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Medicinal Plants – Chemical, Biochemical, and Pharmacological Approaches
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the rhizosphere, under limiting conditions, siderophores may also inhibit the growth of pathogens that could potentially cause damage to the plant [64]. Trichoderma spp. producing siderophores in rhizospheres can restrict iron and make it less available to pathogens, indirectly promoting plant growth [65]. Studies conducted by [59] showed the role of siderophores produced by T. asperellum T34 in controlling F. oxysporum, reducing tomato infestation and stimulating plant root growth.
. Biomolecules enhancing plant growth
. Production of phytohormones
Phytohormones play an important role in agriculture [66]; they are synthe­sized by many rhizosphere microorganisms including Trichoderma spp. They have various roles such as modification of the physiological functions of plants to accelerate their growth by intensive cell division in callus tissue, promotion of phloem development, enhance lateral root development, plant growth stimula­tion and prevention of leaf aging by slowing down the breakdown of chlorophyll pigments in plants as well as improving metabolism even at low concentrations [67–69]. IAA and gibberellins (GAs) are among the most important phytohor­mones that regulate the plant’s development and enhance plant growth through several processes [70–74].
. Acquisition and nutrients solubilization
Various fungi such as Trichoderma spp. are associated to the plant roots’ rhizo- sphere, they provide nutrients, protection against biotic and abiotic stresses, and stimulate plant growth [75, 76]. Trichoderma species have the ability to acquire nutrients in the rhizosphere through various mechanisms.
.. Phosphate solubilization
Phosphorus is important elements for plant growth. It can be found in two forms: organic phosphorus and inorganic phosphorus, which usually forms insoluble mineral compounds with calcium, aluminum, or manganese [77, 78]. The distribution of these forms in soils is influenced by several factors such as microbial activity, pH, soil type, and organic matter availability [1]. Recently, phosphate solubilizing microorganisms have attracted the attention of agrono­mists; these microorganisms were used as soil inoculum to improve plant growth [79]. Plants and fungi including Trichoderma spp. compete for the limited avail- able phosphorus through various processes, such as solubilization, precipitation, absorption, and desorption. Inorganic phosphate and organic phosphorus can be mineralized through enzymatic action [1]. Trichoderma species have the ability to solubilize insoluble phosphate into soluble phosphate [80, 81]. In previous studies; [82] reported that Trichoderma atroviride LBM 112 and T. stilbohypoxyli LBM 120 revealed positive results for phosphate solubilization with formation of halo-zone on the solid medium containing insoluble inorganic phosphorus source. In addi­tion, T. harzianum T11 (OL587563) isolated from rhizosphere soil of olive trees has several plant growth-promoting traits, such as the phosphate-solubilizing ability and the production of siderophores [74].
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.. Nitrification and nitrogen fixation
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Nitrogen fixation processes have significant ecological importance in various
ecosystems, including those of agricultural interest. Nitrogen plays a critical role in plant synthesis as it is a component of important biomolecules such as nucleic acids, peptides, organic acids, and fatty acids, which are necessary for the structure and activity of all organisms. Nitrogen-fixing by microorganisms play a key role on growth-promoting plant. It has been suggested that the promotion effect on plant growth might be mediated by providing nitrogen through biological nitrogen fixation and hormones [83, 84]. Production of ammonia and nitrogen-fixing abil­ity by Trichoderma strains are reported in previous findings. Ahemad and Kibret [85] reported that ammonia is useful for plants as directly or indirectly. Ammonia production by the Trichoderma isolates may influence plant growth indirectly;
Figure 2. Schematic description of the main mechanisms used by Trichoderma spp. to competitively colonize the rhizosphere of host plants [74].
Chemical nature Secondary metabolites Trichoderma
Alcohol 2-Phenylethanol T. harzianum Reduces the
Anthraquinone Pachybasin T. harzianum Increases the
species
Emodin T. viride Antimicrobial and
Bio-activity observed
growth of Aspergillus flavus and aflatoxin production
number of coils of the biocontrol agent against R.
solani
antineoplasic agent
Reference
[87, 88]
[89]
[90–92]
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Chemical nature Secondary metabolites Trichoderma
Azaphilone T22azaphilone T. harzianum Inhibits the growth
Bisorbicillinoid Bisvertinolone T. longibrachiatum Antifungal
Butenolide Dehydro derivative of
Hydrolytic
enzymes
Indolic
compound
Koninginins Koninginins A–E T. koningii
Monoterpene β-Myrcene T. virens Regulates the
harzianolide
Cellulases T. re es e i Degrades cellulase
β-1,6- Glucanases Trichoderma sp. Hydrolyses
Chitinases Trichoderma sp. Hydrolytic
Indole-3- acetic acid (IAA)
Indole-3- acetaldehyde T. atroviride, T.
Indole-3­carboxaldehyde
species
T. harzianum Antifungal
T. atroviride, T. virens
virens
T. atroviride, T. virens
T. harzianum
Bio-activity observed
of R. solani, Pythium ultimum and Gaeuman nomyces graminis
properties via inhibition of β- (1,6)-glucan biosynthesis
activity against
Gaeumannomy ces graminis var. tritici
during root colonization to penetrate the plant tissue
fungal pathogen cell walls of B.
cinerea, R. solani, Phytophthora citrophthora
enzymes of the fungal cell wall
Controls a number of growth and development processes in plants
Controls root growth in
Arabidopsis thaliana
Induces adventitious root formation in
A. thaliana
Antifungal activity against
F. oxysporum, Fusarium solani, and Alternaria panax
expression of genes
Reference
[93]
[94]
[95]
[96]
[17]
[97, 98]
[99]
[99]
[100]
[101, 102]
[22, 103]
Biomolecules Produced by Trichoderma Species as Eco-Friendly Alternative Suppressing… ITexLi.112028
Chemical nature Secondary metabolites Trichoderma
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Nitrogen heterocyclic compound
Peptide Trichokonin VI (Tk VI) T. longibrachiatum Inhibits primary
Pyrane Koninginin A T. koningii Plant growth
Pyridones Hharzianopyridone T. harzianum antifungal activity
Pyrones 6-Pentyl-2H- pyran—2-
Siderophore Fusarinine C Trichoderma sp. Fe-chelated, can be
Harzianic acid T. arundinaceum;
Harzianopyridone T. harzianum Antifungal activity
Melanoxadin T. sp. strain
Koninginin D T. koningii Alters pathogen
one
Ferricrocin T. atroviride
Coprogen B Trichoderma spp. Solubilizes iron
species
T. harzianum
ATF-451
Trichoderma viride T. atroviride
a
virens
, T. re e se i
a
, T.
a
Bio-activity observed
Antimicrobial metabolite, siderophore and plant growth regulator
against B. cinerea, R. solani and inhibitor of the protein phosphatase type 2A (PP2A)
Inhibits melanin formation in the larval hemolymph of the silkworm,
Bombyx mori
root growth in A. thaliana
regulator
fungal growth of R. solani,
Phytophthora cinnamomi, Pythium middletonii, F. oxysporum and Bipola ris sorokiniana
against plant pathogenic fungi, such a s P. ultimum, G.
graminis var. tritici, R. solani, and B. cinerea
Antifungal activity against R. solani, F.
available to plants
Key metabolite in the competition for iron in the rhizophere
unavailable to the plant
Reference
[104–106]
[107]
[108]
[109]
[110]
[111]
[112]
[113]
[114]
[115]
[116]
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Chemical nature Secondary metabolites Trichoderma
Steroidal
compound
a
http://genomebiology.com////R
Viridin T. koningii, T.
species
virens, T. viride
Bio-activity observed
Antifungal metabolite that alter the spore germination of Botrytis
allii, Colletotrichum lini and Fusarium caeruleum
Reference
[55]
Table 1. Secondary metabolites secreted by Trichoderma sp. and their bio-active role.
ACC synthesized in plant tissues by ACC synthase is released from plant roots and taken up by neighboring micro-organisms. Then, Trichodrema may hydrolyze ACC (1-aminocyclopropane-1-carboxylic acid) to ammonia. Besides, [74] reported that production of ammonia by Trichoderma species isolated from rhizosphere soil of
Compound Strain Cro ps Application
Biofertilizer Trichoderma
azevedoi
Trichoderma afroharzianum
T. harzianum, T. asperellum, Trichoderma hamatum, T. atroviride
Trichoderma brevicompactum, Trichoderma gamsii, T. harzianum
T. harzianum T. asperellum
T. brevicompactm, T. gamsii, T. harzianum
Lettuce Simple
Tomato Seed
Chinese cabbage
Tomato Seedling
Tomato Seed
Tomato Seed
mode
exposure
inoculation or treatment
Irrigation Increases soil enzyme activity,
drenching
treatment
drenching
Beneficial outcome References
Increases carotenoids and chlorophyll with reduction in the white mold attack to about
78.83%
Helps in the secretion of Phytohormones like homeostasis, antioxidant activity, phenylpropanoid biosynthesis and glutathione metabolism
yield by 37%, and increases the concentration of inorganic nitrogen and phosphorus content of the soil
Improved growth and yield due to the production of IAA
Improves phosphorus uptake
Improves phosphorus solubilization
[117]
[118]
[119]
[35, 120]
[121]
[120]
Table 2.
Trichoderma sp. as bio-fertilizers and their role in promoting plant growth and yield.

Biomolecules Produced by Trichoderma Species as Eco-Friendly Alternative Suppressing… ITexLi.112028
olive is sustained with the results obtained by [86] who reported that among 20
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Trichoderma spp. isolated from chili rhizosphere, 13 isolates were able to produce
ammonia (Figure ; Tables  and ).
. Conclusion
This report reviews the importance of suppressing the growth of the fungal pathogens and as biofertilizer enhancing plant growth. Therefore, the increase use of cides and biofertilizers offers promising prospects for sustainable and environmen­tally friendly agriculture. These eco-friendly alternatives can substitute the excessive use of chemical products that can cause problems in the long term. The biotechno­logical advances from these microorganisms such as fungi are immense and yet to be explored. Thus, more studies need to be explored to elucidate the development of sustainable biotechnological applications of the system.
Trichoderma
Trichoderma
spp. as a biocontrol agent
spp.as commercial mycofungi-
Trichoderma
species on soil–plant
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