Trichoderma

Trichoderma is a genus of fungi in the family Hypocreaceae that

is present in all soils, where they are the most prevalent Trichoderma
culturable fungi. Many species in this genus can be characterized
as opportunistic avirulent plant symbionts.[1] This refers to the
ability of several Trichoderma species to form mutualistic
endophytic relationships with several plant species.[2] The
genomes of several Trichoderma species have been sequenced
and are publicly available from the JGI.[3]

Contents
Taxonomy
Subdivision
Species
T. harzianum
Characteristics
Scientific classification
Teleomorph
Kingdom: Fungi
Occurrence
Division: Ascomycota
Biocontrol agent
Causal agent of disease Class: Sordariomycetes
Toxic house mold Order: Hypocreales
Medical uses Family: Hypocreaceae
Industrial use
Genus: Trichoderma
See also Pers. (1801)
References
Type species
Bibliography
Trichoderma fuliginoides

External links
Pers. (1801)

Species
Taxonomy
see List of Trichoderma species
The genus was described by 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, named for producing green
mold.[4]

Subdivision
In 1991, Bissett divided the genus into five sections, partly based on the aggregate species described by
Rifai:[5]

Pachybasium (20 species)

Longibrachiatum (10 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, Pachybasium was determined to be paraphyletic, many of its species clustering with
Trichoderma. Druzhina and Kubicek (2005) confirmed the genus as circumscribed was holomorphic. They
identified 88 species which they demonstrated could be assigned to two major clades.[4] 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

The belief that Trichoderma was monotypic persisted until the 1969 work of Rifai, who recognised nine
species.[6]
There are currently 89 accepted species in the genus Trichoderma. Hypocrea are teleomorphs of
Trichoderma, which themselves have Hypocrea as anamorphs.[6]

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
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) 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 are typically enlarged in the middle but may be cylindrical or nearly 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 (gliocladium-like). Phialides may be densely clustered on a wide main axis (e.g., T. polysporum,
T. hamatum), or they may be solitary (e.g., T. longibrachiatum).
Conidia typically appear dry, but in some species, they may be held in drops of clear green or yellow liquid
(e.g., T. virens, T. flavofuscum). Conidia of most species are ellipsoidal, 3–5 x 2–4  µm (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 mm for common species. Conidiophores branch
repeatedly, bearing clusters of phialides terminally in most cases. [7]

Synanamorphs are formed by some species that also have typical Trichoderma pustules. Synanamorphs are
recognized by their solitary conidiophores that are verticillately branched and that bear conidia in a drop of
clear green liquid at the tip of each phialide.

Chlamydospores 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).

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

Teleomorph
Teleomorphs of Trichoderma are species of the ascomycete genus 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 species are frequently isolated from forest or
agricultural soils at all latitudes. Hypocrea species are most
frequently found on bark or on decorticated wood but many species
grow on bracket fungi (e.g. H. pulvinata), Exidia (H. sulphurea) or
bird's nest fungi (H. latizonata) or agarics (H. avellanea).

Biocontrol agent
Several strains of Trichoderma have been developed as biocontrol Trichoderma colony in nature
agents against fungal diseases of plants.[8] The various mechanisms
include antibiosis, parasitism, inducing host-plant resistance
, and
competition. Most biocontrol agents are from the species T. asperellum, T. harzianum, T. viride, and T.
hamatum. The biocontrol agent generally grows in its natural habitat on the root surface, and so affects root
disease in particular, but can also be effective against foliar diseases.

Causal agent of disease

T. aggressivum (formerly T. harzianum biotype 4) is the causal agent of green mold, a disease of cultivated
button mushrooms.[9][10] Trichoderma viride is the causal agent of green mold rot of onion. A strain of
Trichoderma viride is a known cause of dieback of Pinus nigra seedlings.[11]

Toxic house mold

The common house mold, Trichoderma longibrachiatum, produces small toxic peptides containing amino
acids not found in common proteins, like 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 that ferry potassium and sodium ions across the cell membrane. This affects in the cells action
potential profile, as seen in cardiomyocytes, pneumocytes and neurons leading to conduction defects.
Trilongins are highly resistant to heat and antimicrobials making primary prevention the only management
option.[12][13][14]

Medical uses
Cyclosporine A (CsA), a calcineurin inhibitor produced by the fungi Trichoderma polysporum,[15]
Tolypocladium inflatum, and Cylindrocarpon lucidum, is an immunosuppressant prescribed in organ
transplants to prevent rejection.[16]

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 is used to produce cellulase and hemicellulase.[17]

Trichoderma longibrachiatum is used to produce xylanase.[18]
Trichoderma harzianum is used to produce chitinase.[19]

See also
List of Trichoderma species
Bisvertinolone

References
1. Harman, G.E.; Howell, C.R.; Viterbo, A.; Chet, I.; Lorito, M. (2004). "Trichoderma species—
opportunistic avirulent plant symbionts" (https://naldc-legacy.nal.usda.gov/naldc/download.x
html?id=25508&content=PDF). Nature Reviews Microbiology. 2 (1): 43–56.
doi:10.1038/nrmicro797 (https://doi.org/10.1038%2Fnrmicro797). PMID 15035008 (https://pu
bmed.ncbi.nlm.nih.gov/15035008).
2. Bae, H; Roberts, DP; Lim, HS; Strem, MD; Park, SC; Ryu, CM; Melnick, RL; Bailey, BA
(2011). "Endophytic Trichoderma isolates from tropical environments delay disease onset
and induce resistance against Phytophthora capsici in hot pepper using multiple
mechanisms" (https://doi.org/10.1094%2FMPMI-09-10-0221). Mol. Plant Microbe Interact. 24
(3): 336–51. doi:10.1094/MPMI-09-10-0221 (https://doi.org/10.1094%2FMPMI-09-10-0221).
PMID 21091159 (https://pubmed.ncbi.nlm.nih.gov/21091159).
 3. "Home - Trichoderma atroviride" (http://genome.jgi-psf.org/Triat1/Triat1.home.html).
4. Druzhinina, Irina; Kubicek, Christian P. (February 2005). "Species concepts and biodiversity
in Trichoderma and Hypocrea : from aggregate species to species clusters?" (https://www.nc
bi.nlm.nih.gov/pmc/articles/PMC1389624). Journal of Zhejiang University Science. 6B (2):
100–112. doi:10.1631/jzus.2005.B0100 (https://doi.org/10.1631%2Fjzus.2005.B0100).
PMC 1389624 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1389624). PMID 15633245
(https://pubmed.ncbi.nlm.nih.gov/15633245).
5. Bissett, John (November 1991). "A revision of the genus Trichoderma. III. Section
Pachybasium". Canadian Journal of Botany. 69 (11): 2373–2417. doi:10.1139/b91-298 (http
s://doi.org/10.1139%2Fb91-298).
6. Samuels, Gary J. (2006). "Trichoderma: Systematics, the Sexual State, and Ecology" (https://
zenodo.org/record/1235933/files/article.pdf) (PDF). Phytopathology. 96 (2): 195–206.
doi:10.1094/PHYTO-96-0195 (https://doi.org/10.1094%2FPHYTO-96-0195). ISSN 0031-
949X (https://www.worldcat.org/issn/0031-949X). PMID 18943925 (https://pubmed.ncbi.nlm.
nih.gov/18943925).
7. Yang, Chin (2007). SAMPLING AND ANALYSIS OF INDOOR MICROORGANISMS. Wiley.
p. 259. ISBN 9780471730934.
8. Harman, G.E. (2006). "Overview of mechanisms and uses of Trichoderma spp" (https://doi.or
g/10.1094%2FPHYTO-96-0190). Phytopathology. 96 (2): 190–194. doi:10.1094/PHYTO-96-
0190 (https://doi.org/10.1094%2FPHYTO-96-0190). PMID 18943924 (https://pubmed.ncbi.nl
m.nih.gov/18943924).
9. 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
10. 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:10.2307/3761854 (https://doi.org/10.2307%2F3761854). JSTOR 3761854 (https://www.j
stor.org/stable/3761854).
11. 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-04-1
4-0433-RE). Plant Disease. 99 (1): 44–49. doi:10.1094/PDIS-04-14-0433-RE (https://doi.org/
10.1094%2FPDIS-04-14-0433-RE). PMID 30699733 (https://pubmed.ncbi.nlm.nih.gov/3069
9733).
12. Reason Discovered for the Toxicity of Indoor Mould – ScienceDaily (Oct. 12, 2012) :
https://www.sciencedaily.com/releases/2012/10/121012074655.htm
13. Mikkola, Raimo; Andersson, Maria A.; Kredics, László; Grigoriev, Pavel A.; Sundell, Nina;
Salkinoja-Salonen, Mirja S. (2012). "20-Residue and 11-residue peptaibols from the fungus
Trichoderma longibrachiatum are synergistic in forming Na + /K + -permeable channels and
adverse action towards mammalian cells" (https://doi.org/10.1111%2Ffebs.12010). FEBS
Journal. 279 (22): 4172–4190. doi:10.1111/febs.12010 (https://doi.org/10.1111%2Ffebs.120
10). PMID 22994321 (https://pubmed.ncbi.nlm.nih.gov/22994321).
14. “Trilongins” Offer Insight into Mold Toxicity (http://ehp.niehs.nih.gov/121-a44/) Archived (http
s://web.archive.org/web/20160311024320/http://ehp.niehs.nih.gov/121-a44/) 2016-03-11 at
the Wayback Machine Environmental health perspectives 2/2013.
15. Dreyfuss, M; Härri, E; Hofmann, H; Kobel, H; Pache, W; Tscherter, H (1976). "Cyclosporin A
and C: new metabolites from Trichoderma polysporum (Link ex Pers.) Rifai". European
Journal of Applied Microbiology. 3: 125–133. doi:10.1007/bf00928431 (https://doi.org/10.100
7%2Fbf00928431).
16. Authors; Chong, FW; Chakravarthi, S; Nagaraja, HS; Thanikachalam, PM; Lee, N (Jun
2009). "Expression of transforming growth factor-beta and determination of apoptotic index
in histopathological sections for assessment of the effects of Apigenin (4', 5', 7'-
Trihydroxyflavone) on Cyclosporine A induced renal damage". Malays J Pathol. 31 (1): 35–
43. PMID 19694312 (https://pubmed.ncbi.nlm.nih.gov/19694312).
17. "Home - Trichoderma reesei v2.0" (http://genome.jgi-psf.org/Trire2/Trire2.home.html).
18. Azin, M.; Moravej, R.; Zareh, D. (2007). "Self-directing optimization of parameters for
extracellular chitinase production by Trichoderma harzianum in batch mode". Process
Biochemistry. 34 (6–7): 563–566. doi:10.1016/S0032-9592(98)00128-9 (https://doi.org/10.10
16%2FS0032-9592%2898%2900128-9).
19. Felse, P.A.; Panda, T. (1999). "Production of xylanase by Trichoderma longibrachiatum on a
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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.nc
bi.nlm.nih.gov/pmc/articles/PMC1389624). Journal of Zhejiang University Science. 6B (2):
100–112. doi:10.1631/jzus.2005.B0100 (https://doi.org/10.1631%2Fjzus.2005.B0100).
PMC 1389624 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1389624). PMID 15633245
(https://pubmed.ncbi.nlm.nih.gov/15633245).

  This article incorporates  public domain material from websites or documents of the United States
Department of Agriculture.

External links
Data related to Trichoderma at Wikispecies
Media related to Trichoderma at Wikimedia Commons
Samuels, G.J.; Chaverri, P.; Farr, D.F.; McCray, E.B. "Trichoderma Online" (https://web.archiv
e.org/web/20100420011149/http://nt.ars-grin.gov/taxadescriptions/keys/TrichodermaIndex.cf
m). Systematic Botany & Mycology Laboratory, ARS, USDA. Archived from the original on
2010-04-20.
International Subcommission on Trichoderma and Hypocrea Taxonomy site (http://www.isth.i
nfo/).

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