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Medicinal Plants – Chemical, Biochemical, and Pharmacological Approaches
351
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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]

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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]

Medicinal Plants – Chemical, Biochemical, and Pharmacological Approaches
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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.

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olive is sustained with the results obtained by [86] who reported that among 20
356
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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

Medicinal Plants – Chemical, Biochemical, and Pharmacological Approaches
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