# Trichoderma

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Genus of fungi

Trichoderma T. harzianum Scientific classification Kingdom: Fungi Division: Ascomycota Class: Sordariomycetes Order: Hypocreales Family: Hypocreaceae Genus: Trichoderma Pers. (1801) Type species Trichoderma fuliginoides Pers. (1801) Species see List of Trichoderma species

***Trichoderma*** is a [genus](/source/Genus) of [fungi](/source/Fungi) in the family [Hypocreaceae](/source/Hypocreaceae) that is present in all soils, where they are the most prevalent [culturable](/source/Cell_culture) fungi. Many species in this genus can be characterized as opportunistic avirulent plant [symbionts](/source/Symbiont).[1] This refers to the ability of several *Trichoderma*species to form mutualistic [endophytic](/source/Endophytic) relationships with several plant species.[2] *Trichoderma* species are also responsible for green mold disease in mushroom cultivation.[3] The genomes of several *Trichoderma* specieshave been sequenced and are publicly available from the [JGI](/source/Joint_Genome_Institute).[4]

## Taxonomy

The genus was described by [Christiaan Hendrik Persoon](/source/Christiaan_Hendrik_Persoon) in 1794, but the taxonomy has remained difficult to resolve. For a long time, it was considered to consist of only one species, *[Trichoderma viride](/source/Trichoderma_viride)*, named for producing green [mold](/source/Mold_(fungus)).[5]

### Subdivision

In 1991, Bissett divided the genus into five [sections](/source/Section_(botany)), partly based on the aggregate species described by Rifai:[6]

- *Pachybasium* (20 species)

- *Longibrachiatum* (21 species)

- *Trichoderma*

- *Saturnisporum* (2 species)

- *Hypocreanum*

With the advent of molecular markers from 1995 onwards, Bissett's scheme was largely confirmed but *Saturnisporum* was merged with *Longibrachiatum*. While *Longibrachiatum* and *Hypocreanum* appeared [monophyletic](/source/Monophyletic), *Pachybasium* was determined to be [paraphyletic](/source/Paraphyletic), many of its species clustering with *Trichoderma*. Druzhina and Kubicek (2005) confirmed the genus as [circumscribed](/source/Circumscription_(taxonomy)) was [holomorphic](/source/Teleomorph%2C_anamorph_and_holomorph). They identified 88 species which they demonstrated could be assigned to two major [clades](/source/Clades).[5] Consequently, the formal description of sections has been largely replaced by informal descriptions of clades, such as the Aureoviride clade or the Gelatinosum clade.

### Species

Main article: [List of Trichoderma species](/source/List_of_Trichoderma_species)

The belief that *Trichoderma* was [monotypic](/source/Monotypic) persisted until the 1969 work of Rifai, who recognised nine species.[7] There are currently 89 accepted species in the genus *Trichoderma*. *[Hypocrea](/source/Hypocrea)* are [teleomorphs](/source/Teleomorph) of *Trichoderma*, with *Trichoderma* being the [anamorphs](/source/Anamorphs) of *Hypocrea*.[7]

## Characteristics

Cultures are typically fast-growing at 25–30 °C (77–86 °F), but some species of *Trichoderma* will grow at 45 °C (113 °F). Colonies are transparent at first on media such as cornmeal dextrose agar (CMD) or white on richer media such as potato dextrose agar (PDA). Mycelium are not typically obvious on CMD, [conidia](/source/Conidia) typically form within one week in compact or loose tufts in shades of green or yellow or less frequently white. A yellow pigment may be secreted into the agar, especially on PDA. Some species produce a characteristic sweet or 'coconut' odor.

Conidiophores are highly branched and thus difficult to define or measure, loosely or compactly tufted, often formed in distinct concentric rings or borne along the scant aerial hyphae. Main branches of the conidiophores produce lateral side branches that may be paired or not, the longest branches distant from the tip and often phialides arising directly from the main axis near the tip. The branches may rebranch, with the secondary branches often paired and longest secondary branches being closest to the main axis. All primary and secondary branches arise at or near 90° with respect to the main axis. The typical *Trichoderma* conidiophore with paired branches assumes a pyramidal aspect. Typically the conidiophore terminates in one or a few phialides. In some species (e.g., *[T. polysporum](https://en.wikipedia.org/w/index.php?title=Trichoderma_polysporum&action=edit&redlink=1)*) the main branches are terminated by long, simple or branched, hooked, straight or sinuous, septate, thin-walled, sterile or terminally fertile elongations. The main axis may be the same width as the base of the phialide or it may be much wider.

[Phialides](/source/Phialide) are typically enlarged in the middle but may be cylindrical or nearly [subglobose](/source/Subglobose). Phialides may be held in whorls, at an angle of 90° with respect to other members of the whorl, or they may be variously [penicillate](https://en.wikipedia.org/w/index.php?title=Penicillate&action=edit&redlink=1) ([gliocladium](/source/Gliocladium)-like). Phialides may be densely clustered on a wide main axis (e.g., *[T. polysporum](https://en.wikipedia.org/w/index.php?title=Trichoderma_polysporum&action=edit&redlink=1)*, *[T. hamatum](/source/Trichoderma_hamatum)*), or they may be solitary (e.g., *[T. longibrachiatum](/source/Trichoderma_longibrachiatum)*).

[Conidia](/source/Conidia) typically appear dry, but in some species, they may be held in drops of clear green or yellow liquid (e.g., *[T. virens](/source/Trichoderma_virens)*, *[T. flavofuscum](https://en.wikipedia.org/w/index.php?title=Trichoderma_flavofuscum&action=edit&redlink=1)*). Conidia of most species are ellipsoidal, 3–5 x 2–4 [μm](/source/Micrometre) (L/W = > 1.3); globose conidia (L/W < 1.3) are rare. Conidia are typically smooth but tuberculate to finely warted conidia are known in a few species. Conidia appear colorless to green, smooth to rough, and are in moist conidial masses, variable in shape and size, small, 2.8– 4.8 [μm](/source/Micrometre) for common species. Conidiophores branch repeatedly, bearing clusters of phialides terminally in most cases.[8]

[Synanamorphs](/source/Synanamorph) are formed by some species that also have typical *Trichoderma* pustules. Synanamorphs are recognized by their solitary [conidiophores](/source/Conidiophore) that are verticillately branched and that bear conidia in a drop of clear green liquid at the tip of each phialide.

[Chlamydospores](/source/Chlamydospore) may be produced by all species, but not all species produce chlamydospores on CMD at 20 °C within 10 days. Chlamydospores are typically unicellular subglobose and terminate short hyphae; they may also be formed within hyphal cells. Chlamydospores of some species are multicellular (e.g., *[T. stromaticum](/source/Trichoderma_stromaticum)*).

*Trichoderma* genomes appear to be in the 30–40 Mb range, with approximately 12,000 genes being identifiable.

## Teleomorph

[Teleomorphs](/source/Teleomorph) of *Trichoderma* are species of the ascomycete genus *[Hypocrea](/source/Hypocrea)*. These are characterized by the formation of fleshy, stromata in shades of light or dark brown, yellow or orange. Typically the stroma is discoidal to pulvinate and limited in extent but stromata of some species are effused, sometimes covering extensive areas. Stromata of some species (Podostroma) are clavate or turbinate. Perithecia are completely immersed. Ascospores are bicellular but disarticulate at the septum early in development into 16 part-ascospores so that the ascus appears to contain 16 ascospores. Ascospores are hyaline or green and typically spinulose. More than 200 species of *Hypocrea* have been described but few have been grown in pure culture and even fewer have been described in modern terms.

## Occurrence

*Trichoderma* colony in nature

*Trichoderma* species are frequently isolated from forest or agricultural soils at all [latitudes](/source/Latitude). *Hypocrea* species are most frequently found on bark or on decorticated wood but many species grow on bracket fungi (e.g. *[H. pulvinata](https://en.wikipedia.org/w/index.php?title=Hypocrea_pulvinata&action=edit&redlink=1)*), Exidia (*[H. sulphurea](https://en.wikipedia.org/w/index.php?title=Hypocrea_sulphurea&action=edit&redlink=1)*) or bird's nest fungi (*[H. latizonata](https://en.wikipedia.org/w/index.php?title=Hypocrea_latizonata&action=edit&redlink=1)*) or agarics (*[H. avellanea](https://en.wikipedia.org/w/index.php?title=Hypocrea_avellanea&action=edit&redlink=1)*).

## Biocontrol agent

Several strains of *Trichoderma* have been developed as biocontrol agents against fungal diseases of plants.[9] The various mechanisms include [antibiosis](/source/Antibiosis), [parasitism](/source/Parasitism), inducing [host-plant resistance](/source/Plant_defense_against_herbivory) , and [competition](/source/Interspecific_competition). Most biocontrol agents are from the species *[T. asperellum](/source/Trichoderma_asperellum)*, *[T. harzianum](/source/Trichoderma_harzianum)*, *[T. viride](/source/Trichoderma_viride)*, and *[T. hamatum](/source/Trichoderma_hamatum)*. The biocontrol agent generally grows in its natural habitat on the root surface, and so affects [root disease](/source/Soil_borne_pathogen) in particular, but can also be effective against [foliar diseases](/source/Plant_pathology).

## Causal agent of disease

Many species of *Trichoderma* are major pathogens of cultivated mushrooms, with infections being referred to as "green mold." *Trichoderma* species have been found to infect [button mushrooms](/source/Button_mushrooms), [shiitake](/source/Shiitake), [oyster mushrooms](/source/Oyster_mushrooms), among others. *Trichoderma* infections have caused major crop losses and widespread epidemics in mushroom growing regions, with estimates of damages caused by infections totaling to tens of millions of dollars. Infected substrate leads to lower yields due to competition, and also causes mushrooms to be malformed, discolored, lesioned, and of lower mass than is typical.[3] The species *[T. aggressivum](https://en.wikipedia.org/w/index.php?title=Trichoderma_aggressivum&action=edit&redlink=1)* (formerly *T. harzianum* biotype 4) has been found to infect button mushrooms.[10][11]

*Trichoderma* spp. can also be pathogenic to plants. *[Trichoderma viride](/source/Trichoderma_viride)* is the causal agent of green mold rot of onion.[12] A strain of *Trichoderma viride* is a known cause of dieback of *[Pinus nigra](/source/Pinus_nigra)* seedlings.[13]

### Toxic house mold

The common [house mold](/source/Mold_health_issues), *[Trichoderma longibrachiatum](/source/Trichoderma_longibrachiatum)*, produces small toxic peptides containing amino acids not found in common proteins, like **[alpha-aminoisobutyric acid](/source/Alpha-aminoisobutyric_acid)**, called **trilongins** (up to 10% w/w). Their toxicity is due to absorption into human cells and production of nano-channels that obstruct vital [ion channels](/source/Ion_channels) that ferry potassium and sodium ions across the [cell membrane](/source/Cell_membrane). This affects in the cells [action potential](/source/Action_potential) profile, as seen in [cardiomyocytes](/source/Cardiomyocytes), [pneumocytes](/source/Pneumocytes) and [neurons](/source/Neurons) leading to conduction defects. Trilongins are highly resistant to heat and [antimicrobials](/source/Antimicrobials) making [primary prevention](/source/Primary_prevention) the only management option.[14][15][16]

## Medical uses

[Cyclosporine A](/source/Cyclosporine_A) (CsA), a [calcineurin](/source/Calcineurin) inhibitor produced by the fungi *[Trichoderma polysporum](https://en.wikipedia.org/w/index.php?title=Trichoderma_polysporum&action=edit&redlink=1)*,[17] *[Tolypocladium inflatum](/source/Tolypocladium_inflatum)*, and *[Cylindrocarpon lucidum](https://en.wikipedia.org/w/index.php?title=Cylindrocarpon_lucidum&action=edit&redlink=1)*, is an immunosuppressant prescribed in organ transplants to prevent rejection.[18]

## Industrial use

*Trichoderma*, being a saprophyte adapted to thrive in diverse situations, produces a wide array of enzymes. By selecting strains that produce a particular kind of enzyme, and culturing these in suspension, industrial quantities of enzyme can be produced.

- *[Trichoderma reesei](/source/Trichoderma_reesei)* is used to produce [cellulase](/source/Cellulase) and hemicellulase.[19]

- *[Trichoderma longibrachiatum](/source/Trichoderma_longibrachiatum)* is used to produce [xylanase](/source/Xylanase).[20]

- *[Trichoderma harzianum](/source/Trichoderma_harzianum)* is used to produce [chitinase](/source/Chitinase).[21]

## See also

- [List of *Trichoderma* species](/source/List_of_Trichoderma_species)

- [Bisvertinolone](/source/Bisvertinolone)

## References

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1. **[^](#cite_ref-6)** Bissett, John (November 1991). "A revision of the genus *Trichoderma*. III. Section *Pachybasium*". *Canadian Journal of Botany*. **69** (11): 2373–2417. [Bibcode](/source/Bibcode_(identifier)):[1991CaJB...69.2373B](https://ui.adsabs.harvard.edu/abs/1991CaJB...69.2373B). [doi](/source/Doi_(identifier)):[10.1139/b91-298](https://doi.org/10.1139%2Fb91-298).

1. ^ [***a***](#cite_ref-Samuels2006_7-0) [***b***](#cite_ref-Samuels2006_7-1) Samuels, Gary J. (2006). ["Trichoderma: Systematics, the Sexual State, and Ecology"](https://zenodo.org/record/1235933). *Phytopathology*. **96** (2): 195–206. [Bibcode](/source/Bibcode_(identifier)):[2006PhPat..96..195S](https://ui.adsabs.harvard.edu/abs/2006PhPat..96..195S). [doi](/source/Doi_(identifier)):[10.1094/PHYTO-96-0195](https://doi.org/10.1094%2FPHYTO-96-0195). [ISSN](/source/ISSN_(identifier)) [0031-949X](https://search.worldcat.org/issn/0031-949X). [PMID](/source/PMID_(identifier)) [18943925](https://pubmed.ncbi.nlm.nih.gov/18943925).

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1. **[^](#cite_ref-10)** Beyer, W.M.; Wuest, P.J.; Anderson, M.G. ["Green mold of Mushrooms"](http://www.ppath.cas.psu.edu/MushGrowInfo/Trichoderma%20Green%20Mold.html). Pennsylvania State University. Retrieved 2007-08-02. Pennsylvania State University extension bulletin

1. **[^](#cite_ref-11)** Samuels, G.J.; Dodd, S.L.; Gams, W.; Castlebury, L.A.; Petrini, O. (2002). ["Trichoderma species associated with the green mold epidemic of commercially grown Agaricus bisporus"](http://www.mycologia.org/cgi/content/full/94/1/146). *Mycologia*. **94** (1): 146–170. [doi](/source/Doi_(identifier)):[10.2307/3761854](https://doi.org/10.2307%2F3761854). [JSTOR](/source/JSTOR_(identifier)) [3761854](https://www.jstor.org/stable/3761854). [PMID](/source/PMID_(identifier)) [21156486](https://pubmed.ncbi.nlm.nih.gov/21156486).

1. **[^](#cite_ref-12)** Sandle, T. (2014-01-01), ["Trichoderma"](https://www.sciencedirect.com/science/article/pii/B9780123847300003372), in Batt, Carl A.; Tortorello, Mary Lou (eds.), *Encyclopedia of Food Microbiology (Second Edition)*, Oxford: Academic Press, pp. 644–646, [doi](/source/Doi_(identifier)):[10.1016/b978-0-12-384730-0.00337-2](https://doi.org/10.1016%2Fb978-0-12-384730-0.00337-2), [ISBN](/source/ISBN_(identifier)) [978-0-12-384733-1](https://en.wikipedia.org/wiki/Special:BookSources/978-0-12-384733-1), retrieved 2024-02-27

1. **[^](#cite_ref-13)** Li Destri Nicosia, M.G.; Mosca, S.; Mercurio, R.; Schena, L. (2015). ["Dieback of Pinus nigra Seedlings Caused by a Strain of Trichoderma viride"](https://doi.org/10.1094%2FPDIS-04-14-0433-RE). *Plant Disease*. **99** (1): 44–49. [Bibcode](/source/Bibcode_(identifier)):[2015PlDis..99...44L](https://ui.adsabs.harvard.edu/abs/2015PlDis..99...44L). [doi](/source/Doi_(identifier)):[10.1094/PDIS-04-14-0433-RE](https://doi.org/10.1094%2FPDIS-04-14-0433-RE). [PMID](/source/PMID_(identifier)) [30699733](https://pubmed.ncbi.nlm.nih.gov/30699733).

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## Bibliography

- Rifai, M. A. 1969. A revision of the genus *Trichoderma*. Mycol. Pap. 116:1-56.

- Druzhinina, Irina; Kubicek, Christian P. (February 2005). ["Species concepts and biodiversity in *Trichoderma* and *Hypocrea* : from aggregate species to species clusters?"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1389624). *Journal of Zhejiang University Science*. **6B** (2): 100–112. [doi](/source/Doi_(identifier)):[10.1631/jzus.2005.B0100](https://doi.org/10.1631%2Fjzus.2005.B0100). [PMC](/source/PMC_(identifier)) [1389624](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1389624). [PMID](/source/PMID_(identifier)) [15633245](https://pubmed.ncbi.nlm.nih.gov/15633245).

This article incorporates [public domain material](/source/Copyright_status_of_works_by_the_federal_government_of_the_United_States) from websites or documents of the [United States Department of Agriculture](/source/United_States_Department_of_Agriculture).

## External links

- Data related to [*Trichoderma*](https://species.wikimedia.org/wiki/Trichoderma) at Wikispecies

- Media related to [*Trichoderma*](https://commons.wikimedia.org/wiki/Category:Trichoderma) at Wikimedia Commons

- Samuels, G.J.; Chaverri, P.; Farr, D.F.; McCray, E.B. ["*Trichoderma* Online"](https://web.archive.org/web/20100420011149/http://nt.ars-grin.gov/taxadescriptions/keys/TrichodermaIndex.cfm). Systematic Botany & Mycology Laboratory, ARS, USDA. Archived from the original on 2010-04-20.{{[cite web](https://en.wikipedia.org/wiki/Template:Cite_web)}}: CS1 maint: bot: original URL status unknown ([link](https://en.wikipedia.org/wiki/Category:CS1_maint:_bot:_original_URL_status_unknown))

- [International Subcommission on Trichoderma and Hypocrea Taxonomy site](http://www.isth.info/).

Taxon identifiers Trichoderma Wikidata: Q135322 Wikispecies: Trichoderma AusFungi: 60014672 CoL: 63W3K EoL: 21368 EPPO: 1TRCDG FloraBase: 50410 GBIF: 2542714 iNaturalist: 352744 IndexFungorum: 10282 IRMNG: 1307461 ITIS: 926486 MycoBank: 10282 NBN: BMSSYS0000018868 NCBI: 5543 NZOR: 17f9a813-6942-4b03-9b47-f3d803444996 SpeciesFungorum: 10282 WoRMS: 100248

Authority control databases: National Czech Republic Israel

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Adapted from the Wikipedia article [Trichoderma](https://en.wikipedia.org/wiki/Trichoderma) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Trichoderma?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.