Trichoderma in Agriculture,
Industry and Medicine: An Overview
Prasun K. Mukherjee,1* Benjamin A. Horwitz,2 Uma S. Singh,3
Mala Mukherjee1 and Monika Schmoll4
Central Institute for Cotton Research, Nagpur, India; 2The Technion – Israel
Institute for Technology, Haifa, Israel; 3International Rice Research Institute,
New Delhi, India; 4Austrian Institute of Technology GmbH (AIT), Tulln, Austria



Species of the genus Trichoderma belong to one
of the most useful groups of microbes to have
had an impact on human welfare in recent
times. These filamentous fungi have many
applications (Fig. 1.1). They are the most widely
used biofungicides and plant growth modifiers,
and are sources of enzymes of industrial utility, including those used in the biofuels industry.
Furthermore, they are prolific producers of secondary metabolites, some of which have clinical significance, and some species have been
engineered to act as microbial cell factories for
the heterologous production of important proteins. In the soil, Trichoderma species are used in
the bioremediation of organic and inorganic
wastes including heavy metals (Schuster and
Schmoll, 2010; Harman, 2011a,b).
No doubt, the volume of literature
(Google Scholar database ~138,000 hits) and
the number of patents (Google patents database ~56,100 hits) being filed/granted on
Trichoderma have seen exponential growth in
the past few decades. Centuries ago, this
genus was erected by Persoon (1794); however, the potential in human welfare was
realized only in the 1930s after Weindling

(1932, 1934) demonstrated that these fungi
are mycoparasites, antibiotic producers and
plant disease biocontrol agents. There was little research in the area of biological control
using Trichoderma for the next three decades
but then it picked up again, thanks to the
pioneering and dedicated work of several
research groups who demonstrated the bioefficacy of Trichoderma under field conditions
in the 1970s. Today, these fungi are among
the most widely researched microbes, as is
evident from the huge volume of published
literature. Unfortunately, the taxonomy of
these economically important fungi, which
relied mostly on their morphology, was
rather messy until the advent of molecular
tools and barcodes (http://www.isth.info/).
Even today, many of the strains deposited in
culture collections across the world and many
DNA sequences deposited in the databases
are yet to be corrected (see Chapter 2, this volume). Correct identification (often neglected)
is a must for these economically and clinically
significant fungi. This is especially important
for two reasons: (i) some of the beneficial
traits are species (and often strain) specific and
hence it is important to identify the species/
strain to realize its full potential; and (ii), more

* E-mail: prasunmukherjee1@gmail.com
© CAB International 2013. Trichoderma: Biology and Applications
(eds P.K. Mukherjee et al.)


as es . most of the companies produce Trichoderma on grains and. have had a major impact on agriculture. more than 250 Trichodermabased formulations are sold commercially and many are added each year (Singh et al. .2 Trichoderma in Agriculture Trichoderma spp. yield improvement and control of soil-borne pathogens is striking. 2004. It is unfortunate that even with the advent of freely available online software tools such as TrichoKey. This is one aspect that needs the immediate attention of scientists. green mould of mushrooms A G R I C U LT U R E Fig. Impact (positive and negative) of Trichoderma on human welfare. this volume). One interesting thing about Trichoderma is that it can control fungi belonging to taxonomically diverse groups as well as oomycetes. The adverse side effect of such advancements is the quality control.. improved nutrient use efficiency. Mukherjee et al.. 1. improved seed germination. 2012). breaking of seed dormancy. especially threatening immuno-compromised individuals (see Chapter 17. source of transgenes. me zy En n NT ma ME .. hu nt ON oxins eme IR ot ag yc an NV m /E . biofertilizers. government regulators and the production industry. it is necessary for the safety of the handler and the users (many are illiterate farmers/farm workers). This is in sharp contrast with other countries.info/) many of the widely used commercial strains are not yet correctly identified. 1. Although their interaction with plants is not a strict symbiotic one as with rhizobia and mycorrhiza. where only a handful of products are available. According to a conservative estimate. U e t S na s). this volume). Harman et al. about 60% of all the registered biofungicides worldwide are Trichoderma based (Verma et al.1. which is quite often compromised (use of Trichoderma as biofungicides is discussed in detail in Chapter 14. in industrialized nations. whereas in India and perhaps in other developing nations. In India alone. h extile no et T pa ero and R r tic lo bio Y g les ou fu .b tia ns ten g e Po atho p s he for f mi oo I N ce d. 2007). Selling Trichoderma has thus become a very profitable business with small entrepreneurs. The plant defence induced by Trichoderma helps in restricting pathogenic bacterial growth on foliage (see Chapter 10. whereas others are human pathogens. Some species/strains are producers of mycotoxins. this volume. 2007). 2 Biofungicides. herbicides.isth. llu fee D las d. the set-up needs very small initial investments and the returns are high (Verma et al. TrichoBLAST (http://www. mainly the industrialized nations.P. importantly.K.m TH nce ution AL ll ica HE ignif nd po N a ls MA ica tion HU f clin edia s o em rug ior ld . because the costs of labour are low. This can be explained by the fact that. c s p el ell ro ind fac tei us tor n p trie ies rod s ( uc cel tio lul n.. Trichoderma are produced in large fermenters that need huge investment. abiotic stress management.

2012). it is expected that our understanding of biocontrol in a holistic manner will become more robust in the next few years (Druzhinina et al. addition of glutathione also had similar effects. 2010). Bae et al. the role of individual mechanisms in vivo is not well understood. 2011). The amelioration of oxidative stress has been suggested to be one of the major mechanisms of Trichoderma-mediated mitigation of abiotic stresses. 2010. 2009.. Interestingly.g. 2011. 2009. Biocontrol is a complex mechanism mediated by direct parasitism (hyphae and resting structures/fruiting bodies) of plant pathogens. 2006. especially in developing countries. Mastouri et al.. 2012). salinity and so on. The plants also respond by inducing many stress-responsive genes (Alfano et al.g.. thus opening a new frontier in Trichoderma research (see Chapter 14. Kubicek et al. 2011a.g. 2004. The beneficial effects of Trichoderma genes in plants have been reviewed recently (Hermosa et al. One example worth mentioning is the evaluation of marine isolates of Trichoderma for the biological control of plant diseases in saline and arid soils (Gal-Hemed et al... Distefano et al. 2012a). the endochitinase gene that was originally transferred to plants for the control of plant pathogens was also reported to be imparting abiotic stress tolerance. induction of plant defences. 2008). this volume. The isolation of endophytic Trichoderma strains is an emerging area that is yielding interesting results.. have also been reported to be parasitic on nematodes. Shoresh et al. One of the major issues is the availability of quality products. Shoresh et al. a Trichoderma strain or a combination of strains that are effective against both fungi and nematodes might prove to be very effective in managing these complex diseases. Industry and Medicine Druzhinina et al.. Harman. Trichoderma seed treatment also ameliorates physiological stresses such as ageing and seed dormancy (Delgado-Sanchez et al. These fungi colonize plant roots internally and externally and take up sucrose from roots. Brotman et al. 2010). the expression of a Trichoderma glutathione S-transferase (GST) in tobacco enhanced tolerance to heavy metals and anthracene (a polycyclic aromatic hydrocarbon) through countering the oxidative stresses generated (Dixit et al. high soil pH. e. It has also been proposed that Trichoderma improves plant growth through nutrient solubilization and hence shows better effects under nutrient stress conditions (Mastouri et al.. however. Trichoderma spp. 2012). In addition. 2010). competition for iron and other nutrients. 2011). Shoresh et al.. 2011).. and consumption of germination stimulants from seeds/rhizosphere (Harman et al. Sharon et al.... this volume. owing to the action of release of elicitors (de las Mercedes Dana et al. 2010). Howell. extended shelf-life at ambient temperatures and efficacy under conditions of abiotic stresses (e.b). 2006.Trichoderma in Agriculture. are long known to improve plant (root and shoot) growth (Harman et al. Sustainable growth of the Trichoderma-based biofungicide market will require some modifications. Trichoderma spp.. Apart from biocontrol. Gene function studies with individual gene knockout mutants have set the pace for understanding the mechanisms of biocontrol at the molecular level (for a review see Mukherjee et al. 2004. production of secondary metabolites. Even though it is well researched. 2011). recently shown to be through the production of phytohormones and certain other secondary metabolites (see Chapter 10. Now that we have the genome sequences available for seven species of Trichoderma (with more species scheduled for sequencing already). Indeed. 2006. nematode-assisted penetration of the wilt pathogen Fusarium). 2007. Szabó et al.. 2012. . 2012. 2008. In addition to the control of plant pathogenic fungi and bacteria. Lorito et al.. 2011. 2011). salinity and low moisture).. In addition to imparting tolerance to several abiotic stresses such as drought. 2010. including isolates that can confer biotic and abiotic stress tolerance and produce novel secondary metabolites (see Chapter 9. but simultaneously enhance photosynthetic ability and induced defence (Vargas et al. 3 this volume. Because many plant pathogenic nematodes and fungi ‘collaborate’ in causing plant diseases (e... There is a need for improving the quality of formulations for higher initial colony-forming units (CFUs)... 2010).. there should be more exploratory research to identify new Trichoderma strains with novel applications.... which are otherwise difficult to control. Moran-Diez et al. 2001.

lead to the achievement of mating with other Trichoderma spp. boosted research on the degradation of plant material and nowadays this fungus. T. 1953). Having become one of the most prolific producers of plant cell wall degrading enzymes. because they are mycoparasites. reesei opened up a whole new field of research within the genus Trichoderma. it is now grown in liquid culture in huge steel fermenters. At the same time.. especially for cellulases. reesei becoming a model system for plant cell wall degradation.K. a species specificity seems to be there that restricts the epidemics to only certain species (Samuels et al. this fungus has undergone countless rounds of mutation by chemicals and radiation. reesei as a cell factory for the production of valuable chemical compounds (see Chapter 16. have herbicidal properties too (Javaid and Ali. however. Trichoderma spp. the discovery of sexual development in T.3 Trichoderma in Industry An interesting chance discovery that changed the face of the enzyme industry happened during the Second World War: a filamentous fungus thriving in its tropical home of the Solomon Islands made a crucial mistake. The investigation of secondary metabolism with Trichoderma spp. 2002. .. Hutchinson. 2011). Since then. Another rather novel tool has become available for Trichoderma in recent years: sexual crossing (see Chapter 4. have also negatively impacted agriculture.. 1988. But soon. this volume). sexual crossing will considerably speed up strain improvement and research. the investigation of the possibility of Trichoderma undergoing sexual development will. in the early days. Mukherjee et al. is mainly aimed at the role of bioactive compounds in biocontrol. Komon´-Zelazowska et al. and. many fungi were evaluated for the potential to degrade plant cell wall material along with Trichoderma reesei (Siu and Reese. Jones et al. 2009.4 P. A proof-of-principle study recently revealed the possibility of using T. Interestingly. Besides providing important insights into the physiology of Trichoderma.. a secondary metabolite of Trichoderma virens. Thanks to its strong cellulase promoters and the versatile tools developed for genetic engineering of this fungus (Kubicek et al.. At first. together with other species of the genus. Initially. 2007). 2012). 1999). It took some time. 2011). which happened to be using its habitat... 1. the interest in T. numerous applications. It happily degraded tents and uniforms of the military. Schuster et al. 1984. along with improvement by genetic engineering (Seiboth et al. and. Hatvani et al. These perspectives will be of great importance to all research areas where this genus plays a role – from biocontrol to human pathogenicity. the mushroom fungi could also be targets of parasitism. in the future. instead of the humid and rich tropical soil to which it was adapted. was reported earlier as having herbicidal properties (Howell. however. this volume). probably because of the availability of typical phytopathogens as more effective bioherbicides. 2007. reesei is also used as a host for heterologous protein production. including biofuels (see Chapter 13. because Trichoderma spp. This is one area that has not been explored rigorously. With the advent of the biofuel industry. this volume). are vigorous competitors and hence could easily infest the compost. reesei (then called Trichoderma viride) was not even among the isolates studied for efficient cellulase production. offers a complete toolbox from plant protection to industrial fermentation for the production of enzymes and chemicals. degrading the leftovers of plant growth and keeping the carbon cycle in balance. research on the organisms involved focusing on preventing the damage done by the hydrolytic enzymes of the fungi. Viridiol. This is not unexpected though. Its efficient enzyme system later resulted in T. reesei has been renewed and is being actively investigated to improve its potential to degrade plant-based wastes. T. from discovery to industrial application. with several reports on the ‘green mould’ epidemics that caused havoc in mushroom cultivation. The interest in this field has been renewed recently with the demonstration that the metabolites from some other Trichoderma spp. Although currently a major challenge.

can become a valuable source of new antibiotics. 2012). virens. Gliotoxin. but at the same time is produced by the human pathogen Aspergillus fumigatus and is branded as a ‘mycotoxin’ (Scharf et al.. With the advent of novel challenges in infectious diseases. The availability of genome sequences is expected to yield more novel secondary metabolites with novel functions (Mukherjee et al.. this volume). Fayaz et al. 2012c). Metabolites of Trichoderma spp. reesei. 2010). 2002. 2009. Mukherjee et al. 2010. 2010). Industry and Medicine but these compounds are of industrial value as well. Apart from the continued discovery of novel secondary metabolites from terrestrial strains. is that some recent cases are reported where Trichoderma is causing infection in humans with normal immunity (see Chapter 17. Trichoderma spp. 2012). has a role in biocontrol. 2008. Although T. Trichoderma harzianum.5 Concluding Remarks The importance of the genus Trichoderma is reflected by the completion of whole-genome sequencing of as many as seven species by the Joint Genome Institute (JGI. This is one example showing that not only Trichoderma but also its genes can be used for bioremediation of pollutants. Viswanathan et al. T.Trichoderma in Agriculture. reesei is not known to secrete appreciable amounts of secondary metabolites. 1. the first antibiotic discovered from Trichoderma (virens) has interesting medicinal properties. produce mycotoxins of the trichothecins group (Tijerino et al. These fungi play immensely important roles in biodegradation and recycling of complex polymers such as the ligno-cellulosic wastes and chitins (Schuster and Schmoll. The research community thus should avoid promoting Trichoderma species that are potential human pathogens. 2007. toxins (such as cyanides) and xenobiotics. T. The direct bearing of Trichoderma in human health is multifaceted. What is more disturbing. other species of the genus are important producers of metabolites with antibiotic activity including polyketides.. have applications as aroma compounds in the food industry. have temperature optima between 25°C and 30°C). 2012b. Consequently. pyrones. 2010.4 Trichoderma in Human Health and the Environment Apart from their use in agriculture and industry. such as Trichoderma brevicompactum. and more recently from the endophytic strains of Trichoderma (Li et al. thus reducing the residue loads in the environment (Harman. Trichoderma longibrachiatum and Trichoderma citrinoviride). 2012. The novel field of application of fungal biotechnology to produce nanoparticles opened up intriguing possibilities with Trichoderma spp.. Besides the classical industrial products of enzymes and metabolites. 2012). Some species. Trichoderma asperellum.. As a rule of thumb.. 2011b). are also important from an environmental and health perspective.. T. some recent innovations in nanotechnology also involve fungi. 1998). These fungi are also capable of remediating heavy metals. strains that grow at or near 37°C should not be used in crop protection (fortunately. Ding et al. this volume). 2011. 2011). 1.. reesei . These fungi are potential sources of medicines. Frenzel et al. A few Trichoderma secondary metabolites such as the viridin-analogue wortmannin 5 (Smith et al. terpenes and polypeptides (Sivasithamparam and Ghisalberti... virens GST took up and converted anthracene to non-toxic naphthalenes (Dixit et al. Ming et al. 2006). some are producers of mycotoxins and some are human pathogens. it can be expected that this field of research will increase in importance in the near future and Trichoderma spp. Rether et al. trichosetin and peptaibiotics are potential anticancer and anti-microbial drugs (Marfori et al. Trichoderma atroviride.. research towards efficient biocontrol fungi now faces the challenge of protecting the plants and the people growing them alike. Shi et al.. Tobacco plants expressing a T. 2012c). however. Other species such as Trichoderma longibrachiatum and Hypocrea orientalis are known to be pathogens in immune-suppressed humans. most Trichoderma spp... several compounds of medicinal importance are being discovered from marine strains (see Chapter 15.. being important organisms for its development (Mukherjee et al..

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