Array ( [0] => {{Short description|Biological kingdom, separate from plants and animals}} [1] => {{Redirect|Fungi|other uses|Fungi (disambiguation)|and|Fungus (disambiguation)}} [2] => {{Featured article}} [3] => {{pp-semi|small=yes}} [4] => {{pp-move-indef}} [5] => {{Use dmy dates|date=August 2023}} [6] => {{cs1 config |name-list-style=vanc |display-authors=6}} [7] => {{Automatic taxobox [8] => | name=Fungi [9] => | fossil_range=Middle [[Ordovician]] – [[Holocene|Present]] (but [[#Evolution|see text]]) {{Fossilrange|460|0|earliest=Ediacaran}} [10] => | image=Fungi_collage.jpg [11] => | image_upright=1.3 [12] => | image_caption=Clockwise from top left: {{plainlist| [13] => * ''[[Amanita muscaria]]'', a basidiomycete; [14] => * ''[[Sarcoscypha coccinea]]'', an ascomycete; [15] => * bread covered in [[Mold (fungus)|mold]]; [16] => * a [[Chytridiomycota|chytrid]]; [17] => * an ''[[Aspergillus]]'' [[conidiophore]]. [18] => }} [19] => | image_alt=A collage of five fungi (clockwise from top left): a mushroom with a flat red top with white spots and a white stem growing on the ground; a red cup-shaped fungus growing on wood; a stack of green and white moldy bread slices on a plate; a microscopic spherical grey semitransparent cell with a smaller spherical cell beside it; a microscopic view of an elongated cellular structure shaped like a microphone, attached to the larger end is a number of smaller roughly circular elements that collectively form a mass around it [20] => | display_parents=3 [21] => | taxon=Fungi [22] => | authority=([[Carl Linnaeus|L.]]) [[Royall T. Moore|R.T.Moore]] [23] => | subdivision_ranks=Subkingdoms/Phyla [24] => | subdivision = [25] => * [[Rozellomyceta]] [26] => ** [[Rozellomycota]] [27] => ** [[Microsporidia]] [28] => * [[Aphelidiomyceta]] [29] => ** [[Aphelidiomycota]] [30] => * [[Amastigomycota|Eumycota]] [31] => ** [[Chytridiomyceta]] [32] => *** [[Neocallimastigomycota]] [33] => *** [[Chytridiomycota]] [34] => ** [[Blastocladiomyceta]] [35] => *** [[Blastocladiomycota]] [36] => ** [[Zoopagomyceta]] [37] => *** [[Basidiobolomycota]] [38] => *** [[Entomophthoromycota]] [39] => *** [[Kickxellomycota]] [40] => ** [[Mortierellomycota]] [41] => ** [[Mucoromyceta]] [42] => *** [[Calcarisporiellomycota]] [43] => *** [[Mucoromycota]] [44] => ** [[Symbiomycota]] [45] => *** [[Glomeromycota]] [46] => *** [[Entorrhizomycota]] [47] => *** [[Dikarya]] [48] => **** [[Basidiomycota]] [49] => **** [[Ascomycota]] [50] => }} [51] => [52] => A '''fungus''' ({{plural form}}: '''fungi'''{{IPAc-en|audio=En-us-fungi.ogg|ˈ|f|ʌ|n|dʒ|aɪ}}, {{IPAc-en|audio=En-us-fungi-2.ogg|ˈ|f|ʌ|ŋ|ɡ|aɪ}}, {{IPAc-en|audio=En-us-fungi-3.ogg|ˈ|f|ʌ|ŋ|ɡ|i}} or {{IPAc-en|audio=En-us-fungi-4.ogg|ˈ|f|ʌ|n|dʒ|i}}. The first two pronunciations are favored more in the US and the others in the UK, however all pronunciations can be heard in any English-speaking country. or '''funguses''') is any member of the group of [[Eukaryote|eukaryotic]] organisms that includes microorganisms such as [[yeast]]s and [[Mold (fungus)|mold]]s, as well as the more familiar [[mushroom]]s. These organisms are classified as one of the [[Kingdom_(biology)#Six kingdoms (1998)|traditional eukaryotic kingdoms]], along with [[Animal]]ia, [[Plant]]ae and either [[Protist]]a{{cite journal |last=Whittaker |first=R.H. |date=January 1969 |title=New concepts of kingdoms or organisms. Evolutionary relations are better represented by new classifications than by the traditional two kingdoms |journal=[[Science (journal)|Science]] |volume=163 |issue=3863 |pages=150–60 |pmid=5762760 |doi=10.1126/science.163.3863.150|bibcode = 1969Sci...163..150W |citeseerx=10.1.1.403.5430 }} or [[Protozoa]] and [[Chromista]].{{cite journal |last=Cavalier-Smith |first=T. |year=1998 |title=A revised six-kingdom system of life |journal=[[Biological Reviews]] |volume=73 |pages=203–66 |url=http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=685 |issue=3 |doi=10.1111/j.1469-185X.1998.tb00030.x |pmid=9809012|s2cid=6557779 }} [53] => [54] => A characteristic that places fungi in a different kingdom from [[plants]], [[bacteria]], and some [[protists]] is [[chitin]] in their [[cell wall]]s. Fungi, like animals, are [[heterotroph]]s; they acquire their food by absorbing dissolved molecules, typically by secreting [[digestive enzyme]]s into their environment. Fungi do not [[photosynthesize]]. Growth is their means of [[motility|mobility]], except for [[spore]]s (a few of which are [[flagellate]]d), which may travel through the air or water. Fungi are the principal [[decomposer]]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]] (i.e. they form a ''[[monophyletic]] group''), an interpretation that is also strongly supported by [[molecular phylogenetics]]. This fungal group is distinct from the structurally similar [[Mycetozoa|myxomycetes]] (slime molds) and [[oomycete]]s (water molds). The discipline of [[biology]] devoted to the study of fungi is known as [[mycology]] (from the [[Greek language|Greek]] {{Lang|grc|μύκης|italic=no}} ''{{lang|grc-Latn|mykes}}'', mushroom). In the past, mycology was regarded as a branch of [[botany]], although it is now known that fungi are genetically more closely related to animals than to plants. [55] => [56] => Abundant worldwide, most fungi are inconspicuous because of the small size of their structures, and their [[Crypsis|cryptic]] lifestyles in soil or on dead matter. Fungi include [[Symbiosis|symbionts]] of plants, animals, or other fungi and also [[parasites]]. They may become noticeable when [[Sporocarp (fungi)|fruiting]], either as mushrooms or as molds. Fungi perform an essential role in the decomposition of organic matter and have fundamental roles in nutrient [[Biogeochemical cycle|cycling]] and exchange in the environment. They have long been used as a direct [[Fungivore|source of human food]], in the form of mushrooms and [[truffles]]; as a [[leavening agent]] for bread; and in the [[Fermentation (food)|fermentation]] of various food products, such as [[Fermentation in winemaking|wine]], [[Beer fermentation|beer]], and [[soy sauce]]. Since the 1940s, fungi have been used for the production of [[antibiotic]]s, and, more recently, various [[enzyme]]s produced by fungi are used [[Enzyme#Industrial applications|industrially]] and in [[protease#Biodiversity of proteases|detergents]]. Fungi are also used as [[biological pesticide]]s to control weeds, plant diseases, and insect pests. Many species produce [[bioactive compound]]s called [[mycotoxin]]s, such as [[alkaloid]]s and [[polyketide]]s, that are toxic to animals, including humans. The fruiting structures of [[Psychoactive mushrooms|a few species]] contain [[psychotropic]] compounds and are consumed [[Recreational drug use|recreationally]] or in traditional [[Entheogen|spiritual ceremonies]]. Fungi can break down manufactured materials and buildings, and become significant [[Pathogenic fungi|pathogens]] of humans and other animals. Losses of crops due to fungal diseases (e.g., [[rice blast disease]]) or [[food spoilage]] can have a large impact on human [[Food security|food supplies]] and local economies. [57] => [58] => The fungus kingdom encompasses an enormous diversity of [[taxon|taxa]] with varied ecologies, [[biological life cycle|life cycle]] strategies, and [[morphology (biology)|morphologies]] ranging from unicellular aquatic [[chytrid]]s to large mushrooms. However, little is known of the true [[biodiversity]] of the fungus kingdom, which has been estimated at 2.2 million to 3.8 million species. Of these, only about 148,000 have been described, with over 8,000 species known to be detrimental to plants and at least 300 that can be pathogenic to humans.{{cite journal |title=Stop neglecting fungi|date=25 July 2017|journal=Nature Microbiology |volume=2 |issue=8 |pages=17120 |doi=10.1038/nmicrobiol.2017.120 |doi-access=free | title-link=doi |pmid=28741610}} Ever since the pioneering 18th and 19th century [[Taxonomy (biology)|taxonomical]] works of [[Carl Linnaeus]], [[Christiaan Hendrik Persoon]], and [[Elias Magnus Fries]], fungi have been classified according to their morphology (e.g., characteristics such as spore color or microscopic features) or [[physiology]]. Advances in [[molecular genetics]] have opened the way for [[DNA sequencing|DNA analysis]] to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits. [[Phylogenetic]] studies published in the first decade of the 21st century have helped reshape the classification within the fungi kingdom, which is divided into one [[Kingdom (biology)#Definition and associated terms|subkingdom]], seven [[phylum|phyla]], and ten [[subphyla]]. [59] => {{toclimit}} [60] => [61] => == Etymology == [62] => The English word ''fungus'' is directly adopted from the [[Latin]] ''fungus'' (mushroom), used in the writings of [[Horace]] and [[Pliny the Elder|Pliny]]. This in turn is derived from the [[Ancient Greek|Greek]] word ''sphongos'' (σφόγγος 'sponge'), which refers to the [[macroscopic]] structures and morphology of mushrooms and molds;{{sfn|Ainsworth|1976|p=2}} the [[Root (linguistics)|root]] is also used in other languages, such as the German ''[[wikt:Schwamm#German|Schwamm]]'' ('sponge') and ''[[wikt:Schimmel#German|Schimmel]]'' ('mold'). [63] => [64] => The word ''[[mycology]]'' is derived from the Greek {{transl|grc|mykes}} (μύκης 'mushroom') and ''logos'' (λόγος 'discourse').{{sfn|Alexopoulos|Mims|Blackwell|1996|p=1}} 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]].{{cite book |last1=Persoon |first1=Christiaan Hendrik |title=Observationes Mycologicae: Part 1 |date=1796 |publisher=Peter Philipp Wolf |location=Leipzig, (Germany) |language=la |url=http://bibdigital.rjb.csic.es/ing/Libro.php?Libro=5680 |access-date=30 March 2019 |archive-url=https://web.archive.org/web/20131219091343/http://bibdigital.rjb.csic.es/ing/Libro.php?Libro=5680 |archive-date=19 December 2013 |url-status=live }} The word appeared in English as early as 1824 in a book by [[Robert Kaye Greville]].{{cite book |last1=Greville |first1=Robert Kaye |title=Scottish Cryptogamie Flora: Or Coloured Figures and Descriptions of Cryptogamic Plants, Belonging Chiefly to the Order Fungi |date=1824 |publisher=Maclachland and Stewart |location=Edinburgh, Scotland |volume=2 |page=65 |url=https://babel.hathitrust.org/cgi/pt?id=nyp.33433008943957;view=1up;seq=45}} From p. 65: "This little plant will probably not prove rare in Great Britain, when mycology shall be more studied." In 1836 the English naturalist [[Miles Joseph Berkeley]]'s publication ''The English Flora of Sir James Edward Smith, Vol. 5.'' also refers to mycology as the study of fungi.{{sfn|Ainsworth|1976|p=2}}{{cite book |last1=Smith |first1=James Edward |editor1-last=Hooker |editor1-first=William Jackson |editor2-last=Berkeley |editor2-first=Miles Joseph |title=The English Flora of Sir James Edward Smith |date=1836 |publisher=Longman, Rees, Orme, Brown, Green & Longman |location=London, England |series=Vol. 5, part II: "Class XXIV. Cryptogamia" |page=7 |url=https://babel.hathitrust.org/cgi/pt?id=msu.31293010136830;view=1up;seq=403}} From p. 7: "This has arisen, I conceive, partly from the practical difficulty of preserving specimens for the herbarium, partly from the absence of any general work, adapted to the immense advances which have of late years been made in the study of Mycology." [65] => [66] => A group of all the fungi present in a particular region is known as ''[[mycobiota]]'' (plural noun, no singular).{{cite web |url=http://glossary.lias.net/wiki/Mycobiota |title=LIAS Glossary |access-date=14 August 2013 |archive-url=https://web.archive.org/web/20131211113036/http://glossary.lias.net/wiki/Mycobiota |archive-date=11 December 2013 |url-status=live }} 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 morphologically similar to [[fauna]] and [[flora]]. The [[Species Survival Commission]] (SSC) of the [[International Union for Conservation of Nature]] (IUCN) in August 2021 asked that the phrase ''fauna and flora'' be replaced by ''fauna, flora, and funga''.{{cite web |url=https://www.iucn.org/sites/dev/files/statement-3f.pdf |title=IUCN SSC acceptance of Fauna Flora Funga |publisher=Fungal Conservation Committee, [[IUCN]] SSC |date=2021 |quote=The IUCN Species Survival Commission calls for the due recognition of fungi as major components of biodiversity in legislation and policy. It fully endorses the Fauna Flora Funga Initiative and asks that the phrases '''animals and plants''' and '''fauna and flora''' be replaced with '''animals, fungi, and plants''' and '''fauna, flora, and funga'''. |access-date=11 November 2021 |archive-date=11 November 2021 |archive-url=https://web.archive.org/web/20211111141833/https://www.iucn.org/sites/dev/files/statement-3f.pdf |url-status=dead }} [67] => [68] => == Characteristics == [69] => [[File:HYPHAE.png|thumb|'''Fungal hyphae cells''' [70] => {{image key [71] => |list type=ordered [72] => |Hyphal wall [73] => |[[Septum (cell biology)|Septum]] [74] => |[[Mitochondrion]] [75] => |[[Vacuole]] [76] => |[[Ergosterol]] crystal [77] => |[[Ribosome]] [78] => |Nucleus [79] => |[[Endoplasmic reticulum]] [80] => |[[Lipid body]] [81] => |[[Plasma membrane]] [82] => |[[Spitzenkörper]] [83] => |[[Golgi apparatus]] [84] => }} [85] => ]] [86] => [[File:Fungus cell cycle-en.svg|thumb|Fungal cell cycle showing [[dikaryon]]s typical of [[Dikarya|higher fungi]]]] [87] => Before the introduction of [[Molecular phylogenetics|molecular methods]] for phylogenetic analysis, [[Taxonomy (biology)|taxonomists]] considered fungi to be members of the [[Plant|plant kingdom]] because of similarities in lifestyle: both fungi and plants are mainly [[Sessility (zoology)|immobile]], and have similarities in general morphology and growth habitat. Although inaccurate, the common misconception that fungi are plants persists among the general public due to their historical classification, as well as several similarities.{{cite web |url=https://www.researchgate.net/publication/26571382 |title=Fifth-Grade Elementary School Students' Conceptions and Misconceptions about the Fungus Kingdom |access-date=5 October 2022}}{{cite web |url=https://www.cpp.edu/respect/resources/documents_kinder/pa_lessons_1-3/resources/gr0.kpa_common_student_ideas.pdf |title=Common Student Ideas about Plants and Animals |access-date=5 October 2022}} Like plants, fungi often grow in soil and, in the case of [[mushroom]]s, form conspicuous [[fruit bodies]], which sometimes resemble plants such as [[mosses]]. The fungi are now considered a separate kingdom, distinct from both plants and animals, from which they appear to have [[genetic divergence|diverged]] around one billion years ago (around the start of the [[Neoproterozoic]] 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: [88] => [89] => Shared features: [90] => * With other [[eukaryote]]s: Fungal cells contain [[membrane-bound]] [[cell nucleus|nuclei]] with [[chromosomes]] that contain [[DNA]] with [[Noncoding DNA|noncoding regions]] called [[intron]]s and coding regions called [[exons]]. Fungi have membrane-bound cytoplasmic [[organelles]] such as [[mitochondria]], [[sterol]]-containing membranes, and [[ribosomes]] of the [[80S]] type.{{sfn|Deacon|2005|p=4}} They have a characteristic range of soluble carbohydrates and storage compounds, including [[sugar alcohol]]s (e.g., [[mannitol]]), [[disaccharide]]s, (e.g., [[trehalose]]), and [[polysaccharide]]s (e.g., [[glycogen]], which is also found in animals{{sfn|Deacon|2005|pp=128–129}}). [91] => * With animals: Fungi lack [[chloroplast]]s and are [[heterotroph]]ic organisms and so require preformed [[organic compound]]s as energy sources.{{sfn|Alexopoulos|Mims|Blackwell|1996|pp=28–33}} [92] => * With plants: Fungi have a cell wall{{sfn|Alexopoulos|Mims|Blackwell|1996|pp=31–32}} and [[vacuole]]s. They reproduce by both sexual and asexual means, and like [[Primitive (phylogenetics)|basal]] plant groups (such as [[fern]]s and [[moss]]es) produce [[spore]]s. Similar to mosses and algae, fungi typically have [[haploid]] nuclei.{{sfn|Deacon|2005|p=58}} [93] => * With [[euglenoid]]s and bacteria: Higher fungi, euglenoids, and some bacteria produce the [[amino acid]] L-lysine in specific [[biosynthesis]] steps, called the [[α-aminoadipate pathway]]. [94] => * The cells of most fungi grow as tubular, elongated, and thread-like (filamentous) structures called [[hypha]]e, which may contain multiple nuclei and extend by growing at their tips. Each tip contains a set of aggregated [[vesicle (biology)|vesicles]]—cellular structures consisting of [[protein]]s, [[lipid]]s, and other organic molecules—called the [[Spitzenkörper]].{{sfn|Alexopoulos|Mims|Blackwell|1996|pp=27–28}} Both fungi and [[oomycete]]s grow as filamentous hyphal cells.{{sfn|Alexopoulos|Mims|Blackwell|1996|p=685}} In contrast, similar-looking organisms, such as filamentous [[green algae]], grow by repeated cell division within a chain of cells.{{sfn|Deacon|2005|pp=128–129}} There are also single-celled fungi ([[yeast]]s) that do not form hyphae, and some fungi have both hyphal and yeast forms.{{sfn|Alexopoulos|Mims|Blackwell|1996|p=30}} [95] => * In common with some plant and animal species, [[List of bioluminescent fungi|more than one hundred fungal species]] display [[bioluminescence]]. [96] => [97] => Unique features: [98] => * 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.{{sfn|Alexopoulos|Mims|Blackwell|1996|p=30}} [99] => * The fungal cell wall is made of a [[chitin-glucan complex]]; while glucans are also found in plants and chitin in the [[exoskeleton]] of [[arthropods]],{{sfn|Alexopoulos|Mims|Blackwell|1996|pp=32–33}} 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.{{sfn|Alexopoulos|Mims|Blackwell|1996|p=33}} [100] => [101] => [[File:Omphalotus nidiformis Binnamittalong 2 email.jpg|thumb|right|alt=A whitish fan or funnel-shaped mushroom growing at the base of a tree.|''[[Omphalotus nidiformis]]'', a bioluminescent mushroom]] [102] => 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 [[root]]s of plants. As eukaryotes, fungi possess a [[Biochemical pathway|biosynthetic pathway]] for producing [[terpene]]s 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 metabolite]]s that are similar or identical in structure to those made by plants. Many of the plant and fungal enzymes that make these compounds differ from each other in [[peptide sequence|sequence]] and other characteristics, which indicates separate origins and [[convergent evolution]] of these enzymes in the fungi and plants. [103] => [104] => == Diversity == [105] => [[File:Fungus in a Wood.JPG|thumb|[[Bracket fungus|Bracket fungi]] on a tree stump]] [106] => [107] => 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]] fungi ''[[Batrachochytrium dendrobatidis]]'' and ''[[Batrachochytrium salamandrivorans|B. salamandrivorans]]'', [[parasite]]s 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 themselves through water and enter their amphibian host. Other examples of aquatic fungi include those living in [[hydrothermal]] areas of the ocean. [108] => [109] => [[File:White fungus in wood chips.jpg|thumb|left|Widespread white fungus in wood chip mulch in an [[Oklahoma]] garden{{cite web |title=Fungi in Mulches and Composts |url=https://ag.umass.edu/landscape/fact-sheets/fungi-in-mulches-composts |website=[[University of Massachusetts Amherst]] |date=6 March 2015 |access-date=15 December 2022}}]] {{As of|2020|post=,}} around 148,000 species of fungi have been [[species description|described]] by [[Taxonomy (biology)|taxonomists]], but the global biodiversity of the fungus kingdom is not fully understood. A 2017 estimate suggests there may be between 2.2 and 3.8 million species. The number of new fungi species discovered yearly has increased from 1,000 to 1,500 per year about 10 years ago, to about 2,000 with a peak of more than 2,500 species in 2016. In the year 2019, 1,882 new species of fungi were described, and it was estimated that more than 90% of fungi remain unknown. The following year, 2,905 new species were described—the highest annual record of new fungus names. In mycology, species have historically been distinguished by a variety of methods and concepts. Classification based on [[morphology (biology)|morphological]] characteristics, such as the size and shape of spores or fruiting structures, has traditionally dominated fungal taxonomy.{{sfn|Kirk|Cannon|Minter|Stalpers|2008|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. [110] => [111] => == Mycology == [112] => [[File:Pier Antonio Micheli.jpg|thumb|upright|In 1729, [[Pier Antonio Micheli]] first published descriptions of fungi.]] [113] => [114] => [[Mycology]] is the branch of [[biology]] concerned with the systematic study of fungi, including their genetic and biochemical properties, their taxonomy, and their use to humans as a source of medicine, food, and [[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. [115] => [116] => 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.{{sfn|Ainsworth|1976|p=1}} [117] => [118] => === History === [119] => Mycology became a systematic science 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''.{{sfn|Alexopoulos|Mims|Blackwell|1996|pp=1–2}} 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.{{sfn|Ainsworth|1976|p=18}} Extending the use of the [[binomial nomenclature|binomial system of nomenclature]] introduced by [[Carl Linnaeus]] in his ''[[Species plantarum]]'' (1753), the Dutch [[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]] (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]], [[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]], and has challenged traditional morphology-based groupings in fungal [[Taxonomy (biology)|taxonomy]]. [120] => [121] => == Morphology == [122] => [123] => === Microscopic structures === [124] => [[File:Penicillium labeled cropped.jpg|thumb|upright=1.5|right|alt=Monochrome micrograph showing ''Penicillium'' hyphae as long, transparent, tube-like structures a few micrometres across. Conidiophores branch out laterally from the hyphae, terminating in bundles of phialides on which spherical condidiophores are arranged like beads on a string. Septa are faintly visible as dark lines crossing the hyphae.|An environmental isolate of ''[[Penicillium]]'' [125] => {{image key [126] => |list type=ordered [127] => |[[Hypha]] [128] => |[[Conidiophore]] [129] => |[[Phialide]] [130] => |[[Conidia]] [131] => |[[septum|Septa]] [132] => }} [133] => ]] [134] => Most fungi grow as [[hypha]]e, which are cylindrical, thread-like structures 2–10{{nbsp}}[[micrometres|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.{{sfn|Alexopoulos|Mims|Blackwell|1996|p=30}} 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 angle]]s to the cell wall giving the hypha its shape), with each compartment containing one or more nuclei; coenocytic hyphae are not compartmentalized.{{sfn|Deacon|2005|p=51}} Septa have [[Pit connection#Characteristics|pores]] that allow [[cytoplasm]], [[organelle]]s, and sometimes nuclei to pass through; an example is the [[dolipore septum]] in fungi of the phylum Basidiomycota.{{sfn|Deacon|2005|p=57}} Coenocytic hyphae are in essence [[multinucleate]] supercells. [135] => [136] => 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 [[mycorrhiza]]l fungi, which penetrate into the host cells to consume nutrients. [137] => [138] => Although fungi are [[opisthokont]]s—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 [[glucan]]s (e.g., [[Beta-glucan|β-1,3-glucan]]) and other typical components, also contains the [[biopolymer]] chitin. [139] => [140] => === Macroscopic structures === [141] => [[File:Armillaria ostoyae MO.jpg|right|thumb|alt=A cluster of large, thick-stem, light-brown gilled mushrooms growing at the base of a tree|''[[Armillaria solidipes]]'']] [142] => 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 (fungus)|mold]]s. Mycelia grown on solid [[agar]] media in laboratory [[petri dish]]es 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.{{sfn|Hanson|2008|pp=127–141}} 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 900{{nbsp}}[[hectare|ha]] (3.5 square miles), with an estimated age of nearly 9,000{{nbsp}}years. [143] => [144] => 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.{{sfn|Alexopoulos|Mims|Blackwell|1996|pp=204–205}} The fruit bodies of the basidiomycetes ([[basidiocarp]]s) and some ascomycetes can sometimes grow very large, and many are well known as [[mushroom]]s. [145] => [146] => == Growth and physiology == [147] => [[File:DecayingPeachSmall.gif|frame|left|alt=Time-lapse photography sequence of a [[peach]] becoming progressively discolored and disfigured|[[Mold (fungus)|Mold]] growth covering a decaying [[peach]]. The frames were taken approximately 12 hours apart over a period of six days.]] [148] => [149] => 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 ratio]]s. 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 pathogen]]s, 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 {{convert|8|MPa|lk=in}}. The filamentous fungus ''[[Paecilomyces lilacinus]]'' uses a similar structure to penetrate the eggs of [[nematode]]s. [150] => [151] => The mechanical pressure exerted by the appressorium is generated from physiological processes that increase intracellular [[turgor]] by producing [[osmolyte]]s 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]]s, [[protein]]s, and [[lipid]]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 [[biofilm]]s for substrate colonization and [[intercellular communication]]. [152] => [153] => Fungi are traditionally considered [[heterotroph]]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. [154] => [155] => == Reproduction == [156] => [[File:Polyporus squamosus Molter.jpg|thumb|right|alt=Two thickly stemmed brownish mushrooms with scales on the upper surface, growing out of a tree trunk|''[[Polyporus squamosus]]'']] [157] => Fungal reproduction is complex, reflecting the differences in lifestyles and genetic makeup within this diverse kingdom of organisms.{{sfn|Alexopoulos|Mims|Blackwell|1996|pp=48–56}} 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).{{sfn|Kirk|Cannon|Minter|Stalpers|2008|p=633}} 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 [[propagule]]s. [158] => [159] => === Asexual reproduction === [160] => [[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 (alternatively known as Deuteromycetes, conidial fungi, or mitosporic fungi) is not an accepted taxonomic clade and is now taken to mean simply fungi that lack a known sexual stage. [161] => [162] => === Sexual reproduction === [163] => {{see also|Mating in fungi|Sexual selection in fungi}} [164] => 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 system]]s: [[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. [165] => [166] => Most fungi have both a [[haploid]] and a [[diploid]] stage in their life cycles. In sexually reproducing fungi, compatible individuals may combine by fusing their hyphae together into an interconnected network; this process, [[anastomosis]], is required for the initiation of the sexual cycle. Many ascomycetes and basidiomycetes go through a [[dikaryotic]] stage, in which the nuclei inherited from the two parents do not combine immediately after cell fusion, but remain separate in the hyphal cells (see [[heterokaryosis]]).{{sfn|Jennings|Lysek|1996|pp=107–114}} [167] => [[File:Morelasci.jpg|thumb|left|alt=Microscopic view of numerous translucent or transparent elongated sac-like structures each containing eight spheres lined up in a row|The 8-spore [[ascus|asci]] of ''[[Morchella elata]]'', viewed with [[phase contrast microscopy]]]] [168] => [169] => 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 [[ascospore]]s. After dispersal, the ascospores may germinate and form a new haploid mycelium.{{sfn|Deacon|2005|p=31}} [170] => [171] => 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.{{sfn|Alexopoulos|Mims|Blackwell|1996|pp=492–493}} A [[basidiocarp]] is formed in which club-like structures known as [[basidia]] generate haploid [[basidiospores]] after karyogamy and meiosis.{{sfn|Jennings|Lysek|1996|p=142}} The most commonly known basidiocarps are mushrooms, but they may also take other forms (see [[#Morphology|Morphology]] section). [172] => [173] => 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 [[sporangiospore]]s. These sporangiospores allow the fungus to rapidly disperse and germinate into new genetically identical haploid fungal mycelia.{{sfn|Deacon|2005|pp=21–24}} [174] => [175] => === Spore dispersal === [176] => The spores of most of the researched species of fungi are transported by wind.{{cite web |url=http://www.botany.hawaii.edu/faculty/wong/BOT135/Lect05_b.htm |title=Spore Dispersal in Fungi |website=botany.hawaii.edu |access-date=28 December 2018 |archive-url=https://web.archive.org/web/20111117180734/http://www.botany.hawaii.edu/faculty/wong/BOT135/Lect05_b.htm|archive-date=17 November 2011 |url-status=live}}{{cite web |url=https://herbarium.usu.edu/fun-with-fungi/dispersal |title=Dispersal |website=herbarium.usu.edu |language=en |access-date=28 December 2018 |archive-url=https://web.archive.org/web/20181228223648/https://herbarium.usu.edu/fun-with-fungi/dispersal |archive-date=28 December 2018 |url-status=live}} Such species often produce dry or [[Hydrophobe|hydrophobic]] spores that do not absorb water and are readily scattered by raindrops, for example.{{cite journal |last1=Hassett |first1=Maribeth O. |last2=Fischer |first2=Mark W. F. |last3=Money |first3=Nicholas P. |title=Mushrooms as Rainmakers: How Spores Act as Nuclei for Raindrops |journal=[[PLOS ONE]] |date=28 October 2015 |volume=10 |issue=10 |pages=e0140407 |doi=10.1371/journal.pone.0140407 |pmid=26509436 |pmc=4624964 |bibcode=2015PLoSO..1040407H |language=en |issn=1932-6203 |doi-access=free | title-link=doi }}{{cite journal |last1=Kim |first1=Seungho |last2=Park |first2=Hyunggon |last3=Gruszewski |first3=Hope A. |last4=Schmale |first4=David G. |last5=Jung |first5=Sunghwan |title=Vortex-induced dispersal of a plant pathogen by raindrop impact |journal=[[Proceedings of the National Academy of Sciences]] |date=12 March 2019 |volume=116 |issue=11 |pages=4917–4922 |doi=10.1073/pnas.1820318116 |pmid=30804195 |pmc=6421443 |bibcode=2019PNAS..116.4917K |language=en |issn=0027-8424|doi-access=free | title-link=doi }} 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.[[File:Cyathus stercoreus Fruchtkörper.JPG|thumb|right|alt=A brown, cup-shaped fungus with several greyish disc-shaped structures lying within|The bird's nest fungus ''[[Cyathus stercoreus]]'']] Specialized mechanical and physiological mechanisms, as well as spore surface structures (such as [[hydrophobin]]s), 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 10,000{{nbsp}}[[G-force|g]]; the net result is that the spore is ejected 0.01–0.02{{nbsp}}cm, sufficient distance for it to fall through the [[Agaricales|gills]] or [[polypore|pores]] into the air below.{{sfn|Kirk|Cannon|Minter|Stalpers|2008|p=495}} 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.{{cite web |url=http://www.hampshirebiodiversity.org.uk/pdf/PublishedPlans/ToothFungiSAPfinal.pdf |title=Stipitate hydnoid fungi, Hampshire Biodiversity Partnership |access-date=13 November 2019 |archive-url=https://web.archive.org/web/20160304034104/http://www.hampshirebiodiversity.org.uk/pdf/PublishedPlans/ToothFungiSAPfinal.pdf |archive-date=4 March 2016 |url-status=live}} 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.{{sfn|Alexopoulos|Mims|Blackwell|1996|p=545}} [177] => [178] => === Homothallism === [179] => 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,{{cite journal |author=Dyer PS, O'Gorman CM |date=Jan 2012 |title=Sexual development and cryptic sexuality in fungi: insights from ''Aspergillus'' species |journal=[[FEMS Microbiology Reviews]] |volume=36 |issue=1| pages=165–192 |doi=10.1111/j.1574-6976.2011.00308.x |pmid=22091779 |doi-access=free | title-link=doi }} several species of the [[Ascomycota|ascomycete]] genus ''[[Cochliobolus]]'',{{cite journal |vauthors=Yun SH, Berbee ML, Yoder OC, Turgeon BG |year=1999 |title=Evolution of the fungal self-fertile reproductive life style from self-sterile ancestors |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=96 |issue=10| pages=5592–7 |doi=10.1073/pnas.96.10.5592 |pmid=10318929 |pmc=21905 |bibcode=1999PNAS...96.5592Y |doi-access=free | title-link=doi }} and the ascomycete ''[[Pneumocystis jirovecii]]''.{{cite journal |vauthors=Richard S, Almeida Jmgcf CO, Luraschi A, Nielsen O, Pagni M, Hauser PM |year=2018 |title=Functional and expression analyses of the ''Pneumocystis'' MAT genes suggest obligate sexuality through primary homothallism within host lungs |journal=[[mBio]] |volume=9| issue=1| doi=10.1128/mBio.02201-17 |pmid=29463658 |pmc=5821091}} The earliest mode of sexual reproduction among eukaryotes was likely homothallism, that is, [[selfing|self-fertile unisexual reproduction]].{{cite journal |last1=Heitman |first1=Joseph |title=Evolution of sexual reproduction: A view from the fungal kingdom supports an evolutionary epoch with sex before sexes |journal=[[Fungal Biology Reviews]] |volume=29 |issue=3–4 |year=2015 |pages=108–117 |doi=10.1016/j.fbr.2015.08.002 |pmid=26834823 |pmc=4730888 |doi-access=free | title-link=doi }} [180] => [181] => === Other sexual processes === [182] => 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.{{sfn|Jennings|Lysek|1996|pp=114–115}} 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. [183] => [184] => == Evolution == [185] => {{Main|Evolution of fungi}} [186] => 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]].{{sfn|Taylor|Taylor|1993|p=19}} Researchers study [[compression fossil]]s by dissolving the surrounding matrix with acid and then using light or [[scanning electron microscopy]] to examine surface details.{{sfn|Taylor|Taylor|1993|pp=7–12}} [187] => [[File:Prototaxites milwaukeensis.jpg|thumb|upright=0.8|left|''[[Prototaxites]] milwaukeensis'' (Penhallow, 1908)—a Middle [[Devonian]] fungus from [[Wisconsin]]]] [188] => The earliest fossils possessing features typical of fungi date to the [[Paleoproterozoic]] era, some {{ma|2400}} ([[annum|Ma]]); these multicellular [[benthic]] organisms had filamentous structures capable of [[anastomosis]].{{cite journal |last1=Bengtson |first1=Stefan |last2=Rasmussen |first2=Birger |last3=Ivarsson |first3=Magnus |last4=Muhling |first4=Janet |last5=Broman |first5=Curt |last6=Marone |first6=Federica |last7=Stampanoni |first7=Marco |last8=Bekker |first8=Andrey |s2cid=25586788 |title=Fungus-like mycelial fossils in 2.4-billion-year-old vesicular basalt |journal=[[Nature Ecology & Evolution]] |date=24 April 2017 |volume=1 |issue=6 |pages=0141 |doi=10.1038/s41559-017-0141 |pmid=28812648 |bibcode=2017NatEE...1..141B |hdl=20.500.11937/67718 |hdl-access=free |url=https://escholarship.org/uc/item/4883d4qh |access-date=15 July 2019 |archive-url=https://web.archive.org/web/20190715234418/https://escholarship.org/uc/item/4883d4qh |archive-date=15 July 2019 |url-status=live}} Other studies (2009) estimate the arrival of fungal organisms at about 760–1060{{nbsp}}Ma on the basis of comparisons of the rate of evolution in closely related groups. The oldest fossilizied mycelium to be identified from its molecular composition is between 715 and 810 million years old.[https://phys.org/news/2020-01-mushrooms-earlier-previously-thought.html First mushrooms appeared earlier than previously thought] For much of the [[Paleozoic]] Era (542–251{{nbsp}}Ma), the fungi appear to have been aquatic and consisted of organisms similar to the extant [[chytrid]]s in having flagellum-bearing spores. The evolutionary adaptation from an aquatic to a terrestrial lifestyle necessitated a diversification of ecological strategies for obtaining nutrients, including [[parasitism]], [[Saphrotrophic nutrition|saprobism]], and the development of [[Mutualism (biology)|mutualistic]] relationships such as [[mycorrhiza]] and lichenization.{{sfn|Taylor|Taylor|1993|pp=84–94 & 106–107}} Studies suggest that the ancestral ecological state of the [[Ascomycota]] was saprobism, and that independent [[lichen]]ization events have occurred multiple times. [189] => [190] => In May 2019, scientists reported the discovery of a [[fossil]]ized fungus, named ''[[Ourasphaira giraldae]]'', in the [[Northern Canada|Canadian Arctic]], that may have grown on land a billion years ago, well before [[plant]]s were living on land.{{cite news |last=Zimmer |first=Carl |author-link=Carl Zimmer |title=How Did Life Arrive on Land? A Billion-Year-Old Fungus May Hold Clues – A cache of microscopic fossils from the Arctic hints that fungi reached land long before plants. |date=22 May 2019 |work=[[The New York Times]] |url=https://www.nytimes.com/2019/05/22/science/fungi-fossils-plants.html |access-date=23 May 2019 |archive-url=https://web.archive.org/web/20190523011853/https://www.nytimes.com/2019/05/22/science/fungi-fossils-plants.html |archive-date=23 May 2019 |url-status=live}}{{cite journal |last1=Loron |first1=Corentin C. |last2=François |first2=Camille |last3=Rainbird |first3=Robert H. |last4=Turner |first4=Elizabeth C. |last5=Borensztajn |first5=Stephan |last6=Javaux |first6=Emmanuelle J. |s2cid=162180486 |title=Early fungi from the Proterozoic era in Arctic Canada |journal=[[Nature (journal)|Nature]] |volume=570 |issue=7760 |pages=232–235 |publisher=[[Springer Science+Business Media|Springer Science and Business Media LLC]] |date=22 May 2019 |issn=0028-0836 |doi=10.1038/s41586-019-1217-0 |pmid=31118507 |bibcode=2019Natur.570..232L}}{{cite web |last=Timmer |first=John |title=Billion-year-old fossils may be early fungus |website=[[Ars Technica]] |date=22 May 2019 |url=https://arstechnica.com/science/2019/05/billion-year-old-fossils-may-be-early-fungus/ |access-date=23 May 2019 |archive-url=https://web.archive.org/web/20190523003551/https://arstechnica.com/science/2019/05/billion-year-old-fossils-may-be-early-fungus/ |archive-date=23 May 2019 |url-status=live}} [[Permineralization#Pyritization|Pyritized]] fungus-like [[microfossil]]s preserved in the basal Ediacaran Doushantuo Formation (~635 Ma) have been reported in South China.{{cite journal |last1=Gan |first1=Tian |last2=Luo |first2=Taiyi |last3=Pang |first3=Ke |last4=Zhou |first4=Chuanming |last5=Zhou |first5=Guanghong |last6=Wan |first6=Bin |last7=Li |first7=Gang |last8=Yi |first8=Qiru |last9=Czaja |first9=Andrew D. |last10=Xiao |first10=Shuhai |date=28 January 2021 |title=Cryptic terrestrial fungus-like fossils of the early Ediacaran Period |journal=[[Nature Communications]] |language=en |volume=12 |issue=1 |pages=641 |doi=10.1038/s41467-021-20975-1 |pmid=33510166 |pmc=7843733 |bibcode=2021NatCo..12..641G |issn=2041-1723 |doi-access=free | title-link=doi }} Earlier, it had been presumed that the fungi colonized the land during the [[Cambrian]] (542–488.3{{nbsp}}Ma), also long before land plants. Fossilized hyphae and spores recovered from the [[Ordovician]] of Wisconsin (460{{nbsp}}Ma) 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]] (416–359.2{{nbsp}}Ma), when they occur abundantly in the [[Rhynie chert]], mostly as [[Zygomycota]] and [[Chytridiomycota]]. At about this same time, approximately 400{{nbsp}}Ma, the Ascomycota and Basidiomycota diverged, and all modern [[Class (biology)|classes]] of fungi were present by the Late [[Carboniferous]] ([[Pennsylvanian (geology)|Pennsylvanian]], 318.1–299{{nbsp}}Ma). [191] => [192] => Lichens formed a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 415{{nbsp}}Ma; 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]] from the Pennsylvanian. Rare in the fossil record are the Homobasidiomycetes (a [[taxon]] roughly equivalent to the mushroom-producing species of the [[Agaricomycetes]]). Two [[amber]]-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species ''[[Archaeomarasmius leggetti]]'') appeared during the late [[Cretaceous]], 90{{nbsp}}Ma. [193] => [194] => Some time after the [[Permian–Triassic extinction event]] (251.4{{nbsp}}Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in [[sediment]]s) 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. [195] => [196] => 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".{{cite journal |last1=Casadevall |first1=Arturo |last2=Heitman |first2=Joseph |title=Fungi and the Rise of Mammals |journal=PLOS Pathogens |date=16 August 2012 |volume=8 |issue=8 |pages=e1002808 |doi=10.1371/journal.ppat.1002808 |pmid=22916007 |pmc=3420938 |quote=That ecological calamity was accompanied by massive deforestation, an event followed by a fungal bloom, as the earth became a massive compost. |doi-access=free | title-link=doi }} [197] => [198] => == Taxonomy == [199] => Although commonly included in botany curricula and textbooks, fungi are more closely related to [[animal]]s than to plants and are placed with the animals in the [[monophyletic]] group of [[opisthokont]]s. Analyses using [[molecular phylogenetics]] 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 [[mating]]s.{{cite web |url=http://palaeos.com/fungi/fungi.html |title=Palaeos Fungi: Fungi |archive-url=https://web.archive.org/web/20120620205340/http://palaeos.com/fungi/fungi.html |archive-date=20 June 2012}} for an introduction to fungal taxonomy, including controversies. [https://web.archive.org/web/20041114121617/http://www.palaeos.com/Fungi/default.htm archive] [200] => [201] => 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]]). [202] => [203] => 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 [[phylum|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.{{cite journal |last1=Tedersoo |first1=Leho |last2=Sanchez-Ramırez |first2=Santiago |last3=Koljalg |first3=Urmas |last4=Bahram |first4=Mohammad |last5=Doring |first5=Markus |last6=Schigel |first6=Dmitry |last7=May |first7=Tom |last8=Ryberg |first8=Martin |last9=Abarenkov |first9=Kessy |title=High-level classification of the Fungi and a tool for evolutionary ecological analyses |journal=[[Fungal Diversity]] |date=22 February 2018 |volume=90 |issue=1 |pages=135–159 |doi=10.1007/s13225-018-0401-0 |doi-access=free | title-link=doi }} The lengths of the branches are not proportional to [[evolutionary]] distances. [204] => [205] => {{Clade [206] => |style=font-size:100%; line-height:80% [207] => |label1=[[Zoosporia]] [208] => |1={{clade [209] => |1={{clade [210] => |label1=Rozellomyceta [211] => |1={{clade [212] => |label1=[[Rozellomycota]] [213] => |1=[[Rozellomycetes]] [214] => |label2=[[Microsporidia|Microsporidiomycota]] [215] => |2={{clade [216] => |1=''[[Mitosporidium]]'' [217] => |2={{clade [218] => |1=''[[Paramicrosporidium]]'' [219] => |2={{clade [220] => |1=''[[Nucleophaga]]'' [221] => |2={{clade [222] => |1=[[Metchnikovellea]] [223] => |2=[[Microsporea]] [224] => }} [225] => }} [226] => }} [227] => }} [228] => }} [229] => }} [230] => |2={{clade [231] => |label1=Aphelidiomyceta [232] => |1={{clade [233] => |label1=[[Aphelida|Aphelidiomycota]] [234] => |1=[[Aphelida|Aphelidiomycetes]] [235] => }} [236] => |label2=[[Eumycota]] [237] => |2={{clade [238] => |1={{clade [239] => |label1=Chytridiomyceta [240] => |1={{clade [241] => |label1=[[Neocallimastigomycota]] [242] => |1=[[Neocallimastigomycetes]] [243] => |label2=[[Chytridiomycota]] [244] => |2={{clade [245] => |label1=[[Monoblepharomycotina]] [246] => |1={{clade [247] => |1=[[Hyaloraphidiomycetes]] [248] => |3=[[Sanchytriomycetes]] [249] => |2=[[Monoblepharidomycetes]] [250] => }} [251] => |label2=[[Chytridiomycotina]] [252] => |2={{clade [253] => |1=[[Mesochytriomycetes]] [254] => |2=[[Chytridiomycetes]] [255] => }} [256] => }} [257] => }} [258] => }} [259] => |2={{clade [260] => |label1=Blastocladiomyceta [261] => |label2=[[Amastigomycota]] [262] => |1={{clade [263] => |label1=[[Blastocladiomycota]] [264] => |1={{clade [265] => |1=[[Blastocladiomycetes]] [266] => |2=[[Physodermatomycetes]] [267] => }} [268] => }} [269] => |2={{clade [270] => |1={{clade [271] => |label1=Zoopagomyceta [272] => |1={{clade [273] => |label1=[[Basidiobolomycota]] [274] => |1={{clade [275] => |1=[[Basidiobolomycetes]] [276] => |2=[[Olpidiomycetes]] [277] => }} [278] => |2={{clade [279] => |label1=[[Entomophthoromycota]] [280] => |1={{clade [281] => |1=[[Neozygitomycetes]] [282] => |2=[[Entomophthoromycetes]] [283] => }} [284] => |label2=[[Kickxellomycota]] [285] => |2={{clade [286] => |label1=[[Zoopagomycotina]] [287] => |1=[[Zoopagomycetes]] [288] => |label2=[[Kickxellomycotina]] [289] => |2={{clade [290] => |1=[[Dimargaritomycetes]] [291] => |2=[[Kickxellomycetes]] [292] => }} [293] => }} [294] => }} [295] => }} [296] => }} [297] => |2={{clade [298] => |1={{clade [299] => |label1=[[Mortierellomycota]] [300] => |1=[[Mortierellomycetes]] [301] => }} [302] => |2={{clade [303] => |label1=Mucoromyceta [304] => |1={{clade [305] => |label1=[[Calcarisporiellomycota]] [306] => |1=[[Calcarisporiellomycetes]] [307] => |label2=[[Mucoromycota]] [308] => |2={{clade [309] => |1=[[Umbelopsidomycetes]] [310] => |2=[[Mucoromycetes]] [311] => }} [312] => }} [313] => |label2=[[Symbiomycota]] [314] => |2={{clade [315] => |label1=[[Glomeromycota]] [316] => |1={{clade [317] => |1=[[Paraglomeromycetes]] [318] => |2={{clade [319] => |1=[[Archaeosporomycetes]] [320] => |2=[[Glomeromycetes]] [321] => }} [322] => }} [323] => |label2=[[Dikarya]] [324] => |2={{clade [325] => |1={{clade [326] => |label1=[[Entorrhizomycota]] [327] => |1=[[Entorrhizomycetes]] [328] => |2=[[Basidiomycota]] [329] => }} [330] => |2=[[Ascomycota]] [331] => }} [332] => }} [333] => }} [334] => }} [335] => }} [336] => }} [337] => }} [338] => }} [339] => }} [340] => }} [341] => [342] => {{Clade [343] => |style=font-size:100%; line-height:80% [344] => |label1=[[Basidiomycota]] [345] => |1={{clade [346] => |label1=[[Pucciniomycotina]] [347] => |1={{clade [348] => |1={{clade [349] => |1=[[Tritirachiomycetes]] [350] => |2={{clade [351] => |1=[[Mixiomycetes]] [352] => |2=[[Agaricostilbomycetes]] [353] => }} [354] => }} [355] => |2={{clade [356] => |1=[[Cystobasidiomycetes]] [357] => |2={{clade [358] => |1={{clade [359] => |1=[[Classiculaceae]] [360] => |2=[[Microbotryomycetes]] [361] => }} [362] => |2={{clade [363] => |1=[[Cryptomycocolacomycetes]] [364] => |2={{clade [365] => |1=[[Atractiellomycetes]] [366] => |2=[[Pucciniomycetes]] [367] => }} [368] => }} [369] => }} [370] => }} [371] => }} [372] => |label2=[[Orthomycotina]] [373] => |2={{clade [374] => |label1=[[Ustilaginomycotina]] [375] => |1={{clade [376] => |1={{clade [377] => |1=[[Monilielliomycetes]] [378] => |2=[[Malasseziomycetes]] [379] => }} [380] => |2={{clade [381] => |1=[[Ustilaginomycetes]] [382] => |2=[[Exobasidiomycetes]] [383] => }} [384] => }} [385] => |label2=[[Agaricomycotina]] [386] => |2={{clade [387] => |1=?[[Geminibasidiomycetes]] [388] => |2=?[[Wallemiomycetes]] [389] => |3=[[Bartheletiomycetes]] [390] => |4={{clade [391] => |1=[[Tremellomycetes]] [392] => |2={{clade [393] => |1=[[Dacrymycetes]] [394] => |2=[[Agaricomycetes]] [395] => }} [396] => }} [397] => }} [398] => }} [399] => }} [400] => }} [401] => [402] => {{Clade [403] => |style=font-size:100%; line-height:80% [404] => |label1=[[Ascomycota]] [405] => |1={{clade [406] => |label1=[[Taphrinomycotina]] [407] => |1={{clade [408] => |1={{clade [409] => |1=[[Neolectomycetes]] [410] => |2=[[Taphrinomycetes]] [411] => }} [412] => |label2=[[Schizosaccharomyceta]] [413] => |2={{clade [414] => |1=[[Archaeorhizomycetes]] [415] => |2={{clade [416] => |1=[[Pneumocystidomycetes]] [417] => |2=[[Schizosaccharomycetes]] [418] => }} [419] => }} [420] => }} [421] => |label2=[[Saccharomyceta]] [422] => |2={{clade [423] => |label1=[[Saccharomycotina]] [424] => |1=[[Saccharomycetes]] [425] => |label2=[[Pezizomycotina]] [426] => |2={{clade [427] => |1=?[[Thelocarpales]] [428] => |2=?[[Vezdaeales]] [429] => |3=?[[Lahmiales]] [430] => |4=?[[Triblidiales]] [431] => |5=[[Orbiliomycetes]] [432] => |6={{clade [433] => |1=[[Pezizomycetes]] [434] => |label2=[[Leotiomyceta]] [435] => |2={{clade [436] => |label1=[[Sordariomyceta]] [437] => |1={{clade [438] => |1=[[Xylonomycetes]] [439] => |2={{clade [440] => |1=[[Geoglossomycetes]] [441] => |2={{clade [442] => |1=[[Leotiomycetes]] [443] => |2={{clade [444] => |1=[[Laboulbeniomycetes]] [445] => |2=[[Sordariomycetes]] [446] => }} [447] => }} [448] => }} [449] => }} [450] => |label2=[[Dothideomyceta]] [451] => |2={{clade [452] => |1={{clade [453] => |1=[[Coniocybomycetes]] [454] => |2=[[Lichinomycetes]] [455] => }} [456] => |2={{clade [457] => |1={{clade [458] => |1=[[Eurotiomycetes]] [459] => |2=[[Lecanoromycetes]] [460] => }} [461] => |2={{clade [462] => |1=[[Collemopsidiomycetes]] [463] => |2={{clade [464] => |1=[[Arthoniomycetes]] [465] => |2=[[Dothideomycetes]] [466] => }} [467] => }} [468] => }} [469] => }} [470] => }} [471] => }} [472] => }} [473] => }} [474] => }} [475] => }} [476] => [477] => === Taxonomic groups === [478] => {{See also|List of fungal orders}} [479] => [480] => [[File:02 01 groups of Fungi (M. Piepenbring).png|thumb|right|upright=1.5|Main groups of fungi]] [481] => The major [[phylum|phyla]] (sometimes called divisions) of fungi have been classified mainly on the basis of characteristics of their sexual [[reproduction|reproductive]] structures. {{As of|2019}}, nine major [[Lineage (evolution)|lineages]] have been identified: [[Opisthosporidia]], [[Chytridiomycota]], [[Neocallimastigomycota]], [[Blastocladiomycota]], [[Zoopagomycotina|Zoopagomycotina]], [[Mucoromycota]], [[Glomeromycota]], [[Ascomycota]] and [[Basidiomycota]]. [482] => [483] => 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. [484] => [485] => 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 [[zoospore]]s that are capable of active movement through aqueous phases with a single [[flagellum]], leading early [[taxonomist]]s to classify them as [[protist]]s. [[Molecular phylogenetics|Molecular phylogenies]], inferred from [[rRNA]] sequences in [[ribosome]]s, suggest that the Chytrids are a [[Basal (phylogenetics)|basal]] group divergent from the other fungal phyla, consisting of four major [[clade]]s with suggestive evidence for [[paraphyly]] or possibly [[polyphyly]]. [486] => [487] => 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]]. [488] => [489] => The [[Neocallimastigomycota]] were earlier placed in the phylum Chytridiomycota. Members of this small phylum are [[anaerobic organism]]s, living in the digestive system of larger herbivorous mammals and in other terrestrial and aquatic environments enriched in cellulose (e.g., domestic waste landfill sites). They lack [[mitochondria]] but contain [[hydrogenosome]]s of mitochondrial origin. As in the related chrytrids, neocallimastigomycetes form zoospores that are posteriorly uniflagellate or polyflagellate. [490] => [491] => [[File:Arbuscular mycorrhiza microscope.jpg|thumb|right|alt=Microscopic view of a layer of translucent grayish cells, some containing small dark-color spheres|''[[Arbuscular mycorrhiza]]'' seen under microscope. [[Flax]] root cortical cells containing paired arbuscules.]] [492] => [[File:Ascocarp2.png|thumb|right|alt=Cross-section of a cup-shaped structure showing locations of developing meiotic asci (upper edge of cup, left side, arrows pointing to two gray cells containing four and two small circles), sterile hyphae (upper edge of cup, right side, arrows pointing to white cells with a single small circle in them), and mature asci (upper edge of cup, pointing to two gray cells with eight small circles in them)|Diagram of an [[apothecium]] (the typical cup-like reproductive structure of Ascomycetes) showing sterile tissues as well as developing and mature asci.]] [493] => 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]]s several meters through the air.{{sfn|Alexopoulos|Mims|Blackwell|1996|p=145}} Medically relevant genera include ''[[Mucor]]'', ''[[Rhizomucor]]'', and ''[[Rhizopus]]''. [494] => [495] => The [[Ascomycota]], commonly known as sac fungi or ascomycetes, constitute the largest taxonomic group within the Eumycota.{{sfn|Kirk|Cannon|Minter|Stalpers|2008|p=489}} These fungi form meiotic spores called [[ascospore]]s, which are enclosed in a special sac-like structure called an [[ascus]]. This phylum includes [[morel]]s, a few [[mushroom]]s and [[Tuber (genus)|truffles]], unicellular [[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 [[anamorph]]ic species), but analysis of molecular data has often been able to identify their closest [[teleomorph]]s 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]]''). [496] => [497] => Members of the [[Basidiomycota]], commonly known as the club fungi or basidiomycetes, produce meiospores called [[basidiospore]]s 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]]''. [498] => [499] => === Fungus-like organisms === [500] => Because of similarities in morphology and lifestyle, the [[slime mold]]s ([[mycetozoa]]ns, [[plasmodiophorid]]s, [[acrasid]]s, ''[[Fonticula]]'' and [[labyrinthulid]]s, now in [[Amoebozoa]], [[Rhizaria]], [[Excavata]], [[Opisthokonta]] and [[Stramenopiles]], respectively), water molds ([[oomycete]]s) and [[hyphochytrid]]s (both [[Stramenopiles]]) were formerly classified in the kingdom Fungi, in groups like [[Mastigomycotina]], [[Gymnomycota]] and [[Phycomycetes]]. The slime molds were studied also as [[protozoa]]ns, leading to an [[ambiregnal]], duplicated taxonomy. [501] => [502] => Unlike true fungi, the [[cell wall]]s of oomycetes contain [[cellulose]] and lack [[chitin]]. Hyphochytrids have both chitin and cellulose. Slime molds lack a cell wall during the assimilative phase (except labyrinthulids, which have a wall of scales), and take in 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, [[taxonomist]]s no longer group them in the kingdom Fungi. Nonetheless, studies of the oomycetes and myxomycetes are still often included in [[mycology]] textbooks and primary research literature. [503] => [504] => The [[Eccrinales]] and [[Amoebidiales]] are [[opisthokont]] [[protist]]s, 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. [505] => [506] => 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. [507] => [508] => The [[nucleariid]]s may be the next sister group to the eumycete clade, and as such could be included in an expanded fungal kingdom. [509] => Many [[Actinomycetales]] ([[Actinomycetota]]), a group with many filamentous bacteria, were also long believed to be fungi.{{cite book |last1=Amoroso |first1=Maria Julia |last2=Benimeli |first2=Claudia Susana |last3=Cuozzo |first3=Sergio Antonio |title=Actinobacteria : application in bioremediation and production of industrial enzymes |date=2013 |publisher=[[CRC Press]], [[Taylor & Francis]] Group |isbn=9781466578739 |page=33 |url=https://www.crcpress.com/Actinobacteria-Application-in-Bioremediation-and-Production-of-Industrial/Amoroso-Benimeli-Cuozzo/p/book/9781466578739 |language=en}}{{cite web |url=https://www.humankindoregon.com/soil-biology |title=An Introduction to Soil Biology |website=Humankind Oregon}} [510] => [511] => == Ecology == [512] => [[File:PinMould on Peach LowMag Scale.jpg|thumb|A pin mold decomposing a peach]] [513] => 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 [[saprotroph]]s and [[symbiont]]s, degrading [[organic matter]] to inorganic molecules, which can then re-enter anabolic metabolic pathways in plants or other organisms. [514] => [515] => === Symbiosis === [516] => 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. [517] => [518] => ==== With plants ==== [519] => [[Mycorrhiza]]l 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. [520] => [[File:Neotyphodium coenophialum.jpg|thumb|left|upright|alt=A microscopic view of blue-stained cells, some with dark wavy lines in them|The dark filaments are [[hyphae]] of the endophytic fungus ''[[Epichloë coenophiala]]'' in the intercellular spaces of [[Festuca arundinacea|tall fescue]] leaf sheath tissue]] [521] => 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.{{cite journal |last=Heijden |first=Marcel G. A. van der |s2cid=133399719 |date=15 April 2016 |title=Underground networking |journal=[[Science (journal)|Science]] |language=en |volume=352 |issue=6283 |pages=290–291 |doi=10.1126/science.aaf4694 |issn=0036-8075 |pmid=27081054 |bibcode=2016Sci...352..290H}} Such mycorrhizal communities are called "common [[mycorrhizal network]]s".{{cite web |url=https://www.theatlantic.com/science/archive/2016/04/the-wood-wide-web/478224/ |title=Trees Have Their Own Internet |last=Yong |first=Ed |date=14 April 2016 |website=[[The Atlantic]] |language=en-US |access-date=9 March 2019 |archive-url=https://web.archive.org/web/20190328164827/https://www.theatlantic.com/science/archive/2016/04/the-wood-wide-web/478224/ |archive-date=28 March 2019 |url-status=live}} 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. [522] => [523] => ==== With algae and cyanobacteria ==== [524] => [[File:Lobaria pulmonaria 010108a.jpg|right|thumb|alt=A green, leaf-like structure attached to a tree, with a pattern of ridges and depression on the bottom surface|The lichen ''[[Lobaria pulmonaria]]'', a symbiosis of fungal, [[algae|algal]], and [[cyanobacteria]]l species]] [525] => [[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 [[ascomycete]]s and a few [[basidiomycete]]s. 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.{{sfn|Deacon|2005|p=267}} They are able to grow on inhospitable surfaces, including bare soil, rocks, [[Bark (botany)|tree bark]], wood, shells, barnacles and leaves. As in [[mycorrhiza]]s, 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.{{sfn|Kirk|Cannon|Minter|Stalpers|2008|p=378}} 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.{{sfn|Deacon|2005|pp=267–276}} [526] => [527] => ==== With insects ==== [528] => 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.{{sfn|Deacon|2005|p=277}} 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 [[neuroptera]]ns, [[beetle]]s, and [[cockroach]]es; 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]]).{{cite journal |last1=Filipiak |first1=Michał |last2=Weiner |first2=January |date=March 2017 |title=Nutritional dynamics during the development of xylophagous beetles related to changes in the stoichiometry of 11 elements |journal= [[Physiological Entomology]] |volume=42 |issue=1 |pages=73–84 |doi=10.1111/phen.12168 |doi-access=free | title-link=doi }}{{cite book |last1=Ulyshen |first1=Michael D. |title=Saproxylic Insects |chapter=Nutrient Dynamics in Decomposing Dead Wood in the Context of Wood Eater Requirements: The Ecological Stoichiometry of Saproxylophagous Insects |series=Zoological Monographs |date=2018 |volume=1 |publisher=Springer, Cham |isbn=978-3-319-75937-1 |pages=429–469 |doi=10.1007/978-3-319-75937-1_13 |url=https://depot.ceon.pl/handle/123456789/15394?show=full}}{{cite book |last1=Ulyshen |first1=Michael D. |title=Saproxylic Insects |chapter=Insect-Fungus Interactions in Dead Wood Systems |series=Zoological Monographs |date=2018 |volume=1 |publisher=Springer, Cham |isbn=978-3-319-75936-4 |pages=377–427 |doi=10.1007/978-3-319-75937-1_12}} They deliver nutrients needed by [[Xylophagy|xylophages]] to nutritionally scarce dead wood.{{cite journal |last1=Filipiak |first1=Michał |last2=Sobczyk |first2=Łukasz |last3=Weiner |first3=January |year=2016 |title=Fungal transformation of tree stumps into a suitable resource for xylophagous beetles via changes in elemental ratios |journal=Insects |volume=7 |issue=2 |pages=13 |doi=10.3390/insects7020013 |pmc=4931425 |doi-access=free | title-link=doi }} 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]]. [529] => [530] => {{anchor|Parasite}} [531] => {{anchor|Pathogen}} [532] => [533] => {{anchor|Necrotroph}} [534] => [535] => ==== As pathogens and parasites ==== [536] => [[File:Aecidium magnellanicum.jpg|right|thumb|alt=A thin brown stick positioned horizontally with roughly two dozen clustered orange-red leaves originating from a single point in the middle of the stick. These orange leaves are three to four times larger than the few other green leaves growing out of the stick, and are covered on the lower leaf surface with hundreds of tiny bumps. The background shows the green leaves and branches of neighboring shrubs.|The plant pathogen ''Puccinia magellanicum'' ([[calafate rust]]) causes the defect known as [[witch's broom]], seen here on a [[barberry]] shrub in Chile.]] [[File:Candida Gram stain.jpg|thumb|right|[[Gram stain]] of ''Candida albicans'' from a vaginal swab from a woman with [[candidiasis]], showing hyphae, and chlamydospores, which are 2–4 [[micrometre|µm]] in diameter.]] [537] => [538] => Many fungi are [[parasite]]s on plants, animals (including humans), and other fungi. Serious pathogens of many cultivated plants causing extensive damage and losses to agriculture and forestry include the [[rice blast]] fungus ''[[Magnaporthe oryzae]]'', tree pathogens such as ''[[Ophiostoma ulmi]]'' and ''[[Ophiostoma novo-ulmi]]'' causing [[Dutch elm disease]], ''[[Cryphonectria parasitica]]'' responsible for [[chestnut blight]], and ''[[Texas Root Rot|Phymatotrichopsis omnivora]]'' causing [[Texas Root Rot]], 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).{{cite journal|last1=Koeck |first1=M. |last2=Hardham |first2=A.R. |last3=Dodds |last4=P.N. |date=2011 |volume=13 |issue=12 |pages=1849–1857 |title=The role of effectors of biotrophic and hemibiotrophic fungi in infection |journal=Cellular Microbiology |doi=10.1111/j.1462-5822.2011.01665.x|pmid=21848815 |pmc=3218205 }} 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.{{cite web |url=https://www.researchgate.net/publication/6114636 |title=''Asterotremella gen. nov. albida'', an anamorphic tremelloid yeast isolated from the agarics ''Asterophora lycoperdoides'' and ''Asterophora parasitica'' |via=[[ResearchGate]] |language=en |access-date=19 April 2019}} [539] => [540] => 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, a person with [[immunodeficiency]] is more 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. [541] => [542] => ==== As targets of mycoparasites ==== [543] => 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. [544] => Fungi can also become the target of infection by [[mycovirus]]es.{{cite journal |vauthors=Pearson MN, Beever RE, Boine B, Arthur K |title=Mycoviruses of filamentous fungi and their relevance to plant pathology |journal=[[Molecular Plant Pathology]] |volume=10 |issue=1 |pages=115–28 |date=January 2009 |pmid=19161358 |doi=10.1111/j.1364-3703.2008.00503.x |pmc=6640375}}{{cite journal |vauthors=Bozarth RF |title=Mycoviruses: a new dimension in microbiology |journal=[[Environmental Health Perspectives]] |volume=2 |issue=1 |pages=23–39 |date=October 1972 |pmid=4628853 |pmc=1474899 |doi=10.1289/ehp.720223}} [545] => [546] => === Communication === [547] => {{Main|Mycorrhizal networks}} [548] => There appears to be electrical communication between fungi in word-like components according to spiking characteristics.{{cite journal |last1=Adamatzky |first1=Andrew |title=Language of fungi derived from their electrical spiking activity |journal=[[Royal Society Open Science]] |year=2022 |volume=9 |issue=4 |pages=211926 |doi=10.1098/rsos.211926|pmid=35425630 |pmc=8984380 |arxiv=2112.09907 |bibcode=2022RSOS....911926A}} [549] => [550] => === Possible impact on climate === [551] => According to a study published in the academic journal [[Current Biology]], fungi can soak from the atmosphere around 36% of global fossil fuel [[greenhouse gas emissions]].{{cite news |last1=ELBEIN |first1=SAUL |title=Fungi may offer 'jaw-dropping' solution to climate change |url=https://thehill.com/policy/equilibrium-sustainability/4034986-fungi-may-offer-jaw-dropping-solution-to-climate-change/ |access-date=6 June 2023 |agency=The Hill |date=6 June 2023}}{{cite web |title=Fungi stores a third of carbon from fossil fuel emissions and could be essential to reaching net zero, new study reveals |url=https://www.eurekalert.org/news-releases/991288 |website=EurekAlert |publisher=UNIVERSITY OF SHEFFIELD |access-date=6 June 2023}} [552] => [553] => == Mycotoxins == [554] => [[File:Ergotamine3.png|thumb|right|alt=(6aR,9R)-N-((2R,5S,10aS,10bS)-5-benzyl-10b-hydroxy-2-methyl-3,6-dioxooctahydro-2H-oxazolo[3,2-a] pyrrolo[2,1-c]pyrazin-2-yl)-7-methyl-4,6,6a,7,8,9-hexahydroindolo[4,3-fg] quinoline-9-carboxamide|[[Ergotamine]], a major mycotoxin produced by ''[[Claviceps]]'' species, which if ingested can cause [[gangrene]], convulsions, and [[hallucination]]s]] [555] => 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 [[amatoxin]]s 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 [[cereal]]s contaminated with [[sclerotia]] of the ergot fungus, ''[[Claviceps purpurea]]''. Other notable mycotoxins include the [[aflatoxin]]s, 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, [[ochratoxin]]s, [[patulin]], and [[trichothecene]]s (e.g., [[T-2 mycotoxin]]) and [[fumonisin]]s, which have significant impact on human food supplies or animal [[livestock]]. [556] => [557] => Mycotoxins are secondary metabolites (or [[natural product]]s), and research has established the existence of biochemical pathways solely for the purpose of producing mycotoxins and other natural products in fungi. Mycotoxins may provide [[Fitness (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|Human use below]]. [558] => [559] => == Pathogenic mechanisms == [560] => ''[[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. [561] => [562] => ''[[Cryptococcus neoformans]]'' is an encapsulated yeast that can live in both plants and animals. ''C.{{nbsp}}neoformans'' usually infects the lungs, where it is phagocytosed by [[alveolar macrophage]]s. Some ''C.{{nbsp}}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.{{nbsp}}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.{{nbsp}}neoformans'' are mating "type a". Filaments of mating "type a" ordinarily have haploid nuclei, but they can become diploid (perhaps by endoduplication or by stimulated nuclear fusion) to form [[blastospore]]s. The diploid nuclei of blastospores can undergo meiosis, including recombination, to form haploid basidiospores that can be dispersed. This process is referred to as monokaryotic fruiting. This process requires a gene called ''[[DMC1]]'', 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.{{nbsp}}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. [563] => [564] => == Human use == [565] => {{See also|Human interactions with fungi}} [566] => [[File:S cerevisiae under DIC microscopy.jpg|thumb|upright|right|alt=Microscopic view of five spherical structures; one of the spheres is considerably smaller than the rest and attached to one of the larger spheres|''[[Saccharomyces cerevisiae]]'' cells shown with [[Differential interference contrast microscopy|DIC microscopy]]]] [567] => 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. Fungi-based industries are sometimes considered to be a major part of a growing [[bioeconomy]], with applications under [[research and development]] including use for textiles, [[Environmental impact of meat production|meat]] substitution and general fungal biotechnology.{{cite journal |last1=Meyer |first1=Vera |last2=Basenko |first2=Evelina Y. |last3=Benz |first3=J. Philipp |last4=Braus |first4=Gerhard H. |last5=Caddick |first5=Mark X. |last6=Csukai |first6=Michael |last7=de Vries |first7=Ronald P. |last8=Endy |first8=Drew |last9=Frisvad |first9=Jens C. |last10=Gunde-Cimerman |first10=Nina |last11=Haarmann |first11=Thomas |last12=Hadar |first12=Yitzhak |last13=Hansen |first13=Kim |last14=Johnson |first14=Robert I. |last15=Keller |first15=Nancy P. |last16=Kraševec |first16=Nada |last17=Mortensen |first17=Uffe H. |last18=Perez |first18=Rolando |last19=Ram |first19=Arthur F. J. |last20=Record |first20=Eric |last21=Ross |first21=Phil |last22=Shapaval |first22=Volha |last23=Steiniger |first23=Charlotte |last24=van den Brink |first24=Hans |last25=van Munster |first25=Jolanda |last26=Yarden |first26=Oded |last27=Wösten |first27=Han A. B. |title=Growing a circular economy with fungal biotechnology: a white paper |journal=[[Fungal Biology and Biotechnology]] |date=2 April 2020 |volume=7 |issue=1 |pages=5 |doi=10.1186/s40694-020-00095-z |pmid=32280481 |pmc=7140391 |s2cid=215411291 |issn=2054-3085 |doi-access=free | title-link=doi }}{{cite journal |last1=Jones |first1=Mitchell |last2=Gandia |first2=Antoni |last3=John |first3=Sabu |last4=Bismarck |first4=Alexander |title=Leather-like material biofabrication using fungi |journal=[[Nature Sustainability]] |date=January 2021 |volume=4 |issue=1 |pages=9–16 |doi=10.1038/s41893-020-00606-1 |s2cid=221522085 |language=en |issn=2398-9629}}{{cite web |title=Plant-based meat substitutes - products with future potential {{!}} Bioökonomie.de |url=https://biooekonomie.de/en/topics/in-depth-reports/plant-based-meat-substitutes-products-future-potential |website=biooekonomie.de |access-date=25 May 2022 |language=en}}{{cite web |last1=Berlin |first1=Kustrim CerimiKustrim Cerimi studied biotechnology at the Technical University in |last2=biotechnology |first2=is currently doing his PhD He is interested in the broad field of fungal |last3=Artists |first3=Has Collaborated in Various Interdisciplinary Projects with |last4=Artists |first4=Hybrid |title=Mushroom meat substitutes: A brief patent overview |url=https://blogs.biomedcentral.com/on-biology/2022/01/28/mushroom-meat-substitutes-a-brief-patent-overview/ |website=On Biology |access-date=25 May 2022 |date=28 January 2022}}{{cite journal |last1=Lange |first1=Lene |title=The importance of fungi and mycology for addressing major global challenges* |journal=IMA Fungus |date=December 2014 |volume=5 |issue=2 |pages=463–471 |doi=10.5598/imafungus.2014.05.02.10 |pmid=25734035 |pmc=4329327 |issn=2210-6340}} [568] => [569] => === Therapeutic uses === [570] => [571] => ==== Modern chemotherapeutics ==== [572] => {{See also|Medicinal fungi}} [573] => [[File:Penicillium rubens (type specimen).png|thumb|left|The mold ''[[Penicillium rubens]]'' was the source of [[penicillin G]].{{cite journal |vauthors=Pathak A, Nowell RW, Wilson CG, Ryan MJ, Barraclough TG |date=September 2020 |title=Comparative genomics of Alexander Fleming's original ''Penicillium'' isolate (IMI 15378) reveals sequence divergence of penicillin synthesis genes |journal=[[Scientific Reports]] |volume=10 |issue=1 |pages=Article 15705 |doi=10.1038/s41598-020-72584-5 |pmid=32973216 |pmc=7515868 |bibcode=2020NatSR..1015705P}}]] [574] => [575] => Many species produce metabolites that are major sources of [[pharmacology|pharmacologically]] active drugs. [576] => [577] => ===== Antibiotics ===== [578] => Particularly important are the antibiotics, including the [[penicillin]]s, a structurally related group of [[β-lactam antibiotic]]s that are synthesized from small [[peptide]]s. 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. [579] => [580] => ===== Other ===== [581] => Other drugs produced by fungi include [[griseofulvin]] isolated from ''[[Penicillium griseofulvum]]'', used to treat fungal infections, and [[statin]]s ([[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]]. [[Psilocybin]] from [[Psilocybin mushroom|fungi]] is investigated [[Psilocybin therapy|for therapeutic use]] and appears to cause [[Brain connectivity estimators|global increases in brain]] [[Large-scale brain network|network]] [[Functional integration (neurobiology)|integration]].{{cite journal |last1=Daws |first1=Richard E. |last2=Timmermann |first2=Christopher |last3=Giribaldi |first3=Bruna |last4=Sexton |first4=James D. |last5=Wall |first5=Matthew B. |last6=Erritzoe |first6=David |last7=Roseman |first7=Leor |last8=Nutt |first8=David |last9=Carhart-Harris |first9=Robin|author9-link=Robin Carhart-Harris |title=Increased global integration in the brain after psilocybin therapy for depression |journal=[[Nature Medicine]] |date=April 2022 |volume=28 |issue=4 |pages=844–851 |doi=10.1038/s41591-022-01744-z |pmid=35411074 |s2cid=248099554 |language=en |issn=1546-170X|doi-access=free | title-link=doi |hdl=10044/1/95521 |hdl-access=free }} Fungi produce compounds that inhibit [[virus]]es 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. [582] => [583] => === Traditional medicine === [584] => {{multiple image [585] => | align = right [586] => | image1 = Ganoderma lucidum 01.jpg [587] => | width1 = 140 [588] => | alt1 = Upper surface view of a kidney-shaped fungus, brownish-red with a lighter yellow-brown margin, and a somewhat varnished or shiny appearance [589] => | caption1 = [590] => | image2 = CordycepsSinensis.jpg [591] => | width2 = 180 [592] => | alt2 = Two dried yellow-orange caterpillars, one with a curly grayish fungus growing out of one of its ends. The grayish fungus is roughly equal to or slightly greater in length than the caterpillar, and tapers in thickness to a narrow end. [593] => | caption2 = [594] => | footer = The fungi ''[[Ganoderma lucidum]]'' (left) and ''[[Ophiocordyceps sinensis]]'' (right) are used in traditional medicine practices [595] => }} [596] => [597] => 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]]''. [598] => [599] => === Cultured foods === [600] => [[Baker's yeast]] or ''[[Saccharomyces cerevisiae]]'', a unicellular fungus, is used to make [[bread]] and other wheat-based products, such as [[pizza]] dough and [[dumpling]]s. Yeast species of the genus ''[[Saccharomyces]]'' are also used to produce [[alcoholic beverage]]s through fermentation. Shoyu koji mold (''[[Aspergillus oryzae]]'') is an essential ingredient in brewing [[Shoyu]] ([[soy sauce]]) and [[sake]], and the preparation of [[miso]], while ''[[Rhizopus]]'' species are used for making [[tempeh]]. Several of these fungi are [[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]]''. [601] => [602] => === In food === [603] => [[File:Asian mushrooms.jpg|thumb|left|upright=1.35|A selection of [[edible mushroom]]s eaten in Asia]] [604] => [605] => [[Edible mushroom]]s 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 store]]s and markets, including straw mushrooms (''[[Volvariella volvacea]]''), oyster mushrooms (''[[Pleurotus ostreatus]]''), shiitakes (''[[Lentinula edodes]]''), and [[enokitake]] (''[[Flammulina]]'' spp.). [606] => [607] => [[File:Blue Stilton Quarter Front.jpg|thumb|alt=A corner of cheese with greenish streaks through it|[[Stilton cheese]] veined with ''[[Penicillium roqueforti]]'']] [608] => [609] => Many other mushroom species are [[Mushroom hunting|harvested from the wild]] for personal consumption or commercial sale. [[Lactarius deliciosus|Milk mushrooms]], [[morel]]s, [[chanterelle]]s, [[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.{{sfn|Hall|2003|pp=13–26}} [610] => [611] => 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. [612] => [613] => === Poisonous fungi === [614] => [[File:Amanita phalloides 1.JPG|upright|thumb|alt=Two light yellow-green mushrooms with stems and caps, one smaller and still in the ground, the larger one pulled out and laid beside the other to show its bulbous stem with a ring|''[[Amanita phalloides]]'' accounts for the majority of fatal [[mushroom poisoning]]s worldwide. It sometimes lacks the greenish color seen here.]] [615] => [616] => Many mushroom species are [[Mushroom poisoning|poisonous]] to humans and cause a range of reactions including slight digestive problems, [[allergy|allergic]] reactions, [[hallucination]]s, 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.{{nbsp}}virosa]])'' and the death cap ''([[Amanita phalloides|A.{{nbsp}}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]]. [617] => [618] => 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.{{sfn|Hall|2003|p=7}} [619] => [620] => === Pest control === [621] => {{Main|Biological pest control#Fungi}} [622] => [[File:Beauveria.jpg|thumb|left|alt=Two dead grasshoppers with a whitish fuzz growing on them|Grasshoppers killed by ''[[Beauveria bassiana]]'']] [623] => [624] => In agriculture, fungi may be useful if they actively compete for nutrients and space with [[pathogen]]ic microorganisms such as bacteria or other fungi via the [[competitive exclusion principle]], or if they are [[parasitism|parasites]] of these pathogens. For example, certain species eliminate or suppress the growth of harmful plant pathogens, such as insects, [[mites]], [[weed]]s, [[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 insecticide]]s 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. [625] => [626] => === Bioremediation === [627] => {{See also|Mycoremediation}} [628] => Certain fungi, in particular [[white rot|white-rot]] fungi, can degrade [[insecticide]]s, [[herbicide]]s, [[pentachlorophenol]], [[creosote]], [[coal tar]]s, and heavy fuels and turn them into [[carbon dioxide]], water, and basic elements. Fungi have been shown to [[biomineralization|biomineralize]] [[uranium#Oxides|uranium oxides]], suggesting they may have application in the [[bioremediation]] of radioactively polluted sites. [629] => [630] => === Model organisms === [631] => 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]]. [632] => [633] => === Others === [634] => 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 [[lipase]]s used in [[biological detergent]]s, [[cellulase]]s used in making [[cellulosic ethanol]] and [[stonewashed jeans]], and [[amylase]]s, [[invertase]]s, [[protease]]s and [[xylanase]]s. [635] => [636] => == See also == [637] => {{Portal|Fungi}} [638] => {{div col}} [639] => * [[Conservation of fungi]] [640] => * [[Fantastic Fungi]] [641] => * [[Glossary of mycology]] [642] => * [[Marine fungi]] [643] => * [[Fungal infection]] [644] => * [[Outline of fungi]] [645] => ** [[Outline of lichens]] [646] => * [[Fungi in art]] [647] => {{div col end}} [648] => {{clear}} [649] => [650] => == References == [651] => === Citations === [652] => {{Reflist|refs= [653] => [654] => {{cite journal |vauthors=Aanen DK |title=As you reap, so shall you sow: coupling of harvesting and inoculating stabilizes the mutualism between termites and fungi |journal=[[Biology Letters]] |volume=2 |issue=2 |pages=209–12 |date=June 2006 |pmid=17148364 |pmc=1618886 |doi=10.1098/rsbl.2005.0424}} [655] => [656] => {{cite journal |vauthors=Abe K, Gomi K, Hasegawa F, Machida M |s2cid=36874528 |title=Impact of ''Aspergillus oryzae'' genomics on industrial production of metabolites |journal=[[Mycopathologia]] 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and [[Alison Pouliot]], ''Meetings with Remarkable Mushrooms: Forays with Fungi Across Hemispheres'', University of Chicago Press, 278 pp.), ''[[The New York Review of Books]]'', vol. LXX, no.14 (21 September 2023), pp. 41–42. "Fungi sicken us and fungi sustain us. In either case, we ignore them at our peril." (p. 42.) [1078] => [1079] => == External links == [1080] => {{Sister project links|c=Fungi |commonscat=Fungi |wikt=fungus |1= |author= |auto= |b=no |collapsible= |cookbook= |d= |display= |iw= |m= |mw= |n=no |position= |q=no |qid= |s=no |species=Fungi |species_author= |style= |v=no |voy=no}} [1081] => {{Library resources box [1082] => |onlinebooks=yes [1083] => |by=no [1084] => |lcheading= Fungi [1085] => |label=Fungus [1086] => }} [1087] => * [http://tolweb.org/fungi Tree of Life web project: Fungi] [1088] => * [https://www.eol.org/pages/5559 Encyclopedia of Life: Fungus] [1089] => * [http://www.botanical-dermatology-database.info/BotDermFolder/FUNGI.html FUNGI] in [http://www.botanical-dermatology-database.info/index.html BoDD – Botanical Dermatology Database] [1090] => [1091] => {{Fungi classification}} [1092] => {{Fungus structure}} [1093] => {{Eukaryota}} [1094] => {{Nature nav}} [1095] => {{Life on Earth}} [1096] => {{Organisms et al.}} [1097] => {{Taxonbar|from=Q764}} [1098] => {{Authority control}} [1099] => [1100] => [[Category:Fungi| ]] [1101] => [[Category:Articles containing video clips]] [1102] => [[Category:Cryptogams]] [1103] => [[Category:Extant Early Devonian first appearances]] [1104] => [[Category:Kingdoms (biology)|Fungi]] [] => )
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Fungus

Fungi are a diverse group of organisms that belong to the kingdom Fungi. They are characterized by their ability to break down organic matter and absorb nutrients from their surroundings.

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They are characterized by their ability to break down organic matter and absorb nutrients from their surroundings. This process, known as decomposition, is crucial for the recycling of nutrients in ecosystems. Fungi can be found in various environments, including forests, soils, and on decaying matter. They play important ecological roles, such as symbiotic relationships with plants and other organisms, as well as the decomposition of dead plants and animals. Some fungi are also capable of causing diseases in humans, plants, and animals. The study of fungi, known as mycology, has revealed a vast array of fungal species, estimated to be over 5 million worldwide. These species vary in shape, size, and function. Some fungi, like mushrooms, are visible to the naked eye, while others, like yeast, can only be seen under a microscope. Fungi have a unique life cycle that involves the production of spores, which are akin to seeds in plants. These spores are dispersed into the environment and germinate under favorable conditions, giving rise to new fungal colonies. Fungi can reproduce both sexually and asexually, depending on the species. Humans have been utilizing fungi for various purposes for thousands of years. They have been used for culinary purposes, such as in the production of bread, cheese, and beer. Fungi also have industrial applications, such as the production of antibiotics, enzymes, and biofuels. Additionally, fungi have provided important compounds for the development of medicines, including the famous antibiotic penicillin. Despite their numerous benefits, fungi can also have negative impacts. They can cause damage to crops, leading to significant economic losses in agriculture. Fungal infections in humans, known as mycoses, can be difficult to treat and can have severe consequences, particularly for immunocompromised individuals. Understanding and studying fungi is therefore crucial for various fields, including agriculture, medicine, and ecology. Ongoing research continues to expand knowledge about the diversity, biology, and potential applications of fungi, as well as finding solutions for managing fungal diseases.

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