Fungi Imperfecti
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The fungi imperfecti or imperfect fungi are
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
which do not fit into the commonly established taxonomic classifications of fungi that are based on
biological species concept A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of classification and ...
s or morphological characteristics of sexual structures because their sexual form of
reproduction Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. There are two forms of reproduction: Asexual reproduction, asexual and Sexual ...
has never been observed. They are known as imperfect fungi because only their asexual and vegetative phases are known. They have asexual form of reproduction, meaning that these fungi produce their spores asexually, in the process called sporogenesis. There are about 25,000 species that have been classified in the phylum Deuteromycota and many are Basidiomycota or Ascomycota anamorphs. Fungi producing the antibiotic
penicillin Penicillins (P, PCN or PEN) are a group of beta-lactam antibiotic, β-lactam antibiotics originally obtained from ''Penicillium'' Mold (fungus), moulds, principally ''Penicillium chrysogenum, P. chrysogenum'' and ''Penicillium rubens, P. ru ...
and those that cause
athlete's foot Athlete's foot, known medically as ''tinea pedis'', is a common skin infection of the feet caused by a fungus. Signs and symptoms often include itching, scaling, cracking and redness. In rare cases the skin may blister. Athlete's foot fungus ...
and
yeast infection Candidiasis is a fungal infection due to any species of the genus '' Candida'' (a yeast). When it affects the mouth, in some countries it is commonly called thrush. Signs and symptoms include white patches on the tongue or other areas of the ...
s are algal fungi. In addition, there are a number of edible imperfect fungi, including the ones that provide the distinctive characteristics of
Roquefort Roquefort () is a sheep milk blue cheese from southern France. Though similar cheeses are produced elsewhere, European Union law, EU law dictates that only those cheeses aged in the natural Combalou caves of Roquefort-sur-Soulzon may bear the na ...
and Camembert cheese. Other, more informal names besides phylum Deuteromycota (or class "Deuteromycetes") and fungi imperfecti are anamorphic fungi, or mitosporic fungi, but these are terms without taxonomic rank. Examples are Alternaria, Colletotrichum, Trichoderma etc. The class
Phycomycetes Phycomycetes or algal fungi is an obsolete polyphyletic taxon for certain fungi with aseptate hyphae. It is used in the Engler system. Asexual reproduction takes place by zoospores (motile) or by Aplanospores (non-motile). These spores are endo ...
("algal fungi") has also been used.


Problems in taxonomic classification

Although Fungi imperfecti/Deuteromycota is no longer formally accepted as a
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
, many of the fungi it included have yet to find a place in modern fungal classification. This is because most
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
are classified based on characteristics of the fruiting bodies and spores produced during sexual reproduction, and members of the Deuteromycota have been observed to reproduce only asexually or produce no spores. Mycologists formerly used a unique dual system of nomenclature in classifying fungi, which was permitted by Article 59 of the
International Code of Botanical Nomenclature The ''International Code of Nomenclature for algae, fungi, and plants'' (ICN or ICNafp) is the set of rules and recommendations dealing with the formal botanical names that are given to plants, fungi and a few other groups of organisms, all tho ...
(the rules governing the naming of plants and fungi). However, the system of dual nomenclature for fungi was abolished in the 2011 update of the Code. Under the former system, a name for an asexually reproducing fungus was considered a ''form taxon''. For example, the ubiquitous and industrially important mold, ''
Aspergillus niger ''Aspergillus niger'' is a mold classified within the ''Nigri'' section of the ''Aspergillus'' genus. The ''Aspergillus'' genus consists of common molds found throughout the environment within soil and water, on vegetation, in fecal matter, on de ...
'', has no known sexual cycle. Thus ''Aspergillus niger'' was considered a form taxon. In contrast, isolates of its close relative, '' Aspergillus nidulans'' , revealed it to be the anamorphic stage of a teleomorph (the ascocarp or fruiting body of the sexual reproductive stage of a fungus), which was already named ''Emericella nidulans''. When such a teleomorphic stage became known, that name would take priority over the name of an anamorph (which lacks a sexual reproductive stage). Hence the formerly classified ''Aspergillus nidulans'' would be properly called ''Emericella nidulans'' – note there's no reference to the original author. The system since 2013 instead treats both as the same species typified by the anamorph, and hence the author citation would include the original author as ''Emericella nidulans''


Phylogeny and taxonomy

Phylogenetic classification of asexually reproducing fungi now commonly uses molecular systematics.
Phylogenetic tree A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA. In ...
s constructed from comparative analyses of
DNA sequence A nucleic acid sequence is a succession of bases within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the nu ...
s (such as
rRNA Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosomal ...
or multigene phylogenies )may be used to infer relationships between asexually reproducing fungi and their sexually reproducing counterparts. With these methods, many asexually reproducing fungi have now been placed in the tree of life. However, because phylogenetic methods require sufficient quantities of biological materials (spores or fresh specimens) that are from pure (i.e., uncontaminated) fungal cultures, for many asexual species their exact relationship with other fungal species has yet to be determined. Under the current system of fungal nomenclature, teleomorph names cannot be applied to fungi that lack sexual structures. Classifying and naming asexually reproducing fungi is the subject of ongoing debate in the mycological community.


Historical classification of the imperfect fungi

These groups are no longer formally accepted because they do not adhere to the principle of
monophyly In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent comm ...
. The taxon names are sometimes used informally. In particular, the term 'hyphomycetes' is often used to refer to molds, and the term 'coelomycetes' is used to refer to many asexually reproducing plant pathogens that form discrete fruiting bodies. Following, a classification of the Fungi imperfecti: Saccardo et al.(1882-1972) * Class Hyphomycetes lacking fruiting bodies ** Order Moniliales (producing spores on simple
conidiophore A conidium ( ; : conidia), sometimes termed an asexual chlamydospore or chlamydoconidium (: chlamydoconidia), is an Asexual reproduction, asexual, non-motility, motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word f ...
s) ** Order Stilbellales (producing spores on synnemata) ** Order Tuberculariales (producing spores in sporodochia) * Class Coelomycetes spores produced in fruiting bodies ** Order Melanconiales (producing spores in acervuli) ** Order Sphaeropsidales (producing spores in pycnidia) * Class Agonomycetes lacking spores Other, according to Dörfelt (1989): * ''Form-Klasse:'' Hyphomycetes ** ''Form-Ordnung:'' Agonomycetales *** ''Form-Familie:'' Agonomycetaceae ** ''Form-Ordnung:'' Moniliales *** ''Form-Familie:'' Moniliaceae *** ''Form-Familie:'' Dematiaceae *** ''Form-Familie:'' Stilbellaceae *** ''Form-Familie:'' Tuberculariaceae * ''Form-Klasse:'' Coelomycetes ** ''Form-Ordnung:'' Melanconiales *** ''Form-Familie:'' Melanconiaceae ** ''Form-Ordnung:'' Sphaeropsidales *** ''Form-Familie:'' Sphaeropsidaceae Other systems of classification are reviewed by .


Common species


Industrially relevant fungi

* '' Tolypocladium inflatum'' → from which the
immunosuppressant Immunosuppressive drugs, also known as immunosuppressive agents, immunosuppressants and antirejection medications, are drugs that inhibit or prevent the activity of the immune system. Classification Immunosuppressive drugs can be classified ...
ciclosporin Ciclosporin, also spelled cyclosporine and cyclosporin, is a calcineurin inhibitor, used as an immunosuppressant medication. It is taken Oral administration, orally or intravenously for rheumatoid arthritis, psoriasis, Crohn's disease, nephr ...
is obtained; * '' Penicillium griseofulvum'' * ''
Penicillium roqueforti ''Penicillium roqueforti'' is a common saprotrophic fungus in the genus ''Penicillium''. Widespread in nature, it can be isolated from soil, decaying organic matter, and plants. The major industrial use of this fungus is the production of blu ...
'' * ''
Penicillium camemberti ''Penicillium camemberti'' is a species of fungus in the genus ''Penicillium''. It is used in the production of Camembert, Brie, Langres, Coulommiers, and Cambozola cheeses, on which colonies of ''P. camemberti'' form a hard, white crust ...
'' * Other species of ''Penicillium'' are used to improve both the taste and the texture of
cheese Cheese is a type of dairy product produced in a range of flavors, textures, and forms by coagulation of the milk protein casein. It comprises proteins and fat from milk (usually the milk of cows, buffalo, goats or sheep). During prod ...
s * ''
Aspergillus oryzae ''Aspergillus oryzae'', also known as , is a mold used in East Asia to saccharify rice, sweet potato, and barley in the making of alcoholic beverages such as '' sake'' and '' shōchū'', and also to ferment soybeans for making soy sauce and '' ...
'' * '' Aspergillus sojae'' * ''
Aspergillus niger ''Aspergillus niger'' is a mold classified within the ''Nigri'' section of the ''Aspergillus'' genus. The ''Aspergillus'' genus consists of common molds found throughout the environment within soil and water, on vegetation, in fecal matter, on de ...
'' * '' Amorphotheca resinae'' * ''Lecanicillium'' sp. → these produce
conidia A conidium ( ; : conidia), sometimes termed an asexual chlamydospore or chlamydoconidium (: chlamydoconidia), is an asexual, non- motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also ...
which may control certain species of insect
pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...
* Other entomopathogenic fungi, including ''
Metarhizium ''Metarhizium'' is a genus (biology), genus of entomopathogenic fungus, entomopathogenic fungi in the family Clavicipitaceae. With the advent of genetic profiling, placing these fungi in proper taxa has now become possible. Most turn out to be ...
'' and '' Beauveria'' spp. * '' Pochonia'' spp. are under development for control of
Nematode The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (h ...
pests.


Important pathogens

* '' Magnaporthe oryzae'', a rice pathogen


Evolution

Many fungi imperfecti are unable to reproduce in a sexual way because of identified mutations that disable their ability to sexually reproduce, others may be sterile due to epigenetic changes. These kinds of changes can arise very quickly in laboratory conditions that force 10–19 generations of exclusive asexual reproduction. Female-sterile strains of '' Magnaporthe oryzae'' appear to have a selective advantage in the form of faster growth with more efficient conidia transfer. Nieuwenhuis and James (2016) also discusses the costs and benefits for sexual and reproduction in fungi as well as mechanisms that have evolved to reduce the costs of either. They also describe the so-called "parasexual cycle" in some fungi imperfecti, which allows recombination without sex.


See also

*
Forest pathology Forest pathology is the research of both biotic and abiotic maladies affecting the health of a forest ecosystem, primarily fungal pathogens and their insect vectors. It is a subfield of forestry and plant pathology. Forest pathology is part ...
* List of mitosporic Ascomycota


References


Bibliography

* * * * * {{Authority control . Fungus common names Fungi by classification