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Organic agriculture has been coming up as one of the promising segments of crop production systems in India. There are numerous reasons for it, however; human health, sustainable environment, soil health, etc. are the important ones. As per the latest information, India has about 1.5% of total cultivable land under organic agriculture. The occurrence of plant diseases in this crop production system is one of the limiting factors. For the management of plant diseases in organically grown crops, there are limited resources since there is a restriction on the use of synthetic fungicides. Under such a situation, bio-pesticides have the potency to take care of plant diseases. Although there are certain fungal and bacterial candidates well efficient in controlling diseases, genus Trichoderma has occupied a prestigious position among them. It is capable of managing seed and soil-borne plant diseases. Presently it is available in wettable powder (WP) and liquid formulations in variable concentrations for the application.
Chaudhary Charan Singh Haryana Agricultural University, India
Dalvinder Pal Singh
Forage Section, Department of Genetics and Plant Breeding, India
Anil Kumar
Deendayal Upadhayay Center of Excellence for Organic Farming Chaudhary Charan Singh Haryana Agricultural University, India
*Address all correspondence to: kishorkumarc786@hau.ac.in
1. Introduction
The interest in organic crop production has been increasing day-by-day since the last decade because of the increased negative impact of conventional agriculture production systems. Production of crops adopting the inorganic agricultural inputs primarily agrochemicals, which has been used starting from seed treatment till harvesting of the crop for providing protective umbrella against various diseases. The frequent and injudicious application of inorganic agrochemical inputs has invited several environmental and health-related problems.
The demand for organic food commodities has been increased tremendously and hence researchers, as well as growers, have been focusing their attention towards organic crop production wherein, the use of inorganic inputs is completely avoided. However, successful crop production through this system faces various biotic and abiotic challenges too.
Organic farming was first initiated by British botanist Sir Albert Howard in 1905. However, in India, it was initiated during the late 90s. The government of India, in the year 2001, implemented the national program for organic production (NPOP) for the promotion of organic farming. Organic farming was started in several states. Sikkim has converted cent percent of the agriculture into an organic state in this country. Presently, India has about 2.3 million hectares under it [1]. with a production of about 1.70 million MT certified organic products with voluminous export of different organic food items [2].
The occurrence of plant diseases is one of the important concerns in organic agriculture. Various fungi are notable/major phytopathogenic fungi causing huge crop losses [3]. For the management of plant diseases, plant protection measures start as early as seed treatment and would continue until crop harvesting in different crops.
In the recent era, organic agriculture is getting huge popularity and adaptability owing to its several beneficial aspects. The organically produced agricultural commodities ensure pesticide-free items which are fit for human consumption. Under such a crop production system, taking care of plant diseases is one of the major challenges. There are limited plant protection options due to the restricted use of synthetic fungicides in it. It emphasises the use of on-farm inputs for almost all requirements. Under such circumstances, the application of Trichoderma spp. could be an ideal alternative to handle plant diseases. Since the last couple of decades, genus Trichoderma has occupied a reputed position among the microbes possessing disease management potency. As a biocontrol agent, the antagonistic property of Trichoderma viride was recognised [4] while working with the damping-off of citrus seedlings caused by Rhizoctonia. Various chemical fungicides were the most dominant during the green revolution era in the 60s and 70s and hence Trichoderma could not significantly attain popularity among the growers at the grass-root level. Due to excessive usage of chemical fungicides numerous problems related to the environment, soil and human health ecosystem, resistance development, etc. have emerged as a big challenge before everyone, including scientists, consumers, growers, and the overall associated environment. Therefore, majority has been preferring pesticide-free food commodities for the last so many years. It is good news or sign that usage of chemical fungicides has been coming down and usage of biopesticides has been going up nationally as well as internationally [5]. Currently, in India, there are several hundred manufacturers of Trichoderma possessing the registration certificate from the Central Insecticide Board and Registration Committee [6]. They are manufacturing and marketing the Trichoderma-based products with variable active ingredients, the dose of application in diversified crop avenues. Although there are four-five species of this antagonist suitable for the management of plant diseases, however only two species namely, T. viride and T. harzianum have been made available to the end-users. The plus point with this antagonist is that it can be used in a wide range of crops, such as cereals, pulses, vegetables, horticultural, and plantation crops [7, 8, 9, 10, 11]. Its wettable powder (WP) and liquid formulations are popular in the market; however, the first one is dominant. In addition to managing the phytopathogen, it is capable of promoting the vegetative growth of plants. Being a natural fungus, it is safer for the environment as well as consumers, capable of adjusting and performing under variable conditions. This chapter highlights various important information related to this genus.
This chapter mainly focuses on the application of Trichoderma spp. for the management of plant diseases. Organic crop production excludes the use of synthetic/inorganic fungicides for the control of plant diseases.
Before touching its actual uses in agriculture, it becomes imperative to focus on some important aspects related to it. For the first time, Persoon introduced the name Trichoderma [12]. In 1865, the Tulasne brothers reported teleomorph (sexual stage) of T. viride Pers as Hypocrea rufa [13]. Till 1969, there were only one species, i.e., T. viride of genus Trichoderma [14].
In 1969, Rifai made out nine ‘aggregate species’ namely, (1) T. harzianum Rifai, (2) T. viride, (3) T. hamatum (Bonord.) Bainier, (4) T. koningii (Oudem.) Duché & R. Heim, (5) T. polysporum (Link) Rifai, (6) T. piluliferum J. Webster & Rifai, (7) T. aureoviride Rifai, (8) T. longibrachiatum Rifai, and (9) T. pseudokoningii Rifai. In the early 1990s, [15, 16, 17] identified five sections and 27 biological species within the genus Trichoderma. The introduction of molecular techniques contributed to the greater extent in identifying the species comparatively more precisely and hence from the late 1990s up to the year 2002, the number of Trichoderma species increased to 47 [18].
Till 2005, the International sub-commission on Trichoderma/Hypocrea listed 104 species on the basis of phylogenetic analyses [19]. Till the year 2015, there are 252 species one variety and one form. In addition, T. neocrassum Samuel (syn. Hypocrea crassa P. Chaverri & Samuels) and T. patellotropicum Samuels (syn. Hypocrea patella f. tropicaYoshim.Doi) were proposed as two new names [20].
3. Interaction of Trichoderma with phytopathogens of agricultural crops
Hyperparasitism, competition, and antibiosis are the important mechanisms of this genus through which it interacts with phytopathogenic fungi. This antagonist is well efficient to colonise in variable ecological niches [21, 22, 23].
3.1 Hyperparasitism
In Hyperparasitism, the Trichoderma directly contact with a pathogen and finally cause the death of pathogen cells death [23]. For this purpose, the Trichoderma synthesise cell wall degrading enzymes (CWDE) namely, cellulase, xylanase, pectinase, glucanase, lipase, amylase, arabinase, protease [24], and lytic enzymes. These enzymes degrade the pathogen cell walls, composed of chitin and glucan polysaccharides. Among the chitinolytic enzymes β-N-acetyl glucosaminidase, endochitinase, and chitobiosidase are of key importance responsible for the degradation of the cell wall of other plants pathogenic fungi, which are produced by T. harzianum, T. atroviride P. Karst, and T. asperellum Samuels, Lieckf. & Nirenberg [25]. Several volatile metabolites, such as 6-n-pentyl-2H- -pyran-2-one (6-PAP), are produced by Trichoderma for plant protection [26]. The β-1,3- and β-1,6-glucanases enzymes determine the hyperparasitic capability of Trichoderma to react on species of Phytophthora sp. and Pythium [27, 28]. Proteolytic enzymes, namely, endo- and exoproteases of Trichoderma responsible for enzyme secretion for the control of Botrytis cinerea, Rhizoctonia solani, and Fusarium culmorum [29, 30]. Ceratin cellulase enzymes, such as exo-β-1,4-glucanases, endo-β-1,4-glucanases, and β-glucosidases, produced by antagonists also play a significant role in hyperparasitism [31].
3.2 Competition
Competition of Trichoderma spp. with plant pathogens is another mode of interaction that helps in the control of plant diseases. This phenomenon takes place for utilisation of nutrients, occupying ecological position or infection sites on plant roots. Certain Trichoderma strains produce siderophores, i.e., iron-chelating compounds. Through siderophores, antagonist traps iron from the associated environment and creates nutrient deficiency due to which the growth of pathogenic fungi, such as B. cinerea, is hampered. The Trichoderma creates an acidic environment that has a negative effect on the growth and development of pathogenic fungi and aggressively colonises the host plant’s roots due to the enhanced activity of the hydrophobins [9].
3.3 Antibiosis
It is a biological interaction between two or more organisms that is unfavourable to at least one of them; it can also be an antagonistic association between an organism and the metabolic substances produced by another. Antibiosis in relation to Trichoderma fungi is a specific mechanism of antagonistic interactions with other plant pathogenic fungi. This phenomenon is based on the generation of secondary metabolites, which exhibit an inhibitory or lethal effect on a parasitic fungus. From the genus Trichoderma, over 180 secondary metabolites have been characterised so far, representing different classes of chemical compounds [32, 33]. Such compounds can be divided into volatile antibiotics, water-soluble compounds, and peptaibols. Volatile antibiotic, (6PAP (6-pentyl-α-pyrone) produced by T. viride, T. harzianum, and T. koningii, plays a major role in the biocontrol of Botrytis cinerea, R. solani, and Fusarium oxysporum.
Another category of antibiotics is Peptaibols. They are polypeptide antibiotics comprising of 500–2200 Da, rich in non-proteinogenic amino acids, specifically alfa-aminoisobutyric acid, and their characteristic attributes include the presence of N-acetylated ends and C-end amino alcohols. Peptaibols exhibit potent activity against a number of fungi (Table 1).
Trichoderma spp.
Produced Peptibols antibiotics
Reference
T. viride
trichotoxins A and B, trichodecenins, trichorovins, and trichocellins
Peptibols antibiotics of Trichoderma spp. to control phytopathogens.
In Trichoderma fungi, the activity of antibiotics is enhanced with the activity of lytic enzymes. Their joint activity provides a higher level of antagonism compared to the activity of either enzymes or antibiotics alone [36]. Researchers found preliminary degradation of the cell walls in B. cinerea and F. oxysporum by lytic enzymes; it facilitated easier penetration of antibiotics to pathogen cells [8].
4. Role of Trichoderma spp. as a biological control agent
Trichoderma spp. are the ideal option for safer management of phytopathogens. T. harzianum and T. viride have been occupied the prime position and its highly exploited biological control agent for controlling soil-borne fungal diseases.
There is a list of about 970 registered manufacturers with the Central Insecticide Board & Registration Committee (CIB & RC) for the manufacturing and marketing of biopesticides in India, out of which 558 are involved in Trichoderma spp. (Figure 1). The different formulations of this antagonist comprised of wettable powder (WP), wettable suspension (WS), aqueous suspension (AS), and liquid. The Trichoderma spp. formulations have been manufactured and marketed by private companies, government organisations, and NGOs. Presently, wettable powder (WP) and liquid formulations are available for use by end-users; however, WP formulation in different concentrations (0.5 to 6.0%) is dominant (Figure 2). The liquid formulations contribute nearly 2% of the formulations.
Figure 1.
Number of manufacturers of different Trichoderma formulations.
Figure 2.
Status of Trichoderma WP and liquid formulation manufacturers.
6. Management through seed treatment with Trichoderma spp.
Seed treatment is an important and economical practice to manage the seed-borne phytopathogens at the initial stage with the minimum cost. For this purpose, a small quantity of desired input is required and this practice is very easy in handling. For seed treatment, the formulation of T. harzianum and T. viride can be used. Seedlings of vegetable crops can be treated with T. viride and T. harzianum just before transplanting into the main field (Table 2).
7. Management through soil treatment with Trichoderma spp.
Soil residential phytopathogens, such as Pythium, Phytophthora, Fusarium, Rhizoctonia, Sclerotinia, Macrophomina, play an important role in the development of various soil-borne diseases, such as root rot, wilt, and seedling rot of various crops. Management of such fungal phytopathogens can be achieved through the application of T. harzianum and T. viride through soil drenching with water, enriched farmyard manure, and vermicompost. There are several species of Trichoderma, however T. harzianum, H. lixii, T. atroviride, H. atroviridis, T. asperellum, and T. virens are potential biocontrol agents against phytopathogens [37]. Application of T. viride enriched FYM (5 kg/plant) to two-year guava saplings at the basin near the root zone resulted in decreased wilt incidence and better plant growth in terms of stem girth [38]. T. harzianum when applied (50 g/vine) in black pepper field it had effectively managed the foot rot disease. For the management of pomegranate wilt bio-formulation of T. viride (0.1 and 0.2%) was found significantly superior over the control. Soil application of T. harzianum plus T. virens is efficient in managing stem bleeding disease of coconut [39]. Five monthly applications of T. harzianum (50 g/plant) in bananas reduced 50% of the vascular discoloration index of Fusarium wilt disease and increased the yield [40]. T. asperellum found inhibitory against Pythiumaph anidermatum, Pythium debaryanum, Sclerotium rolfsii, and S. rolfsii, Fusarium oxysporum f.sp. lycopersici and Alternaria solani. Application of Trichoderma (@20 kg/ha) along with 2.0 tonnes castor cake/ha reduced nematode population and increased yield in pomegranate.
Soil application of silver nanoparticles synthesised from T. asperellum resulted in complete control of Fusarium wilt in cv. Grand Nain [41]. T. harzianum talc formulation could control root rot disease of citrus up to 80%, and under field conditions [42]. Some strains of Pseudomonas spp. and Trichoderma spp. found effective in controlling wilt of banana, caused by F. oxysporum f.sp. cubense (Foc) race 1 under field conditions [43]. Isolates of Trichoderma and Aspergillus, when applied in the field for the control of guava wilt disease caused by F. oxysporum f. sp. psidii and F. solani, could reduce disease incidence and promoted plant growth [38]. Antagonistic fungi Aspergillus niger, T. harzianum, and Penicillium citrinum were found effective for the management of the wilt disease of guava [44].
8. Management through foliar application of Trichoderma spp.
Foliar application of Trichoderma spp. is advisable for the management of airborne fungal and bacterial phytopathogens, such as species of Alternaria, Curvularia, Fusarium, Colletotrichum, Pestalotiopsis, Pyricularia, Puccinia, powdery and downy mildew pathogens of cereal, vegetable and other crops (Table 3). The potent candidates include T. viride and T. harzianum. Post-prune foliar application of T. harzianum and T. viride is a common practice in tea (Camellia sp) crop production in Darjeeling and the North East region of India [45, 46]. When a combination of T. harzianum and Pseudomonas fluorescens was sprayed before harvesting mango fruits, it had suppressed post-harvest fruit rot in dasheri mango [47]. Banana hands (cv. Grand Nain) when dipped in T. asperellum suspension and packed without ethylene absorbent extended its shelf life up to 75 days at 13.5°C. There was no incidence of anthracnose and crown rot [41]. It was noted that T. viride, T. harzianum, and T. asperellum were potential antagonists for the management of F. solani and Pestalotiopsis theae causing dieback and grey leaf spot disease of tea [48, 49]. Foliar spray of T. asperellum and T. atroviride could manage the dieback disease of tea (Camelliasp) and enhance the vegetative growth in terms of more number of pluckable shoots [50].
9. Management through paste application of Trichoderma spp.
Application of Trichoderma paste (20% w/v) is generally done after severe pruning operations (rejuvenation prune and medium prune) in tea plantations to protect the plants from airborne pathogens. Such an application can be useful in horticultural crops in which pruning is done.
10. Important considerations for better performance of Trichoderma spp.
There are certain issues/concerns which should be taken into consideration for achieving desired benefits from the use of Trichoderma spp. Important concerns are—(1) reliable source of procurement, (2) assurance of fresh formulations, (3) avoidance of tank mixing with agrochemicals, (4) tank mixing with compatible bio formulations, (5) repeated application at the proper interval, (6) use of neat and clean separate sprayers for such formulations, (7) seeking technical advice from experts, (8) application during early morning and late evening hours, (9) procurement in well-ventilated store under lock and the key, and (10) quality assurance through certified agencies, such as IMO, Ecocert, Lecon, or any other approved agency.
11. Conclusion
T. harzianum and T. viride are efficient in controlling several plant diseases safely. When considering the interactions of Trichoderma fungi, it was found that these antagonistic fungi have an advantageous effect on plants. Stimulation of plant growth and yield takes place in this interaction and the advantageous effects are seen in the production of vitamins, the increased availability of biogenic elements (nitrogen, phosphorus), the mobilisation of nutrients from the soil and organic matter, and the enhanced intensity of mineral uptake and transport. Furthermore, Trichoderma fungi are capable of producing zeaxanthin and gibberellin, i.e., compounds accelerating seed germination. Managing plant diseases through these approaches could be safer for the agroecosystem, overall environment, soil health, and human health and would be the right step in sustaining crop production to meet the demand of the growing population.
Acknowledgments
We, authors, are highly thankful to Dr. Azariah Babu, Deputy Director (Research), Tea Research Association, North Bengal Regional Research and Development Centre, Nagrakata, Jalpaiguri, West Bengal for critically going through the chapter.
Conflict of interest
The authors declare no conflict of interest.
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Written By
Kishor Chand Kumhar, Dalvinder Pal Singh and Anil Kumar
Submitted: 22 January 2022Reviewed: 16 February 2022Published: 05 April 2022
Open access peer-reviewed chapter
