged that address specific biological questions relevant to medicine, plant pathology, and industrial uses; examples include ''Candida albicans'', a dimorphic, opportunistic human pathogen, ''Magnaporthe grisea'', a plant 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 * Marine fungi * Mycosis * Outline of fungi


References

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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 and 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


External links

{{commons category {{Wiktionary {{Library resources box , onlinebooks=yes , by=no , lcheading= Fungi , label=Fungus
Tree of Life web project: Fungi
* Mushroom Observer
mushroomobserver.org
, a collaborative fungus recording and identification project

i

{{Fungi classification {{Fungus structure {{Eukaryota {{Nature nav {{Life on Earth {{Organisms et al. {{Taxonbar, from=Q764 {{Authority control {{Featured article Articles containing video clips Cryptogams Extant Early Devonian first appearances Fungi, Kingdoms (biology), Fungi