Hindawi

Advances in Agriculture
Volume 2023, Article ID 5573662, 11 pages
https://doi.org/10.1155/2023/5573662

Review Article
Potential and Prospects of Trichoderma in Plant Protection

Ritesh Kumar ,1 Pratik Samanta,1 Susanth Vijay Raj,1 Pratush Bera,1 and
Mohammed Naimuddin 2
1
Department of Plant Pathology, M. S. Swaminathan School of Agriculture, Centurion University of Technology and Management,
Bhubaneswar, Odisha, India
2
Department of Applied Biology, School of Applied Natural Sciences, Adama Science and Technology University, Adama, Ethiopia

Correspondence should be addressed to Mohammed Naimuddin; [email protected]

Received 28 November 2022; Revised 13 April 2023; Accepted 6 July 2023; Published 31 July 2023

Academic Editor: Mudassar Iqbal

Copyright © 2023 Ritesh Kumar et al. This is an open access article distributed under the Creative Commons Attribution License,
which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

In order to feed the growing population, agriculture is a vital component of any country; however, pests pose a constant threat to it.
Chemical pesticides are employed to safeguard the crops from the enormous yield loss. These chemical pesticides are boundless in
killing crop pests; however, they have detrimental effects on the health of both humans and the environment. Therefore, biological
control techniques are being utilised after identifying an environmentally suitable substitute. Due to its well-known biological
control mechanism, Trichoderma spp. have been utilised extensively in agricultural applications. The host plant’s soil and rhizo-
sphere serve as shelter place for Trichoderma. It has the ability to create a variety of secondary metabolites and secretion of
important enzymes. Clarification of Trichoderma’s importance in the prevention and treatment of plant diseases is thus important
in order to advance sustainable agriculture. The mechanisms, which include mycoparasitism, antibiosis and competition aid in the
management of insect pests and plant pathogens in the soil, seeds, roots, stems, leaves, etc. It is crucial to create new formulations of
biocontrol microorganisms with a greater level of stability and survivability in order to implement biocontrol technology in the
field and enhance its commercialisation.

1. Introduction benign biocontrol agents to control plant diseases, insect pests,

bioremediation and enhance plant growth. In addition to the
By 2050, it is anticipated that there will be 9.1 billion people mycorrhizal fungi, endophytic fungi, endophytic bacteria,
on the planet earth, consequently, an increase in agricultural plant growth-promoting rhizobacteria, plant growth-promoting
food output of around 70% is required to sustain this grow- fungi, and other organisms have been reported for the enhance-
ing global population [1]. The problems like global warming ment of plant growth under various stress conditions [3].
and environmental degradation led to plants experiencing a One of the many biocontrol agents used to manage biotic
variety of biotic and abiotic stress, that are responsible for and abiotic stress in plants are Trichoderma species, which
yield loss of the plants. Additionally, the plant pathogens are predominate in the soil microflora [4]. Due to their signifi-
forced to undergo genetic alterations as a result of the indis- cant antagonistic action against a variety of phytopathogens,
criminate use of fungicides, which ultimately lead to the they can biologically control a variety of plant pathogens that
selection of fungicide-resistant biotypes. Agronomists and cause diseases, including a range of soil and air-borne plant
business sectors have recently demonstrated a strong interest diseases [3]. The ability to act as a biocontrol agent against
in the creation of eco-friendly and economical techniques for other microbes, specially plant pathogenic fungi is a result of
the management of plant diseases [2]. the production of antibiotics and mycoparasitism that affects
Recent strategy that uses naturally occurring plant growth- the growth of other microbes [5]. In addition to producing
promoting fungi rather than pesticides in order to achieve the antibiotics, enzymes, volatile and non-volatile chemicals,
good soil and plant health is an excellent illustration. There Trichoderma also induces systemic resistance in plants [5].
have been numerous reports on the use of environmentally Trichoderma harzianum solubilises several soil nutrients that
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2 Advances in Agriculture

TABLE 1: The classification taxa of Trichoderma [11]. custom search tool and TrichoBLAST, developed by Inter-
Kingdom Fungi national Subcommission on Trichoderma and Hypocrea have
Division Ascomycota greatly helped in the identification of novel species [17, 18].
Subdivision Pezizomycotina However, there are still a large number of Trichoderma strains
Class Sordariomyctes for which sequences have not yet been identified [19]. Over
Order Hypocreales 20 of the estimated 40 described taxa of Trichoderma have
been found to have some agricultural applications [11].
Family Hypocreaceae
Genus Trichoderma
3. Morphological Features of Trichodermas spp.
Trichoderma is a genus under fungi, which is mostly found in
are present in an inaccessible form and transforms them into
soil and root environments and are free-living, anamorphic,
available forms for the plants, which increases the efficiency
filamentous and mostly asexually reproducing [4, 8, 20]. In
of CO2 and O2 utilisation in plants [6]. Also, it is particularly
temperate and tropical soils, Trichoderma species form asco-
helpful for the potential of Trichoderma to promote plant
mycetes with green spores with hyaline and smooth septate
development and produce resistance through controlling the
expression of genes in plants, in addition to having a direct vegetative hyphae (Figure 1(b)). The conidia are single celled
impact on fungal plant diseases [7]. Maximizing the potential and oval or globose in shape. Also, Trichoderma produce
of Trichoderma as bioinoculants for agriculture, requires branched conidiophores on septate, hyaline and smooth-
proper identification of the native strains that are most effec- walled vegetative hyphae. The primary branches’ lateral side
tive there, which is a prerequisite for the development of a branches may or may not be paired, and they occasionally
bioformulation product for agricultural use. may rebranch. In most cases, the branches develop at an angle
The versatility and growth of Trichoderma is suitable on of 90°. Phialides, also referred to as conidiogenous cells, can
a variety of substrates like grains, millets, husk, etc. under in be cylindrical or almost subglobose in shape [20]. The myce-
vitro condition. Naturally, presence of these organisms has lial colour is generally green, but can sometimes be colourless,
been reported in a wide range of soil types, including agri- grey or brownish as seen in Figure 1(a) [13, 21]. On Potato
cultural soils, forest soils, salt marsh soils and desert soils and Dextrose Agar (PDA) and Cornmeal Dextrose Agar substrates,
also in various climatic zones [8]. On rotting wood and in the colonies are white and transparent, respectively and when
soil, due to their heterotrophic interactions, including para- conidia are produced, sporadic blue–green or yellow–green
sitism, decomposition and even opportunistic endophytism, patches can be seen. One or a few phialides normally emerge
Trichoderma species are known to be widespread. It has been straight off from the tip of the typical conidiophore (Figure 1(b)).
shown that Trichoderma grows on plants as epiphytes, endo- Trichoderma spores, which are propagative structures, can ger-
phytes and even in the rhizosphere. minate and grow on the surfaces of plant roots when they are
introduced to soil and even infect the outermost root cells [20].
2. Systemic Classification of Trichoderma For Trichoderma, characters including growth rate, colony colour
and colony appearance are considered as taxonomically useful
The earliest description of Trichoderma was published in traits. Pale or yellowish colour of reverse of colonies, rapid growth
1794 [9] and a relation to the sexual stage of a Hypocrea, at 25–30°C of majority of Trichoderma isolates and typically not
the species was proposed in 1865 by Tulasne brothers [10]. growing at 35°C [22]. By growing at 35°C, T. harzianum can be
The taxonomy of Trichoderma and Hypocrea has been well differentiated from species like T. aggressivum and T. atroviride
reviewed thorough examination of various life stages (Table 1). that share a similar morphology. While T. harzianum grows well
However, morphologically it was challenging to differentiate and sporulates at 35°C, neither T. aggressivum nor T. atroviride
among the numerous species of Trichoderma. In fact, it has can have colonies greater than 5 mm after 96 hr [22]. Addition-
been suggested that just Trichoderma viride should be consid- ally, some Trichoderma species, like T. viride, emit a distinctive
ered as a single species; an identification concept was not sweet scent similar to ’coconut’ odour [20]. Within 30 samples,
developed until 1969 [12, 13]. Harz first identified the species morphological analysis identified two different conidiophore and
Trichoderma harzianum in 1871 after the study of its micro- phialide arrangement types. The first set of 24 isolates had con-
scopic features, particularly the presence of phialides [14]. idiophores that were spread out at the top and wide near the base
Internal transcribed spacer (ITS) region, restriction frag- that were smooth or rounded. Phialides mostly appeared in
ment length polymorphisms and random amplified poly- densely populated areas, but they also had whorls of two
morphic DNA sequence analysis are among the strategies to six on the terminal stems and an angle with conidio-
to identify microbes using DNA markers. This has piqued phores. The penicillate type second group of six isolates
the interest of Trichoderma workers to incorporate sequence was distinguished by predominate conidiation, numerous
data to the developing taxonomy of Trichoderma and Hypo- divided ranches, collecting all finger to top and fertile to the
crea. Several new Trichoderma that belongs to Hypocrea spe- apex [23]. Chlamydospores serve a role in survival and are
cies were discovered, bringing an increase in number of vegetative cells with thickened walls and compacted cyto-
phylogenetically recognised species [15]. T. harzianum was plasm [24]. These colourless, pale yellowish, or greenish, glo-
misidentified in some situations, mostly in early reports [16]. bose to subglobose structure are produced either within
TrichOKEY, an oligonucleotide barcode, along with a hyphae or at the hyphal tips [20].
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Advances in Agriculture 3

ðaÞ ðbÞ
FIGURE 1: Mycelial (a) and microscopic (b) images of Trichoderma spp. (Structure in the red circle are denoting the Phialides of respective
Trichoderma).

4. Identification of Trichoderma spp. Based on cold-tolerant plant pathogenic fungus. T. aureoviride and
Morphology and Molecular Markers T. viride grow well at 5°C [29], while T. harzianum are
adapted to warm climate [30]. The antagonistic potential
Trichoderma isolated from soil samples, and incubated for of cold-tolerant Trichoderma strains was tested at different
7 days at 28°C on PDA media shows green colony pigmenta- temperatures, and it was discovered that temperature had no
tion as seen in Figure 1. Traditional methods have previously effect on hyphal contacts [31]. When compared to T. harzia-
been used to identify Trichoderma spp. by its morphology like num strains, T. aureoviride and T. viride strains are more
linear growth, colony colour, growth pattern, hyphal pigmen- efficient in vitro antagonists as well as better cold tolerants.
tation and cultural structures include the arrangement of con- Extracellular ß-1,4-N-acetyl-glucosaminidase (NAGase),
idiophores, phialides and shape of conidia (Figure 1) [13, 23]. trypsin and chymotrypsin-like proteases, ß-glucosidase
The rate of growth and microscopic characteristics of isolated and other enzymes involved in the mycoparasitic process
Trichoderma, such as the sizes and shapes of conidia, the were produced at 10°C and found to be quite active at 5°C
branching patterns of conidiophores and the sizes and shapes in the cold-tolerant strains [32].The poor osmotolerance
of phialides, are also crucial physical factors in identifying the level of Trichoderma strains is one of the most significant
various Trichoderma species [25]. Identification of Tricho- drawbacks of using them as a biocontrol agent in field
derma species can also be benefited from observations of condition. Soil water conditions are limiting elements
spore and phialide sizes, but most of the isolates cannot be that affect fungal activity. As a result of natural drying
identified using just the accounts of the conidial shapes as due between two rainfalls, dry conditions can develop even in
to genetic, environmental and nutritional changes, the fungus soils that are usually moist. In dry soils, however, biocon-
has displayed varied morphologies on culturable media that trol chemicals may be required to combat plant pathogens.
makes the identification challenging. Molecular identification Water conditions have been shown to have a significant
by using fungal ITS sequencing is a widespread method,
impact on Trichoderma activities, including spore germi-
which confers a reliable identification. Molecular identifica-
nation and germ tube growth, mycelial growth [33, 34],
tion utilising ITS region and their respective morphological
saprophytic ability [35], interaction with other fungi and
characteristics of different Trichoderma species are sum-
enzyme production [36]. Information on the effects of
marised in the tabular format (Table 2). Additionally, the
water conditions on the metabolic activities of Tricho-
methods for identifying Trichoderma with DNA markers
derma strains is critical for biocontrol strategy develop-
and the use of DNA sequence analysis became new ideas in
ment. At both 25 and 10°C, there is a linear relationship
the field of fungal systematics. Numerous researchers sug-
gested using more variable phylogenetic markers, such as between water potential and colony development rate, with
tef1_int4, tef1_int5 and rpb2, that encode RNA polymerase larger growth rates at higher temperature and water poten-
subunit 2 and translation elongation factor 1-alpha (EF-1) tial. Cellobiohydrolase and NAGase enzymes showed opti-
[12, 13]. A significant amount of novel traits in terms of mal secretion at higher water potential. Aside from those
divergent base pairs are accumulated by DNA sequencing that are best for growth, ß-glucosidase, ß-xylosidase and
data and their incorporation into gene banks, contributing chymotrypsin-like protease enzymes show the greatest
to cladistics analyses of most recent shared ancestry. activity at lower water potential values [36, 37]. A certain
pH is required for the use of biocontrol Trichoderma
5. Effects of Temperature, Water and pH on strains in agricultural soils. Because of this, it is necessary
Growth of Trichoderma to gather data regarding the effect of pH on mycelium
growth and in vitro activities of extracellular enzymes
Trichoderma species can be grown on a wide variation in that participate in nutrition competition and mycoparasit-
optimal temperatures [27]. Most Trichoderma strains are ism in Trichoderma strains with respect to its biocontrol
mesophilic [28], which means that they cannot protect ger- potentiality. Trichoderma strains can grow in a pH range of
minating seeds from soil-borne plant diseases caused by 2.0–6.0, with a preference towards pH 4.0. At this pH, there
 4

TABLE 2: Molecular identification and morphological characteristics of different Trichoderma species [26].
S. No. Trichoderma Locus Primer pair used Amplicon Morphological and cultural characteristics
The colour of mycelium varies from watery white to light green. The petri dish’s reverse side
ITS1-AGAGTTTGATCCTGGCTCAG
1 T. harzianum KC800922 546 base pairs displays areas of uncoloured rings. Conidiophores grow in loose tufts and are heavily branched.
ITS4-GGTTACCTGTTACGACTT
Phialides have a short, skittle-like form with a centre bulge and a base that is narrower
Mycelium is smooth and hairy with a cotton pattern that is often composed of 1–2 ringed
ITS1-TCCGTAGGTGAACCTGCGG
2 T. asperellum KC800921 1,200 base pairs concentrics in a yellowish green colour. Conidiophores have a compact morphology and are
ITS2-TCCTCCGCTTATTGATATGC
heavily branched. Phialides have a nine-pin appearance
The mycelium changes from green to a dark yellowish green colour after 2-3 days. Green
ITS1-TCCGTAGGTGAACCTGCGG
3 T. viride KC800920 641 base pairs conidia are spread out all throughout, with conidia production being lower in the centre than at
ITS4-TCCTCCGCTTATTGATATGC
the borders. Phialides are long, thin, horn-shaped, and bulging in the centre
The mycelium mat looks watery white and appears translucent and smooth. Conidiophores are
ITS1-TCCTCCGCTTATTGATATGC
4 T. atroviride KC008065 627 base pairs highly branched and arise in compact form. Phialides appear in ampulliform and oblong
ITS2-GGAAGTAAAAGTCGTAACAAGG
shaped, curved, and narrow at the base
Mycelium are mostly submerged translucent or watery white. Conidiophores are smooth, form
ITS1-TCCTCCGCTTATTGATATGC
5 T. longibrachiatum JX978542 664 base pairs irregular tuftsand arise from substratum or from aerial hyphae. Phialides can appear
ITS2-GGAAGTAAAAGTCGTAACAAGG
individually or in verticils of two to three. They often have lageniform shapes and a narrow base
The colour of mycelium is creamy white and shifts from white to terreverte. Colonies resemble
ITS1- TCTGTAGGTGAACCTGCGG glaucous membranes and are crusty and dense. Chlamydospores can develop terminally or
6 T. koningii KC800924 206 base pairs
ITS4-GGAAGTAAAAGTCGTAACAAGG intercalarily. At the base, phialides are thin. They seem like a nine-pin bowling ball when they
emerge singly and laterally
Mycelium does not have a distinctive smell and goes from being watery white to a lovely shade
ITS1-TCCTCCGCTTATTGATATGC of green with granules that are dull blackish green. Conidiophores branch irregularly towards
7 T. virens KC800923 635 base pairs
ITS4-GGAAGTAAAAGTCGTAACAAGG the tip, with each branch ending in a group of three to six tightly packed phialides. Phialides are
lageniform to ampulliform in shape, with a bulging midsection and attenuated tip
Advances in Agriculture

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Advances in Agriculture 5

Mechanism for suppression

Suppression of

of plant pathogen
Mycoparasitism
plant pathogens

Antibiosis

Competition

Presence of microbes in the plant rhizopheric region

Trichoderma in the
rhizospheric region

FIGURE 2: Mechanisms of Trichoderma spp. for suppression of growth and viability of plant pathogens.

is a regulation of the synthesis of extracellular enzyme TABLE 3: List of genes involved in mycoparasitism.
ß-1,6-glucanase in T. harzianum [38], also it has been
seen that the optimal pH range for biomass generation in S. No. Gene Protein Reference
Trichoderma is between 4.6 and 6.8 [39]. 1 prb1 Basic proteinase [46]
2 ech42 Endochitinase [47]
6. Microbial Compatibility between 3 chit33 Endochitinase 33 [48]
4 cre1 Carbon catabolite repressor [49]
Trichoderma, Different Bacteria and Plant
5 egl1 β-1,4-Endoglucanase [50]
Pathogenic Fungi
It has been discovered that numerous soil bacteria, such
as fluorescent Pseudomonads, may have an impact on Tri-
against plant pathogenic fungus is due to its essential antag-
choderma development in agricultural soils (primarily sup-
onistic property of mycoparasitism [43]. Cell wall degrading
pressive soil) [40]. There are numerous accounts on the
enzymes (CWDEs) and antimicrobial secondary metabolites
enzymological underpinnings of the mycoparasitic activities
make up the intricate process of mycoparasitism [44]. Endo-
of Trichoderma strains, but very few studies on bacteria that
adversely affect Trichoderma’s biocontrol abilities. Tricho- chitinase, β-1, 3 glucanase and chitobiosidase are the chiti-
derma’s competitive success might be attributed to its sensi- nolytic enzymes produced by T. harzianum, and are efficient
tivity to the soil bacteria’s inhibitory impact. Under inductive against plant pathogenic fungi. Chitobiosidase and endochi-
and noninductive conditions, the activities of NAGase, tinases can suppress the plant pathogenic fungal spore ger-
trypsin-like and chymotrypsin-like proteases were examined mination and hyphal lysis, while endochitinase and β-1-3
in the instance of five strains that demonstrated exceptional glucanase have been demonstrated for antifungal activity
degrading abilities toward Bacillus subtilis [40]. All strains via lysis of spore cell walls, thallus and hyphal tips [45].
generated modest amounts of NAGase and proteases, which Various genes have been identified for mycoparasitism in
may be increased through induction using B. subtilis cells. biocontrol activity (Table 3).
NAGase and proteases were three to six times more abun- Competition for resources and space, or ability to pre-
dant in inductive medium. The disintegration of bacterial vent spore germination, killing of cells (antibiosis), or alter-
cells appears to be significantly influenced by proteases, ing the rhizosphere, such as by making the soil acidic, help
NAGases and muramidases. Testing Trichoderma strains’ Trichoderma spp. in preventing the growth of phytopatho-
capacity to destroy bacteria might be helpful in addition to genic fungi. As most plant pathogenic fungi require iron
determining how well they can fight plant pathogenic fungus uptake for viability [51], competition for this (iron) limited
[41]. This characteristic may help the strains become the nutrients results in the biological control of fungal phyto-
prevalent microorganisms in the environments where they pathogens. When iron is under scarce amount, the fungi
are used. Trichoderma’s ability to parasitise other plant path- release siderophores (low-molecular-weight ferric iron-specific
ogenic fungi prompted the first research on biocontrol with chelators), to mobilise environmental iron [52]. Highly effec-
Trichoderma [5]. However, it required another 40 years to tive siderophores produced by Trichoderma chelate the iron
prove that using Trichoderma in field might stop the spread and prevent the growth of fungi [53]. Interactions with low-
of fungus-related plant diseases [42]. Numerous articles have molecular-weight diffusible substances or antibiotics pro-
since highlighted mycoparasitism as a crucial biological con- duced by Trichoderma strains that prevent the growth of
trol mechanism in fungal diseases. Antagonistic tactics which other plant pathogenic microbes result in antibiosis. Also,
include mycoparasitism, competition for nutrients and the production of 6-penthyl—pyrone, alamethicins, gliovirin,
resources and antibiosis (Figure 2) are used by Trichoderma glisoprenins, harzianic acid, heptelidic acid, massoilactone,
spp. to combat plant diseases. Trichoderma’s effectiveness peptaibols, tricholin and viridin are some of the volatile
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6 Advances in Agriculture

and non-volatile metabolites secreted by most of the Plant pathogens

Trichoderma [54]. Having genes for
cutinase, cellulase,
7. Plant Disease Suppression by Trichoderma hemicellulase, and
pectinase
Mycoparasitism is the act of attacking another pathogenic
fungus directly, in order to use it as a food source. The four
phases of process are detecting prey, growing positively toward
it, chemical and physical attacks, which usually destroy the Cutin
hyphae of the prey and eventually consuming nutrients [13].
Combining mycoparasitism, competitive exclusion and anti- Cellulose
biosis has an impact on Trichoderma-mediated disease sup-
pression [55]. The physical attack during the contact phase Pectin
results in morphological changes, including hyphal coiling,
intense branching or the formation of appressorium-like struc-
tures, whereas the chemical attack occurs before contact, when Cutinase Xylanase
enzymes that breakdown cell walls are produced and released Endoglucanase Endopolygalacturonase
in mass quantities. When the fungi used for biocontrol is cellobyohydrolase rhamnogalacuronan hydrolase
administered to roots or seeds, Trichoderma spp. quickly col-
FIGURE 3: Interaction between a plant pathogen and host cell wall.
onise. Infectious pathogens are avoided by the rhizosphere
(root zone) and the spermosphere (seed zone). In direct
antibiosis, Trichoderma’s secondary metabolites or released by Trichoderma spp. As lysis of the plant pathogenic fungal
enzymes prevent the germination or growth of pathogens cell wall is one of the key steps during mycoparasitism and
[56]. The metabolites may also influence mycoparasitism thereby controlling of plant pathogen, different strains of
and competition. Because mycoparasitism is difficult to Trichoderma harzianum have been found to be effective
study in vivo, there are fewer direct study of mycoparasit- in controlling the plant pathogens [58]. One of the main
ism’s effects on plant diseases. The first detailed study of mechanisms put forth to explain the hostile activity of Tri-
the mycoparasitic behaviour of T. virens (then known as T. choderma species against a variety of soil-borne plant patho-
lignorum) on Rhizoctonia solani showed that there is a coiling genic fungi, such as Sclerotium rolfii, Rhizoctonia solani and
of hyphae, growth in wavy or straight lines, protoplast coagu- Pythium aphanidermatum, is their direct mycoparasitic
lation and loss of vacuolated structures [5]. Gliotoxin was activity. [59]. Trichoderma’s chitinases are mainly involved
ultimately discovered to be the cause of this Trichoderma in (i) mycoparasitism (ii) exogenous chitin degradation and
strain’s antagonistic activity and pot tests revealed that this (iii) recycling of the fungal cell wall [60]. The chitinolytic
strain effectively suppressed the R. solani strain [5]. enzyme system of Trichoderma can be divided into sub-
groups A, B and C and is a member of GH family 18. Sub-
8. Mycoparasitism and Cell Wall- group B is believed to be an endochitinase and several of its
Degrading Enzymes members contain a CBM1 domain at their C terminus, while
subgroups A and C are predicted to be exochitinases that
CWDEs released by phytopathogenic fungi have been shown
cleave close to the ends of the chitin chain and release
to play an important role when the cell wall of a plant cell is
N-acetylglucosamine dimers. CBM 18 and CBM 50 domains
invaded by a pathogenic fungus. When the primary struc-
can be found at the N-terminus of the GH 18 module of
tural polysaccharide components of plant cell walls, such as
subgroup C chitinases [60].
cellulose, hemicellulose and pectin are broken down by fun-
gal CWDEs (Figure 3), pathogenic fungus have the opportu-
nity to utilise small polysaccharide molecules as a source of 9. Trichoderma as an Insect Pest
nutrients. Additionally, CWDEs from fungi have been iden- Biocontrol Agent
tified as important sources of Microbe-Associated Molecular
Pattern that trigger primary immune responses in plant cells, Numerous microorganisms that operate as entomopatho-
or fungal cell wall [57]. Also, induced systemic resistance is a gens and interact with plants in the soil can be employed
type of active resistance that also relies on these types of as efficient biocontrol agents against various agricultural
structural and biochemical changes in plants. Various Tri- insect pests [61]. Beauveria, Cordyceps, Hirsutella, Isaria,
choderma strains may be suggestive for the emergence of Lecanicillium, Metarhizium, Ophiocordyceps, Paecilomyces,
plant-induced systemic resistance [57]. In case of attack by Pochonia, Torubiella and Trichoderma are some of the
the plant pathogens, sequential events, including host recog- most investigated and utilised genus in agriculture [62]. Tri-
nition, attack and subsequent penetration as well as killing choderma is found to be effective against different type of
of the host occur during mycoparasitism. Trichoderma spp. insect pests (Table 4). The phytohormones salicylic acid
have the ability to recognise other fungus, migrate there and (SA), jasmonic acid (JA) and ethylene are the signal trans-
directly parasitise them. Plant pathogenic fungi such as duction pathways that are activated by an arthropod attack
Botrytis cinerea and some Fusarium spp. can be parasitised on plants [70, 71]. The effects that these insects have on
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Advances in Agriculture 7

TABLE 4: Biocontrol capacity of Trichoderma against different pests.

S. No. Trichoderma Effective against Reference
1 T. asperellum Tetranychus urticae [63]
2 T. viride, T. harzianum Odontotermes formosanus [64]
3 Trichoderma sp. Amrasca bigutulla bigutulla [65]
4 Trichoderma sp. Aphis gossypii [65]
5 Trichoderma sp. Myzus persicae [66]
6 Trichoderma sp. Locusta migratoria [67]
7 T. viride Corcyra cephalonica [68]
8 T. viride Helicoverpa armigera [69]

TABLE 5: Various methods to prepare biocontrol formulation of Trichoderma [78].

S. No. Formulation type Methodology
Sterilized talc powder is mixed with mycelia grown in liquid medium at 1 : 2 ratios. When the
1 Talc based medium reaches 8% moisture, it needs to be dried under shade. The Trichoderma talc
formulation can be stored for 3–4 months
In an oven, 33 g of wheat bran and 100 g of vermiculite are sterilised for 3 days at 70°C to
2 Vermiculite-wheat bran based multiply the mycelia. Then 20 g ferment or biomass is mixed with 0.05N medium and the
entire concentrated biomass is mixed with HCl and dried in shade
The Fermentor biomass is mixed with 100 g of wheat flour. Then a layer of 1 mm thick dough is
3 Pesta granules based prepared and air-dried until it breaks crisply. After air drying, the dough sheet is ground and
passed through an 18 mesh to collect the granules
Different banana wastes are chopped into small pieces, such as the sheath, pseudo stem, and
core, and layered in a pit. Different ingredients are added in five layers within the pit. One ton
of banana waste, 5 kg of urea, 125 kg of rock phosphate, and 1 L of broth culture of
4 Banana waste based
Trichoderma are mixed thoroughly and that much of requirements for single layer. Five
different layers are prepared in the same way and mixed thoroughly. Within 45 days, banana
waste decomposes and is easily used in the field
One week old T. viride culture broth evenly mix with 120 kg press mud. To keep moisture in
the mixture, water is sprinkled on. It is covered in gunny bags, which also help air circulation
and trap the moisture. As the same procedure, eight tons of press mud are added to it, mixed it
5 Press mud based
well, and incubated for 8 days under shade. In this way, 8,000 times more inoculums of
biopesticides were added, then recommended, resulting in rapid and visible establishment of
the biopesticides

plants vary depending on how they attack them, with pierc- fungi are required to create a wide range of insecticidal sec-
ing/sucking species having different effects than chewing ondary metabolites throughout the procedure in order to
organisms [72]. Piercing and sucking type of insects like complete their life cycle.
aphids trigger SA-related responses, while chewing type of
insects like caterpillars activate the JA signalling pathway 10. Trichoderma as a Biopesticide
[73]. Despite the fact that their effects on plant pathogens
have been extensively studied in the lab and in the field, The search for eco-friendly biopesticides is critical, as well as
investigations on their impact on insect pests are scarce being able to resist several types of plant pathogens. A highly
and primarily recent. Research in the lab revealed that Tri- effective biocontrol agent, high levels of productive and via-
choderma spp. negatively affect both chewing type of insects ble propagules and bioprotectant-friendly transport mechan-
[74] as well as piercing and sucking type of insects [75]. The isms give the biocontrol agent a competitive edge over other
field study on Trichoderma spp. on the population of pests microflora [77]. Due to the importance of Trichoderma, it is
associated with maize [76] demonstrated that the abundance more commonly used than other biopesticides in agriculture
of piercing and sucking type of insects reduced under field, for the management of plant pathogens. It is risk-free to
while the population of natural enemies increased, support- control plant diseases through biological means when resi-
ing laboratory findings. Entomopathogenic fungi typically dent antagonists are improved. To make adequate quantities
infect insects directly through the cuticle, a process that of active and viable Trichoderma inocula, both solid and
necessitates adhesins and lytic enzymes (chitinases, proteases liquid forms are used. Hyphae, chlamydospores and conidia
and lipases). In the end, the fungus forms and disperses new are the three types of propagules that can be used in formu-
conidia from its dead host after defeating the insect’s lations [56]. A comprehensive integrated pest management
immune system and colonising its body. Entomopathogenic programme promotes disease management in a similar
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8 Advances in Agriculture

TABLE 6: Different methods for the application of Trichoderma.

S. No. Methods Application
1 Seed treatment Delivered in the infection court, i.e., surface of seed coat as protectant at the time of planting
Seeds are hydrated to allow the germination but not sprouting. Two types of priming are available.
Solid matrix priming (SMP) to enhance biocontrol of Trichoderma by regulating water levels in the
2 Seed priming
seed. Osmopriming makes use of aerated aqueous solutions of salts or polyethylene glycols to
generate osmotic potential in the primary solution
3 Liquid coating Application of Trichoderma to the seed with an aqueous, adhesive or binder
4 Double coating Trichoderma is applied directly to the seed coat followed by a particulate to form the second layer

manner to fungicides. The Trichoderma produces a variety of competition for nutrients and resources and antibiosis.
antimicrobial products that act as biocontrol agents and The use of Trichoderma species should be promoted as a
affect other microorganisms [69]. There are various methods viable alternative to pesticides in the era of a green economy,
developed for the mass production of Trichoderma (Table 5). focused on safeguarding both human health and the envi-
ronment in light of the information provided in the present
11. Compatibility of Trichoderma with Other review. The existing methods for detecting and assessing
Biological Systems and their Application these antagonists, which integrate multiple mechanisms of
action and are quick, affordable as well as effective, must be
A single strain of Trichoderma might not be able to control improved upon and used widely. The current compilation of
all plant diseases and pests, therefore may require a combi- works also emphasises the commercialisation of these
nation of strains or another biological control agents as it is biocontrol-based biostimulation and bioremediation pre-
extremely disease specific [79]. The compatibility of the parations for use in farming practises in a more natural
T. harzianum and Glomus trapped in alginate beads with way. However, more research is needed in this area and
no inhibitory effects on each other [80], control of sheath against more diverse groups of plant pathogens as well as
blight of rice caused due to Rhizoctonia solani by the com- insect pests in order to make Trichoderma a viable alternative
bined use of Pseudomonas fluorescens and T. viride in talc for the future of agricultural plant health.
formulation [81], reflected a synergistic type of combining
ability of Trichoderma with other biocontrol agents. Data Availability
Trichoderma’s ability to biologically regulate plant
pathogens depends not only on its ability to act as an antag- All the data are provided in the manuscript.
onist, but also on how it is applied to the soil, roots and seeds.
Additionally, for effective defence and control, all antago- Conflicts of Interest
nists depend on where they are on the infection court [81].
Trichoderma is typically only effective as a preventative mea- The authors declare that they have no conflicts of interest.
sure but can be integrated with other disease management
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