Trichoderma

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Trichoderma
Trichoderma harzianum.jpg
T. harzianum
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
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 of fungi in the family Hypocreaceae that is present in all soils, where they are the most prevalent 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 specieshave been sequenced and are publicly available from the JGI. [3]

Contents

Taxonomy

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]

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 colony in nature Trichoderma colony in nature.jpg
Trichoderma colony in nature

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 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. [11] A strain of Trichoderma viride is a known cause of dieback of Pinus nigra seedlings. [12]

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. [13] [14] [15]

Medical uses

Cyclosporine A (CsA), a calcineurin inhibitor produced by the fungi Trichoderma polysporum , [16] Tolypocladium inflatum , and Cylindrocarpon lucidum , is an immunosuppressant prescribed in organ transplants to prevent rejection. [17]

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.

See also

Related Research Articles

<span class="mw-page-title-main">Ascomycota</span> Division or phylum of fungi

Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The defining feature of this fungal group is the "ascus", a microscopic sexual structure in which nonmotile spores, called ascospores, are formed. However, some species of Ascomycota are asexual and thus do not form asci or ascospores. Familiar examples of sac fungi include morels, truffles, brewers' and bakers' yeast, dead man's fingers, and cup fungi. The fungal symbionts in the majority of lichens such as Cladonia belong to the Ascomycota.

<i>Botrytis cinerea</i> Species of fungus

Botrytis cinerea is a necrotrophic fungus that affects many plant species, although its most notable hosts may be wine grapes. In viticulture, it is commonly known as "botrytis bunch rot"; in horticulture, it is usually called "grey mould" or "gray mold".

<span class="mw-page-title-main">Conidium</span> Asexual, non-motile spore of a fungus

A conidium, sometimes termed an asexual chlamydospore or chlamydoconidium, is an asexual, non-motile spore of a fungus. The word conidium comes from the Ancient Greek word for dust, κόνις (kónis). They are also called mitospores due to the way they are generated through the cellular process of mitosis. They are produced exogenously. The two new haploid cells are genetically identical to the haploid parent, and can develop into new organisms if conditions are favorable, and serve in biological dispersal.

Tolypocladium inflatum is an ascomycete fungus originally isolated from a Norwegian soil sample that, under certain conditions, produces the immunosuppressant drug ciclosporin. In its sexual stage (teleomorph) it is a parasite on scarab beetles. It forms a small, compound ascocarp that arises from the cadaver of its host beetle. In its asexual stage (anamorph) it is a white mold that grows on soil. It is much more commonly found in its asexual stage and this is the stage that was originally given the name Tolypocladium inflatum.

<i>Trichoderma viride</i> Species of fungus

Trichoderma viride is a fungus and a biofungicide.

<i>Penicillium expansum</i> Species of fungus

Penicillium expansum is a psychrophilic blue mold that is common throughout the world in soil. It causes Blue Mold of apples, one of the most prevalent and economically damaging post-harvest diseases of apples.

<i>Ulocladium</i> Genus of fungi

Ulocladium is a genus of fungi. Species of this genus contain both plant pathogens and food spoilage agents. Other species contain enzymes that are biological control agents. Some members of the genus can invade homes and are a sign of moisture because the mold requires water to thrive. They can cause plant diseases or hay fever and more serious infections in immuno-suppressed individuals.

<i>Podostroma cornu-damae</i> Species of fungus

Podostroma cornu-damae, also known as the poison fire coral, is a species of fungus in the family Hypocreaceae. The fruit bodies of the fungus are highly toxic, and have been responsible for several fatalities in Japan. The fungus contains several trichothecene mycotoxins.

<i>Paecilomyces variotii</i> Species of fungus

Paecilomyces variotii, also known by the name Byssochlamys spectabilis for the sexual state, is a common environmental mold from the Phylum Ascomycota. It is widespread in the environment and can be found in composts, soils and wood, as well es a common environmental contaminant in indoor air and carpet dust. Ascospores of the sexual state of P. variotii are strongly heat-resistant. As such the fungus is a common contaminant of heat-treated foods and juices. Paecilomyces variotii has been associated with a number of infective diseases of humans and animals.

<i>Gliocladium</i> Genus of fungi

Gliocladium is an asexual fungal genus in the Hypocreaceae. Certain other species including Gliocladium virens were recently transferred to the genus Trichoderma and G. roseum became Clonostachys rosea f. rosea in the Bionectriaceae. Gliocladium is a mitosporic, filamentous fungus. Species of Gliocladium rarely produce a sexual state. Most pathogenic, disease-causing fungi in humans are mitosporic like Gliocladium. Gliocladium is filamentous; it grows tubular, elongated, and thread-like. It can be considered a contaminant.

Trichoderma longibrachiatum is a fungus in the genus Trichoderma. In addition to being a distinct species, T. longibrachiatum also typifies one of several clades within Trichoderma which comprises 21 different species. Trichoderma longibrachiatum is a soil fungus which is found all over the world but mainly in warmer climates. Many species from this clade have been adopted in various industries because of their ability to secrete large amounts of protein and metabolites.

<i>Penicillium digitatum</i> Species of fungus

Penicillium digitatum is a mesophilic fungus found in the soil of citrus-producing areas. It is a major source of post-harvest decay in fruits and is responsible for the widespread post-harvest disease in Citrus fruit known as green rot or green mould. In nature, this necrotrophic wound pathogen grows in filaments and reproduces asexually through the production of conidiophores and conidia. However, P. digitatum can also be cultivated in the laboratory setting. Alongside its pathogenic life cycle, P. digitatum is also involved in other human, animal and plant interactions and is currently being used in the production of immunologically based mycological detection assays for the food industry.

Paecilomyces marquandii is a soil-borne filamentous fungus distributed throughout temperate to tropical latitudes worldwide including forest, grassland, sewage sludge and strongly metal polluted area characterized by high tolerance in heavy metals. Simultaneous toxic action of zinc and alachlor result an increase in uptake of metal in this fungus but disrupts the cell membrane. Paecilomyces marquandii is known to parasitize the mushroom, Cuphophyllus virgineus, in the family, Hygrophoraceae. Paecilomyces marquandii is categorised as a biosafety risk group 1 in Canada and is not thought to be a significant pathogen of humans or animals.

Trichoderma asperellum Samuels, Lieckf & Nirenberg is a species of fungus in the family Hypocreaceae. It can be distinguished from T. viride by molecular and phenotypic characteristics. The most important molecular characteristics are divergent ITS-1 and 28S sequences and RFLP's of the endochitinase gene. Main phenotypic characters are conidial ornamentation and arrangement and branching of the conidiophores.

<i>Penicillium spinulosum</i> Species of fungus

Penicillium spinulosum is a non-branched, fast-growing fungus with a swelling at the terminal of the stipe (vesiculate) in the genus Penicillium. P. spinulosum is able to grow and reproduce in environment with low temperature and low water availability, and is known to be acidotolerant. P. spinulosum is ubiquitously distributed, and can often be isolated from soil. Each individual strain of P. spinulosum differs from others in their colony morphology, including colony texture, amount of sporulation and roughness of conidia and conidiophores.

Sarocladium kiliense is a saprobic fungus that is occasionally encountered as a opportunistic pathogen of humans, particularly immunocompromised and individuals. The fungus is frequently found in soil and has been linked with skin and systemic infections. This species is also known to cause disease in the green alga, Cladophora glomerata as well as various fruit and vegetable crops grown in warmer climates.

Aspergillus giganteus is a species of fungus in the genus Aspergillus that grows as a mold. It was first described in 1901 by Wehmer, and is one of six Aspergillus species from the Clavati section of the subgenus Fumigati. Its closest taxonomic relatives are Aspergillus rhizopodus and Aspergillus longivescia.

<i>Myxotrichum chartarum</i> Species of fungus

Myxotrichum chartarum is a psychrophilic and cellulolytic fungus first discovered in Germany by Gustav Kunze in 1823. Its classification has changed many times over its history to better reflect the information available at the time. Currently, M. chartarum is known to be an ascomycete surrounded by a gymnothecium composed of ornate spines and releases asexual ascospores. The presence of cellulolytic processes are common in fungi within the family Myxotrichaceae. M. chartarum is one of many Myxotrichum species known to degrade paper and paper products. Evidence of M. chartarum "red spot" mold formation, especially on old books, can be found globally. As a result, this fungal species and other cellulolytic molds are endangering old works of art and books. Currently, there is no evidence that suggests that species within the family Myxotrichaceae are pathogenic.

<i>Trichoderma koningii</i> Species of fungus

Trichoderma koningii is a very common soil dwelling saprotroph with a worldwide distribution. It has been heavily exploited for agricultural use as an effective biopesticide, having been frequently cited as an alternative biological control agent in the regulation of fungi-induced plant diseases. They are endosymbionts associated with plant root tissues, exhibiting mycoparasitism and promoting plant growth due to their capacity to produce different secondary metabolites.

<i>Trichoderma atroviride</i> Species of fungus

Trichoderma atroviride is a filamentous fungal species commonly found in the soil. This fungal species is of particular interest to researchers due to the plethora of secondary metabolites it makes which are used in industry The genus Trichoderma is known for its ubiquity in almost all soils and being easy to culture. Many Trichoderma's are also avirulent plant symbionts.

References

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Bibliography

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