Guzmán-Guzmán et al.

BMC Microbiology (2025) 25:434 BMC Microbiology

[Link]

REVIEW Open Access

Trichoderma: a multifunctional agent in plant

health and microbiome interactions
Paulina Guzmán-Guzmán1, Hassan Etesami2 and Gustavo Santoyo1*

Abstract
The beneficial fungus Trichoderma is a key component of agricultural soils, contributing to sustainable crop
production through multiple mechanisms. Among its major roles are the suppression of plant pathogens,
promotion of plant growth, and activation of plant immune responses. This study reviews recent advances in
understanding the modes of action of Trichoderma spp. related to pathogen control and plant growth promotion,
using comparative analysis of its interactions across different plant-associated habitats. In recent years, considerable
progress has been made in elucidating how Trichoderma interacts with plants in the rhizosphere, endosphere, and
phyllosphere, where it exhibits distinct colonization patterns and functional traits. Additionally, this review explores
emerging but less-studied topics, such as the involvement of Trichoderma in the pathobiome concept. Finally, we
discuss the synergistic interactions between Trichoderma and other plant-associated microorganisms, highlighting
their importance in shaping complex microbial networks within agroecosystems.
Keywords Plant microbiome, Biocontrol, Bioformulations, PGPR

Introduction plant in a well-defined niche with specific physiochemi-

Plants worldwide continuously face biotic and abiotic cal properties, collectively forming what is known as the
stressors such as pathogens, drought, nutrient deficien- plant microbiome [18].
cies, and extreme environmental conditions [19, 79]. As The plant microbiome plays a crucial role in help-
sessile organisms, plants rely heavily on beneficial inter- ing plants cope with abiotic stresses, such as drought,
actions with microorganisms to enhance nutrient uptake salinity, heavy metal exposure, and nutrient acquisition,
and strengthen their defense against these challenges while also enhancing overall health, pathogen resistance,
[146]. In return, plants support soil microbial commu- and productivity [82]. This concept underpins efforts to
nities by secreting a portion of their fixed carbon and manipulate or engineer the plant microbiome as a poten-
nitrogen [117] and providing habitats, such as the rhi- tial biotechnological tool for sustainable agriculture
zosphere, plant endosphere, and phyllosphere [92]. This aimed at boosting crop productivity and reducing plant
unique microbial community, or microbiota, inhabits the disease [139]. For example, in a study conducted by Liu
et al. [74], more than 1,900 fungal operational taxonomic
units (OTUs) and another 2,600 bacterial OTUs were
*Correspondence: identified within the endosphere of banana plants. The
Gustavo Santoyo authors compared the diversity and abundance of these
[Link]@[Link]
1
Institute of Biological and Chemical Research, Universidad Michoacana bacterial and fungal OTUs in root tissues and shoot tips,
de San Nicolás de Hidalgo, Ciudad Universitaria, Morelia, both in healthy plants and in those affected by wilting
Michoacán 58030, México disease caused by Fusarium oxysporum f. sp. cubense.
2
Department of Soil Science, College of Agriculture and Natural
Resources, University of Tehran, Tehran 77871-31587, Iran The most abundant bacterial genera were Caulobacter

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Guzmán-Guzmán et al. BMC Microbiology (2025) 25:434 Page 2 of 17

and Paracoccus, while Cladosporium and Eurotium were are fast-growing fungi with hyaline phialides, septate
predominant among the fungi. Interestingly, three endo- hyphae, and conidiophores that often form tree-like
phytic strains were isolated and engineered to express shapes; their mature conidia are typically yellow to green
1-aminocyclopropane-1-carboxylate (ACC) deaminase in color [21, 59]. Many Trichoderma species are widely
activity on their cell walls: Enterobacter sp. E5, Kosako- used in agriculture because of their beneficial interac-
nia sp. S1, and Klebsiella sp. Kb. Inoculation of banana tions with plants and ability to antagonize various plant
plants with these bacteria enhanced resistance to wilting pathogens [35, 85, 135], making these fungi valuable bio-
and promoted plant growth compared to non-inoculated technological tools for sustainable agriculture.
plants, highlighting the importance of ACC deaminase One of key contributions of Trichoderma in agriculture
activity. is its ability to enhance plant growth and productivity
Diversity within plant microbiomes is substantial, through beneficial plant interactions. Trichoderma pro-
encompassing hundreds to thousands of bacterial and motes plant growth through mechanisms that include
fungal species [62, 88]. For example, the rhizosphere root colonization [134], nutrient solubilization and bio-
microbiome of avocado plants contains 3411 bacterial availability in the soil [133, 138], abiotic stress tolerance
OTUs and 1184 fungal OTUs [129], while 306 bacterial [30, 121], secondary metabolite production [140], and
isolates from the endosphere microbiome of saffron (Cro- the synthesis of plant hormones such as auxins, gibber-
cus sativus) have been identified, with Bacillus being the ellins, and cytokinins [57]. Once Trichoderma colonizes
most abundant genus, exhibiting biocontrol and growth- the root, it establishes a mutualistic interaction, exchang-
promoting traits [5]. The rhizosphere microbiome of the ing secondary metabolites and nutrients with the plant in
five coffee tree species was similarly diverse, predomi- return for carbon sources, such as sugars.
nantly comprising Proteobacteria, Actinobacteria, Asco- For instance, T. viride Tv-1511 colonizes peppermint
mycota, and Mucoromycota [34]. roots, promotes growth, and enhances essential oil pro-
Among these beneficial microorganisms, such as bac- duction [44]. Similarly, T. atroviride colonization in
teria and fungi, Trichoderma spp. have emerged as both wheat, mediated by the glycosyltransferase Taugt17b1,
valuable microbiome components and microbiome mod- enhances plant growth and defense against Rhizoctonia
ulators [34, 50, 62, 148]. Trichoderma has been widely cerealis [26].
recognized as a plant growth promoter and an effective Trichoderma also facilitates the availability of min-
biocontrol agent, benefiting crops such as maize [134], eral nutrients in the soil by solubilizing them, thereby
beans [53], wheat [150], and cotton [126]. improving the soil quality and enhancing plant nutri-
Efforts to manipulate or modify the plant microbiome tion, growth, and productivity. For example, Trichoderma
have enabled researchers to increase the prevalence of koningiopsis can solubilize phosphate under alkaline
beneficial microorganisms and decrease that of patho- and drought conditions, and antagonize Rhizoctonia
gens in agriculturally important plants, with Trichoderma solani [133]. T. harzianum SQR-T037 solubilizes miner-
playing an important role [148, 152]. Given the critical als, such as phosphorus, iron, manganese, copper, and
role that Trichoderma plays in enhancing plant health zinc, promoting the growth and development of tomato
and productivity, understanding its interactions with the plants under nutrient-limited conditions [68]. Differ-
plant microbiome is essential for developing sustainable ent Trichoderma spp. also confer resistance to abiotic
agricultural practices. This review aims to summarize stresses. T. longibrachiatum TG1 and T. harzianum miti-
recent findings on the mechanisms by which Tricho- gate salt stress in wheat and cucumber plants, respec-
derma interacts with plants and their associated micro- tively, whereas T. asperellum helps Solanum lycopersicum
biomes, highlighting its multifaceted contributions to (tomato) resist drought and cold stress [30].
plant growth promotion, pathogen suppression, and In addition, Trichoderma can produce plant hormone-
modulation of microbial communities. By elucidating like compounds, which may explain its plant growth-pro-
these interactions, we sought to provide insights that can moting traits. Illescas et al. [57] found that T. virens T49
inform future research and practical applications, ulti- and T. harzianum T115, which produce gibberellins, aux-
mately contributing to the advancement of agroecosys- ins, and cytokinins, promote growth and reduce drought
tem resilience and sustainability. stress in wheat. However, T. longibrachiatum T68 and T.
spirale T75, which produce these hormones to a lesser
Trichoderma as a plant growth promoter extent, were less effective in promoting growth. Tricho-
Trichoderma spp. are soil-borne fungi belonging to the derma is a prolific producer of secondary metabolites.
phylum Ascomycota and family Hypocreaceae. The Trich- These metabolites serve multiple functions, including
oderma genus comprises over 500 species commonly helping Trichoderma compete for nutrients and space
found in soil, surrounding plant roots, and decayed to inhibit the growth of other microorganisms harmful
organic matter as saprophytes [58]. Trichoderma species to plants, facilitating colonization by modulating plant
Guzmán-Guzmán et al. BMC Microbiology (2025) 25:434 Page 3 of 17

defense responses, and promoting plant productivity. A degrade [35]. This process inhibits the growth of patho-
notable example is the secondary metabolite 6-pentyl- gens and prevents them from causing plant diseases.
α-pyrone (6PP), which enhances plant growth and pro- Studies have shown that Trichoderma is effective
ductivity [23, 33, 61]. Therefore, Trichoderma species are against several fungal pathogens such as Botrytis cinerea
well-suited for enhancing sustainable agriculture by pro- [51], Rhizoctonia solani [48], Fusarium oxysporum [14],
moting plant growth and yield and establishing beneficial Alternaria alternata, Aspergillus flavus [22], and Fusar-
interactions with plant hosts (Fig. 1). ium pseudograminearum [52]. This mycoparasitic activ-
ity in the rhizosphere is crucial because its presence can
Trichoderma as a biocontrol agent directly help manage plant diseases.
Another important and widely recognized trait of Trich- Another important biocontrol trait of Trichoderma
oderma in agriculture is its ability to antagonize plant spp. is antibiosis. During antibiosis, Trichoderma pro-
pathogens through direct and indirect mechanisms, col- duce secondary metabolites that inhibit or limit the
lectively known as its biocontrol capacity [130]. Direct growth of plant pathogens [7, 86]. These compounds
biocontrol mechanisms include antibiosis and mycopara- include gliotoxins from T. virens Q strains, which inhibit
sitism, whereas indirect methods involve competition for Pythium aphanidermatum and prevent damping-off
nutrients and space, as well as activation of plant immune disease in tomato plants [60]. Gliotoxins also damage
defense systems [46, 86]. the hyphal cells of Sclerotium rolfsii, antagonizing this
Trichoderma spp. are often used as biocontrol agents pathogen [54]. Another example is the 6-Pentyl-2H-py-
because of their mycoparasitic abilities. Mycoparasitism ran-2-one (6PP) from T. atroviride, which inhibits Cylin-
involves a series of steps that allow Trichoderma to attack drocarpon destructans [61].
its fungal prey. First, the mycoparasite grows toward its Trichoderma’s antibiotic secondary metabolites also
prey, secreting chitinolytic enzymes to degrade the prey’s inhibit bacterial pathogens such as Xanthomonas ory-
cell wall. Upon reaching its target, Trichoderma attaches zae pv. oryzae, a rice pathogen, is strongly inhibited by
to and coils around the hyphae of prey and begins to Trichokonins A and peptaibols from T. longibrachiatum

Fig. 1 Interactions between Trichoderma and the plant, highlighting the services it provides in conjunction with the host-associated microbiome
Guzmán-Guzmán et al. BMC Microbiology (2025) 25:434 Page 4 of 17

SMF2, which damages bacterial cells [154]. Addition- Trichoderma species, there is a growing interest in com-
ally, T. hamatum FB10 produces secondary metabolites bining Trichoderma with other beneficial microorgan-
with antibacterial effects against Erwinia carotovora and isms, suggesting that it may play a vital role as part of the
Acidovorax avenae, antifungal activity against S. sclerotio- plant microbiome.
rum, R. solani, Alternaria radicina, A. citri, and A. dauci,
and nematicidal effects against M. incognita [12]. Critical roles of Trichoderma in diverse plant-
A significant feature of Trichoderma as a biocon- associated zones
trol agent is its capacity to induce systemic resistance Plants can interact with their associated microbiota
in plants. When a plant encounters a pathogen, Sys- through three main zones: the rhizosphere (the soil zone
temic Acquired Resistance is triggered (SAR), involv- surrounding and influenced by the roots) [25], the endo-
ing salicylic acid– mediated signaling, leading to plant sphere (the interior of plant tissues) [28], and the phyllo-
protection against pathogens; however, when the plant sphere (the surface of leaves and other aerial parts of the
encounters beneficial microbes, such as Trichoderma, plant) [155]. Other zones can also be colonized by endo-
Induced Systemic Resistance is triggered (ISR), which phytic microorganisms, such as the caulosphere (stem),
involves jasmonic acid-and ethylene– mediated signaling anthosphere (flowers), carposphere (fruits), and spermo-
[113]. Induced Systemic Resistance (ISR) primes plants to sphere (seeds) [73, 87]. By colonizing seeds, microbial
respond more quickly and strongly to pathogen attacks, endophytes —including Trichoderma species— can be
and beneficial microorganisms such as Trichoderma vertically transmitted to the next plant generation [43]. In
and plant growth-promoting bacteria (PGPB) can trig- addition, the rhizosphere can be subdivided into the ecto-
ger ISR [90]. For example, T. pubescens Tp21 increases rhizosphere (rhizospheric soil), the rhizoplane (root sur-
the expression of defense-related genes PAL (Phenylala- face), and the endo-rhizosphere (inner root tissue), where
nine Ammonia-Lyase, an enzyme that initiates the phen- plant-Trichoderma interactions can influence plant and
ylpropanoid pathway and contributes to the production fruit growth, development, and health [55, 64, 71, 84].
of antimicrobial compounds), CHS (Chalcone Synthase, In this review, we focus on analyzing studies of Tricho-
a key enzyme in flavonoid biosynthesis involved in anti- derma-plant interactions within these closely connected
oxidant and antimicrobial defenses), and HQT (Hydroxy- regions (rhizosphere, endosphere, and phyllosphere).
cinnamoyl-CoA Quinate Hydroxycinnamoyl Transferase,
responsible for the synthesis of chlorogenic acid, a phe- Trichoderma in the rhizosphere
nolic compound with protective functions) in tomatoes. The rhizosphere is a narrow soil region directly sur-
Additionally, strain Tp21 reduces oxidative stress during rounding plant roots, affected by root exudates, and
R. solani infection, and enhances antioxidant enzyme inhabited by a diverse array of microorganisms associ-
activity such as superoxide dismutase (SOD), catalase ated with plants, known as the “rhizobiome” or “rhizo-
(CAT), polyphenol oxidase (PPO), and peroxidase (POX) sphere microbiome.” [91]. Plant growth-promoting fungi
production, thereby conferring resistance against R. (PGPF) in the rhizosphere, such as Gliocladium spp.,
solani [17]. arbuscular mycorrhizal fungi, and Trichoderma spp., also
Similarly, T. hamatum Th23 protects tomatoes from contribute to increased crop production and plant resis-
Tobacco Mosaic Virus (TMV) by reducing oxidative tance to both abiotic and biotic stress by improving plant
stress, increasing the activities of antioxidant enzymes development and defense systems, mineral solubiliza-
such as CAT, SOD, and PPO, and upregulating genes tion, and pathogen inhibition [4]. Thus, the rhizosphere
involved in the polyphenolic biosynthetic pathway (HQT microbiome plays a crucial role in plant growth, produc-
and CHS) as well as pathogenesis-related proteins (PR- tivity, nutrient acquisition, overall plant health, and soil
1, PR-2, and PR-7), thereby enhancing resistance against quality, thereby enhancing plant resistance to both biotic
TMV [1]. Trichoderma spp. can also induce resistance and abiotic stresses.
against other plant pests, such as insects. When applied The composition of the rhizosphere is determined
to the roots of Lansium domesticum plants, Trichoderma by various factors, including soil properties, plant root
spp. increase salicylic and jasmonic acid concentrations, exudates, plant age or developmental stage, environ-
protecting against infestation by the scale insect Unaspis mental conditions, chemical compounds, and the micro-
mabilis and activating plant defenses [125]. These bio- organisms themselves [129, 151]. Pear fruits treated with
control mechanisms enable Trichoderma to antagonize a 1-methylcyclopropene (1-MCP) to prevent post-harvest
wide range of plant pathogens and protect agriculturally decay caused by pathogenic fungi changed the bacterial
significant crops. Thus, Trichoderma is a valuable tool for and fungal microbiota, decreasing the presence of infect-
sustainable agriculture, serving as an effective alternative ing fungi and improving resistance to post-harvest decay
to chemical fertilizers and pesticides [2, 11]. While much [151]. During flooding stress and elevated CO2 (eCO2)
of the research has focused on using single or multiple conditions, bacterial and fungal communities associated
Guzmán-Guzmán et al. BMC Microbiology (2025) 25:434 Page 5 of 17

with soybean plants increased compared to non-stressed Agri-Biotech Co. Ltd., Beijing, China) helped reduce the
plants, with Trichoderma being among the most abun- disease index of potato common scab in potato plants,
dant fungal genera in the eCO2 and eCO2 + flooding thereby increasing tuber yield [143].
treatments [27], indicating that abiotic stress factors can Adedayo and Babalola [4] reviewed the plant growth-
induce significant shifts in plant microbiome structure. promoting mechanisms of PGPF, including Trichoderma
Root exudates from stressed tomato plants exposed spp., highlighting their significance as bioagents to
to either the foliar pathogen B. cinerea or abiotic stress, enhance crop yields and resistance to abiotic and biotic
such as salinity or wounding, attracted T. harzianum stress, and demonstrating the key role of PGPF, such as
more strongly than non-stressed plants. Furthermore, Trichoderma in maintaining healthy plant environments.
tomato and cucumber plants infected with B. cinerea Table 1 presents some of the most recent studies involv-
exhibited increased T. harzianum growth when inocu- ing Trichoderma strains isolated from, or associated with,
lated near the roots [77], showing that changes in root the rhizosphere.
exudates due to plant stress can attract beneficial soil
microbes, such as Trichoderma. Therefore, any change in Trichoderma in the endosphere
rhizobiome composition can potentially alter its ecologi- Endophytic microorganisms inhabit plant tissues such
cal functions, affecting the overall health of the plant and as roots, stems, leaves, flowers, fruits, and seeds with-
rhizosphere [129]. out causing apparent damage. In this hidden world, they
Trichoderma is present in the root microbiome of can interact with their plant hosts, aiding in their defense
various agriculturally important plants. For instance, against pathogens, stimulating growth, and increasing
Trichoderma has been found in the rhizosphere of cot- production. Among these endophytic microorganisms,
ton plants that are resistant to Verticillium wilt [146]. de we found various fungal species belonging to the genera
Sousa et al. [34] analyzed the microbiome of five coffee Piriformospora, Penicillium, Beauveria, Serendipita, and
species (Coffea arabica, C. canephora, C. stenophylla, C. Glomus. Additionally, nonpathogenic strains have been
racemosa, and C. liberica) and found several bacterial reported as endophytes from genera such as Colletotri-
genera, including Streptomyces, Bradyrhizobium, and chum, Phoma, Fusarium, and Alternaria [15, 98]. In the
Mycobacterium, as well as fungal species, such as Rhi- case of Trichoderma, species have also been reported
zophagus, Fusarium, and Trichoderma among the most living as endophytes in crops, such as T. harzianum, T.
abundant. Olowe et al. [89] isolated seven Trichoderma viride, T. asperellum, to name a few [114].
strains from the rhizospheres of maize, banana, and cas- A recent example of the beneficial effects of endophytic
sava plants. All isolates showed antagonistic capacity strains of Trichoderma was recently published Hasan et al.
against two Fusarium isolates, suggesting the potential [49]. In this study, two strains of Trichoderma spp., ReTk1
of rhizospheric Trichoderma as a biocontrol agent and and ReTv2 were isolated, and their potential for promoting
its presence in maize rhizosphere. Microorganisms in plant growth and biocontrol of clubroot disease caused by
the plant rhizosphere mediate nutrient acquisition under the pathogen Plasmodiophora brassicae in rapeseed plants
stressful conditions. Under nutrient deprivation, plants (Brassica napus L.) was evaluated. Rapeseed is one of the
secrete secondary metabolites into the soil, signaling rhi- most important vegetable oil crops in the world. These two
zospheric microorganisms to make nutrients available to endophytic strains promoted plant growth by increasing
the plant and thereby indirectly promoting growth [93]. parameters such as shoot and root length, leaf diameter,
Trichoderma was no exception: co-inoculation of four and total biomass production. Furthermore, both the ReTk1
T. asperellum isolates and B. subtilis in the rhizosphere and ReTv2 strains showed relevant results in inhibiting the
of Marandu grass (Urochloa brizantha cv. Marandu) germination of resting spores of P. brassicae in root exudate
improved nitrogen, potassium, and phosphorus uptake experiments, indicating a possible mechanism of action.
and promoted plant growth [32]. Inoculation with T. Additionally, the authors reported that both endophytic
viride alone or with Azotobacter chroococcum increased strains stimulated the expression of defense-related mark-
soil micro- and macronutrients, enhancing the growth ers involved in the jasmonate (BnOPR2), ethylene (BnACO
of cotton and wheat [138]. As a rhizosphere inhabitant, and BnSAM3), auxin (BnAAO1), and salicylic acid (BnPR2)
Trichoderma uses biocontrol and plant growth-promot- pathways, demonstrating the excellent potential of these
ing traits to support plant health. For example, seven strains with multiple benefits for their plant host, Brassica
rhizospheric Trichoderma isolates, including T. virens, T. napus L.
harzianum, T. erinaceum, and T. koningiopsis, inhibit the Another study reported Rajani et al. [105] that endo-
growth of Fusarium proliferatum, a plant pathogen [89]. phytic strains of Trichoderma were able to restrict the
A microbial consortium of T. harzianum and B. subtilis growth of pathogens, such as Sclerotinia sclerotiorum,
(a commercial product called Micro-ecological Agents Sclerotium rolfsii, and Fusarium oxysporum, where
for Controlling Continuous Crop Diseases; Sino Green mechanisms such as mycoparasitism and the production
Table 1 Example studies on rhizospheric Trichoderma species with biological control action against various pathogens and benefited crop plants
Trichoderma species/strain Origin Biocontrolled pathogen(s)/Disease Beneficiated crop Mechanisms of action References
T. harzianum MC2 and NBG Rhizosphere of tomato, Suppressed bacterial wilt caused by Tomato (Solanum Antagonistic activity, promotion of plant growth, produc- [104]
brinjal, and chili Ralstonia solanacearum lycopersicum) tion of secondary metabolites
T. asperellum TaspHu1 Rhizosphere soil of Juglans Enhanced growth, resistance to Alternaria Tomato (Solanum Stress tolerance, increased enzyme activity, enhanced [149]
mandshurica alternata lycopersicum) nitrogen absorption
T. atroviride C52 Soil with Sclerotium Controlled onion root rot Onion (Allium cepa) Rhizosphere competence, formulation-dependent [80]
cepivorum effectiveness
T. virens Tomato plant rhizosphere Inhibited growth of rhizosphere bacteria Tomato (Solanum Secreted proteins and metabolites inhibit bacteria, volatile [69]
lycopersicum) compounds affect interactions
T. pseudoharzianum T1 Rhizosphere soil of Syringa Biocontrol of Microsphaera Syringa oblata Induction of ABA production, increase in catalase activity [76]
Guzmán-Guzmán et al. BMC Microbiology

oblata syringejaponicae
T. viride Soybean rhizosphere Reduced disease index in soybean root Soybean (Glycine max) Altered microbial community, increased diversity, and [39]
rot reduced relative abundance of pathogenic fungi
T. asperellum 7316 Castor plant roots Limiting Fusarium oxysporum infection Castor (Ricinus com- High rhizosphere competence, mycoparasitic ability, [100]
munis L.) enhanced plant growth
T. asperelloides, T. koningiopsis Rhizosphere of Ilex Controlled Fusarium oxysporum and Ilex paraguariensis Inhibited mycelial growth, increased oxidative enzymes [103]
(2025) 25:434

paraguariensis Fusarium solani

T. virens T1-02 Rhizosphere of flamingo Controlled flower blight caused by Anthurium andraeanum Production of volatile antifungal compounds, direct [10]
flower Neopestalotiopsis clavispora parasitism
Trichoderma asperellum (TR1, Subabul rhizospheric soil Effective against Fusarium equiseti Subabul (Leucaena Production of volatile metabolites, siderophores, nutrient [95]
TR2, TR3, TR6) (70–90% inhibition) leucocephala) solubilization (P and K)
Trichoderma hamatum (TR4,
TR5)
Trichoderma atroviride Red pepper rhizosphere Effective against Colletotrichum acutatum Red pepper (Capsicum Production of volatile organic compounds (VOCs), inhibit- [63]
(ATR697) annuum) ing mycelial growth, resistance to chemical fungicides
Trichoderma longibrachiatum
(LON701)
Trichoderma gamsii (RH4) Rhizosphere of black gram Protective against Fusarium oxysporum Black gram (Vigna Lytic properties, enzyme production, growth promotion [137]
(Vigna mungo) mungo)
Trichoderma virens Tomato rhizosphere Enhanced growth, suppresses Fusarium Tomato (Solanum Volatile compounds affecting pathogen growth [14]
oxysporum lycopersicum)
Trichoderma harzianum Rhizosphere of Egyp- Antagonistic against crown rot pathogens Egyptian clover Upregulation of defense enzymes, seed treatment, foliar [20]
(TBR-7) tian clover (Trifolium spray
alexandrinum)
Trichoderma asperellum, Rubber tree rhizosphere Inhibitory against Rigidoporus microporus Rubber tree (Hevea Production of antifungal metabolites, hydrolytic enzymes [41]
Trichoderma spirale brasiliensis)
Trichoderma spp. Soil from diverse regions Control of Rhizoctonia solani Strawberry Production of volatile metabolites, enhancing plant [81]
of Lorestan Province, Iran (Fragaria × ananassa) defense mechanisms
Trichoderma virens, Rhizospheric of banana in Biocontrol against Fusarium oxysporum f. Banana (Musa Diversity in Trichoderma species correlates with soil [31]
Trichoderma harzianum Tenerife sp. cubense acuminata) chemistry
Trichoderma spp. (PBT3, PBT4, Various rhizospheric soils Antagonistic against Fusarium oxysporum Chickpea (Cicer Production of antifungal metabolites, cell-wall degrading [66]
PBT9, PBT13) [Link]. ciceri and Sclerotium rolfsii arietinum) enzymes
Page 6 of 17
Guzmán-Guzmán et al. BMC Microbiology (2025) 25:434 Page 7 of 17

of volatile organic compounds are relevant in antago- pathogens. However, these endophytes have tolerance
nism. Similarly, the volatiles produced by the endophyte mechanisms, either by degrading or modifying their
T. asperelloides PSU-P1 demonstrated their function as structure to mitigate their “antimicrobial” effect [8, 120].
antagonists of various pathogens, such as Colletotrichum Table 2 shows recent studies involving endophytic Trich-
sp., Corynespora cassiicola, Curvularia lunata, Gano- oderma strains, as well as some phyllospheric strains, and
derma sp., Penicillium oxalicum, Neopestalotiopsis clav- their biocontrol roles against multiple plant pathogens.
ispora, S. rolfsii, and Stagonosporopsis cucurbitacearum.
Additionally, volatiles such as 2-methyl-1-butanol, Trichoderma in the phyllosphere
2-pentylfuran, acetic acid, and 6-pentyl-2H-pyran-2-one According to Vorholt [141], the surface area of the phyl-
(6-PP) stimulate growth and defense responses in Arabi- losphere is approximately twice the size of the Earth’s
dopsis thaliana plants [97]. surface, providing a vast habitat for a wide variety of
In recent years, there has been an effort to find genetic microorganisms. Naturally, the conditions in the soil or
or genomic differences that result in intrinsic characteris- within plant tissues differ greatly from those encoun-
tics that differentiate Trichoderma strains capable of liv- tered by organisms living as epiphytes on plant surfaces;
ing as endophytes from those that do not. The rationale therefore, they must possess innate colonization abilities
is based on studies showing that plants produce metabo- to adapt and survive in such environments [16]. In fact,
lites, such as flavonoids, phenolics, and terpenes, which the phyllosphere can be a hostile environment, subject to
have defensive functions, and that endophytes must have continuous environmental fluctuations and exposure to
tolerance/resistance mechanisms against these plant UV radiation, sunlight, rain, ice, and more. As a result,
compounds. Therefore, only those species of Tricho- microbial densities are usually lower compared to other
derma with these capabilities will be able to colonize the ecosystems such as the rhizosphere or bulk soil [65, 145].
interior space of a plant, including vascular tissues, with- The topology of plant leaves, as well as their age and leaf
out causing damage or colonizing various aerial and root side (adaxial and abaxial), are important factors in estab-
parts. lishing the structure and diversity of microbial communi-
Thus Scott et al. [120], through a comparative genomic ties in the phyllosphere [40, 70, 128]. In agroecosystems,
analysis of more than 38 genomes from the metabolic the phyllosphere can also be sprayed with agrochemicals
repertoire of endophytic and non-endophytic Tricho- that may be bactericidal or fungicidal, thereby altering
derma strains, we found greater diversity in biosynthetic both pathogenic and beneficial microbial communities
gene clusters (BGC) and degradative gene clusters (DGC) [147]. The same occurs with bioinoculants, which, when
in the endophytic strains. Although variations in BGCs inoculated in the phyllosphere, can alter the microbiota
and DGCs might be attributed to species differences residing there [102]. This is also the case when Tricho-
rather than lifestyle, genome comparisons should be con- derma harzianum (strain T22) was inoculated in the
ducted within a single species using isolates with distinct strawberry phyllosphere, as fungal communities were
lifestyles to confirm this. However, this is an effort to dis- altered but bacterial communities were not [132], sug-
cern intrinsic differences between endophytic strains and gesting some specific interactions between these fun-
those that are not, for example, may only colonize regions gal communities and Trichoderma that require further
such as the rhizosphere or phyllosphere, where plant investigation.
metabolites may not be “toxic or restrictive” for coloniza- In the case of Trichoderma, this mycoparasitic fun-
tion by these Trichoderma strains. Years before, a simi- gus has also been reported as a phyllosphere inhabitant,
lar study attempting to discern the genetic capabilities of although the number of studies reporting its isolation,
endophytism, but in endophytic bacteria versus rhizo- presence, or characterization in this habitat is lower com-
spheric ones, was conducted by Ali et al. [8], comparing pared to rhizospheric or endophytic isolates (Fig. 2). This
the genomes of nine plant growth-promoting endophytic research bias has contributed to a greater understand-
bacteria (PGPB), including genera such as Azoarcus, Azo- ing of Trichoderma as a rhizospheric antagonist [37]. In
spirillum, Burkholderia, Enterobacter, Klebsiella, Pseu- contrast, studies on its interactions within other specific
domonas, and Serratia. From this comparative analysis, plant zones, such as the endosphere and phyllosphere,
genes with potential roles in endophytic colonization remain scarce and remain poorly understood and require
were identified, such as transporter proteins, secretion more focused investigation [29].
and delivery systems, plant polymer degradation/modifi- To address this, Saxena et al. [119] we evaluated and
cation, detoxification, and redox potential maintenance. compared the biocontrol capabilities of a phyllospheric
As can be seen in the studies mentioned above, some Trichoderma strain (BHUF4) and a rhizospheric strain
functions are related to resistance mechanisms to the (T16A) against the pathogen Colletotrichum capsici,
endophytic environment, where the plant, either consti- which causes anthracnose disease in chili (Capsicum
tutively or induced, produces defense compounds against annuum L.). The results showed that the phyllosphere
Table 2 Key studies on endophytic and phyllospheric Trichoderma species with biocontrol activity against various pathogens
Trichoderma species/strain Origin Biocontrolled pathogen(s)/Disease Beneficiated crop Mechanisms of action References
T. harzianum KUFA0436, Endophytic Phytophthora palmivora (Phytophthora leaf fall) Rubber (Hevea Fast growth, competition for resources, antifungal crude [127]
KUFA0437 brasiliensis) extract production
T. longibrachiatum MD33 Endophytic None (focus on enhancing dendrobine production) Dendrobium nobile Enhancement of secondary metabolite production [115]
(dendrobine)
T. asperellum KUFA0702, Endophytic Exserohilum turcicum (Northern leaf blight) Corn (Zea mays) Fast growth, nutrient competition, antifungal crude extract [75]
KUFA0703, T. harzianum production
Guzmán-Guzmán et al. BMC Microbiology

KUFA0710, KUFA0713
T. longibrachiatum WKA55 Endophytic Mycotoxinogenic fungi Peanut (Arachis Enzymatic activity (cellulase, protease, polygalacturonase), [6]
hypogaea) citric acid production
T. erinaceum Endophytic Pythium ultimum Common bean Mycelial growth inhibition through new alkene and known [124]
(Phaseolus vulgaris) bioactive compounds
T. asperellum T-AS2, T-AS7, T. Endophytic Phytophthora cinnamomi (Avocado root rot) Avocado (Persea Competition, resource exclusion, inhibition of pathogen [9]
(2025) 25:434

harzianum T-H5 americana) spread

T. longibrachiatum MD33 Endophytic None (focus on producing dendrobine) Dendrobium nobile Production of bioactive compounds, antibacterial activity [116]
Trichoderma sp. T154 Endophytic Phaeoacremonium minimum (Grapevine trunk disease, Grapevine (Vitis Spore adhesion, niche exclusion, hyphal coiling [24]
esca) vinifera)
Trichoderma gamsii T6085 Endophytic Fusarium head blight (FHB) Wheat (Triticum Induction of defense genes: pal1, pr1, pgip2 and lox1 genes [108]
aestivum’Apogee’)
Trichoderma zelobreve T20 Endophytic Cytospora cincta and Neoscytalidium dimidia- Apple trees Not specified [107]
tum (apple canker)
Trichoderma asperellumstrain Endophytic Fusarium proliferatum f. sp. malus domestica MR5 Apple replant dis- Protease, amylase, cellulase, and laccase activities [144]
6S-2 ease (ARD)
T. asperelloides PSU-P1 Endophytic Colletotrichum sp. (Leaf blight), Corynespora cassiicola A. thaliana Volatiles: 2-methyl-1-butanol, 2-pentylfuran, acetic acid, and [97]
(Leaf spot), Curvularia lunata (Leaf spot), Ganoderma 6-pentyl-2H-pyran-2-one (6-PP)
sp. (Basal stem rot), Penicillium oxalicum(Blue mold),
Neopestalotiopsis clavispora (Flower blight), Sclerotium
rolfsii (Southern blight), Stagonosporopsis cucurbita-
cearum (Gummy stem blight)
Nineteen Trichoderma strains Endophytic Colletotrichum truncatum, Lasiodiplodia theobro- Not specified Antibiosis, together with multiple mechanisms [83]
mae, Macrophomina phaseolina, and Sclerotium
delphinii
Trichoderma sp. strain BHUF4 Phyllospheric Colletotrichum capsici Chili (Capsicum an- Stimulation of antioxidant enzymes; reduced anthracnose [119]
nuum L.) lesions in chili leaves
Trichoderma sp. strain BHUF4 Phyllospheric Colletotrichum truncatum Chili (Capsicum an- Stimulation of the defense responses [118]
nuum L.)
Page 8 of 17
Guzmán-Guzmán et al. BMC Microbiology (2025) 25:434 Page 9 of 17

Fig. 2 Number of documents retrieved from the Scopus database using keywords such as “Trichoderma” AND either “rhizosphere,”“endosphere,” or “phyl-
losphere/epiphyte” ([Link] This illustrates the significant disparity in research attention, with phyllospheric Trichoderma being notably
less studied compared to isolates from endosphere and rhizosphere ecosystems

isolate stimulated defense networks in plants, where demonstrated good colonization and survival capacity in
enzymes such as phenylalanine ammonia-lyase (PAL), the phyllosphere of M. charantia plants, from which they
peroxidase (PO), polyphenol oxidase (PPO), superox- showed a beneficial interaction with their host.
ide dismutase (SOD), and total phenolic content (TPC) In another recent study, the influence of spore concen-
were significantly increased. It also significantly reduced tration on the phyllosphere colonization capacity of a
anthracnose lesions in chili leaves compared to the rhi- Trichoderma consortium in Passiflora caerulea was eval-
zospheric strain, while promoting plant growth. Later uated. The authors reported that morphological, physio-
studies by the same research group demonstrated that logical, and ultrastructural characteristics, as well as fruit
the phyllospheric Trichoderma strain BHUF4 stimu- yield and quality, were improved by these Trichoderma
lated the systemic acquired resistance (SAR) pathway, strains [122]. The inoculated consortium consisted of the
whereas the rhizospheric Trichoderma strain T16A used same two strains reported in previous studies: T. asperel-
the induced systemic response (ISR) pathway to elicit lum T36b and T. harzianum Td50b.
defence responses in the host plant Capsicum annuum L. Due to their sessile nature, plants are subjected to inter-
under C. truncatum inoculation [118]. actions with microorganisms, including beneficial fungi
Recently, the use and application of chlamydospores such as Trichoderma, especially in environments such as
from T. asperellum T36 and T. harzianum Td50b were the phyllosphere [16]. Therefore, it is advisable to further
evaluated through foliar spraying to improve the physi- investigate the succession of phyllospheric Trichoderma
ology and yield of Momordica charantia. M. charan- communities and their influence on plant growth, devel-
tia is a plant from the Cucurbitaceae family, native to opment, and health. It is currently unknown whether
tropical and subtropical regions, with extensive culinary there are specific interactions with certain plant hosts and
and medicinal uses owing to its production of second- whether these are of agro-food importance. Just as some
ary metabolites. The results showed that the chlamydo- interactions occur in the rhizosphere, where root exu-
spore suspension significantly improved physiological dates influence specific interactions to recruit a benefi-
parameters, polyphenol and flavonoid content, and anti- cial microbiome, it is unknown what type of compounds,
oxidant activity in both leaves and fruits. An increase whether diffusible or volatile organic compounds, attract
in yield from 25.33% to 53.07% was also observed [13]. Trichoderma strains and promote their recruitment.
These results suggest that the Trichoderma strains
Guzmán-Guzmán et al. BMC Microbiology (2025) 25:434 Page 10 of 17

Interplay between Trichoderma and plant- harzianum T-78 [94, 131]. This suggests that beneficial
associated microorganisms microorganisms capable of inducing changes in root exu-
Microorganisms do not exist in isolation within their dates may help plants to endure environmental stress.
environments, and Trichoderma not only interacts with Because plants modulate their associated microbiomes,
its plant or fungal hosts but also with other beneficial microorganisms can influence each other. Several studies
microorganisms such as plant growth-promoting bacte- have found that the presence of Trichoderma in the rhi-
ria (PGPB) or PGPF-like mycorrhiza [96]. This interplay, zosphere can alter microbiota composition, favoring the
or crosstalk, between microbiome organisms and plants presence of plant-beneficial microorganisms.
is complex, with Trichoderma engaging in interactions The inoculation of T. harzianum LTR-2 in the cabbage
and communication with other plant-associated organ- rhizosphere increased the abundance of beneficial bacte-
isms. Microorganisms use various molecules for commu- ria, such as Pseudomonas and Bacillus, conferring resis-
nication, including secondary metabolites and effector tance to clubroot disease, compared to non-inoculated
proteins [96]. This communication is essential among rhizospheres [72]. Seed coatings of T. guizhouense on
microorganisms within the plant microbiome, shaping watermelon and maize modified the microbial commu-
their relationships with both plants and each other. nity of the rhizosphere, correlating with improved seed
A study on metabolites, volatile compounds, and pro- germination and plant growth and promoting benefi-
teins produced by T. virens and T. harzianum Li et al. [69] cial microbes such as Trichoderma spp. and Mortierella
demonstrated that these compounds mediate interac- [148].
tions with different rhizospheric bacteria, including phyla The presence of T. viride affected microbial diversity in
such as Firmicutes, Alpha-proteobacteria, and Gamma- the soybean rhizosphere, with Planctomycetes, Patesci-
proteobacteria, affecting bacterial growth to varying bacteria, and Myxococcota being the most abundant bac-
extents. Additionally, compounds secreted from T. atro- terial phyla, whereas Ascomycota and Mortierellomycota
viride were found to inhibit the growth of PGPB, such as were the predominant fungal phyla. These changes were
Bacillus velezensis AF12 and Bacillus halotolerans AF23, correlated with a reduction in soybean root rot disease
but did not affect the growth of Pseudomonas fluorescens [39].
UM270 [47]. In tomato plants, inoculation with T. virens influenced
Effector molecules from Trichoderma have been stud- the relative abundance of fungal and bacterial phyla and
ied as key communication molecules with their hosts genera associated with the plant, promoting tomato
that modulate the nature of these interactions [106, growth, especially when applied at transplant, suggest-
112]. Trichoderma effectors are involved in mycopara- ing that inoculation methods can affect interactions with
sitic interactions with plant pathogens such as R. solani, plants and other microorganisms [14]. The dual inocu-
B. cinerea, S. cepivorum, and C. lindemuthianum [45, lation of T. viride and dark septate fungi effectively pro-
112], as well as in beneficial interactions with plants that tected Astragalus mongholicus plants against drought,
induce resistance and promote growth [110]. Effector- enriching rhizosphere soil fungi, including Stachybotrys
coding genes from Trichoderma also respond to PGPBs, and Trichoderma, with being Ascomycota the main fun-
which are commonly found in the rhizosphere [47]. gal phylum, compared to non-inoculated plants [50].
Thus, secondary metabolites and effector molecules In watermelon, inoculation with T. asperellum M45a
of Trichoderma play a role in communicating with other reduced fungal diversity while increasing beneficial bac-
beneficial microorganisms, potentially influencing their teria such as Pseudomonas, enhancing resistance to F.
interactions. This suggests that as part of the plant micro- oxysporum f. sp. niveum wilt disease, and promoting
biome, Trichoderma utilizes its molecular arsenal to plant growth [153]. Umadevi et al. [136] showed that
communicate and interact with microbial communities the presence of T. harzianum in the rhizosphere of Piper
associated with plants. nigrum plants altered the microbial community and its
functional dynamics, selectively recruiting beneficial
Trichoderma as a key player in plant microbiome microbial communities, as indicated by a reduction in
dynamics pathogenicity islands and an increase in plant growth-
Plants exert strong modulatory effects on soil micro- promoting fungi.
biomes [56]. Root exudates influence the assembly of
microorganisms that are associated with the root micro- Trichoderma and the pathobiome concept
biome. For example, coumarins in the root exudates The term pathobiome was recently used to refer to a group
of Arabidopsis thaliana shape the plant microbiota to of associated microorganisms, as well as their interactions
promote health by activating the MYB72 transcrip- with the plant causing an infection, and consequently,
tion factor, which is induced in the presence of benefi- symptoms of a disease develop [78]. This concept of the
cial microorganisms, such as T. asperellum T-34 and T. pathobiome has previously been employed to define
Guzmán-Guzmán et al. BMC Microbiology (2025) 25:434 Page 11 of 17

Fig. 3 General overview of the pathobiome concept. Under optimal growth conditions, a healthy plant is associated with its beneficial microbiome, or
symbiome—the latter referring to the total assemblage of associated organisms, excluding the plant itself. Over time, pathogen proliferation can disrupt
this balance, leading to the formation of a pathobiome. In this state, interactions among the plant, its microbiome, and pathogens result in infection and
disease. Such a disruption mirrors dysbiosis observed in the human gut microbiome [142]

dysbiosis in human hosts and in studies on gut microbiota. Droby et al. [36] recently reported that the pathobiome
In the case of plants, this concept has gained relevance concept can be applied to postharvest diseases, where
because of its similarities to the human microbiome and changes in the microbiome of fruits and vegetables may
its role in disease development. Therefore, it can be said also be responsible for the onset of diseases. In the case
that the pathobiome arises from the interactions generated of fruits and vegetables, damage such as wounds could be
between the plant, the associated microbiome, and poten- a factor that creates a condition in which the pathobiome
tial pathogens [38]. Under this concept, what role does emerges, and multiple species of pathogens “take advan-
Trichoderma play? How can mycoparasitic fungi control tage” of the availability of resources. Through coopera-
the emergence of a pathobiome? These are questions that tion and colonization, they infect and damage produce.
still lack clear answers, although Trichoderma has been The authors mentioned some examples of postharvest
widely studied as a biocontrol fungus of single-pathogen diseases involving multispecies causal agents, such as
diseases, its role as a regulator of interactions within the banana crown rot, apple diseases originating from latent
pathobiome has not yet been clearly studied. infections by fungal pathogens such as Colletotrichum
Within this concept, Trichoderma could play a signifi- spp. and Neofabraea spp., as well as stem-end rots in
cant role in reducing pathogen proliferation in ecosys- fruits such as mango and avocado.
tems such as the rhizosphere, ultimately affecting the In this context, Trichoderma has been applied as a
assembly of several pathogens that could lead to a state biocontrol agent against postharvest diseases of various
of infection in the plant host. The disease state caused by fruits and vegetables [71], and its role as a biocontrol
multiple species has been reviewed in Abdullah et al. [3], agent for multiple species should continue to be investi-
where they analyzed studies of co-infection in susceptible gated, especially as a regulator of interactions within the
plant hosts and proposed some pathogen interactions pathobiome, as previously mentioned. Figure 3 shows the
that enable infection. Some of these include competition pathobiome concept and how the microbiome, includ-
for physical space and resources, cooperation between ing Trichoderma (and other PGPB, beneficial fungi, etc.),
pathogens, where they beneficially interact through could potentially regulate the proliferation of pathogens
chemical signals, and functional complementation via the through different mechanisms of action. However, the
exchange of resources necessary for survival in the envi- role of Trichoderma in modulating microbiome inter-
ronment. Finally, niche specialization allows the coexis- actions remains unknown. One possibility could be the
tence of pathogens, which can lead to greater infection production of effector-type proteins, which would not
and disease development. only regulate plant metabolism but also affect the effector
Guzmán-Guzmán et al. BMC Microbiology (2025) 25:434 Page 12 of 17

proteins produced by pathogens to infect the plant host microbial communities that are part of the plant micro-
[45]. At the same time, it induces the plant’s immune biome. Its presence as part of plant-associated microbes
system (ISR) and defense mechanisms, making it less is beneficial, demonstrating its potential as a biocontrol
susceptible to the pathogen. However, this hypothesis agent and biostimulant for improved and sustainable
requires further investigation. agriculture. In fact, there are studies in which Arabidopsis
plants have been propagated under sterile conditions and
Perspectives of Trichoderma research inoculated with a synthetic microbiome, clearly show-
The presence of beneficial microorganisms in the plant ing the effects of such inoculation. These studies have
microbiome is crucial for the development of new bio- revealed how complex the environment and the interac-
control agents and promoters of plant growth. An tions between different players can be in a changing envi-
emerging agricultural biotechnology is microbiome engi- ronment, but the mechanisms governing such dynamics
neering, which involves the modulation or manipulation are beginning to be uncovered [142]. Under these con-
of the plant microbiome to harness its benefits in improv- ditions, Trichoderma may play an important role in
ing plant production and resistance to environmental modulating the plant microbiome, recruiting beneficial
stress factors [92, 109, 139]. Thus, manipulating the plant microorganisms, and improving soil quality. Therefore,
microbiome using plant-associated microorganisms such all these interactions are fundamental for achieving con-
as Trichoderma spp. could be of great importance for sistent and positive results under open-field conditions,
enhancing sustainable agricultural techniques. which will help mitigate unpredictable abiotic factors.
A four-species synthetic community (SynCom), Ste-
notrophomonas sp., Rhizobium sp., Advenella sp., and Conclusions
Ochrobactrum sp., selected from the rhizosphere of A. In summary, Trichoderma spp. are pivotal components of
mongholicus plants infected with Fusarium oxysporum, the plant microbiome, significantly contributing to sus-
was effective at reducing root rot disease [67]. Gonçalves tainable agricultural practices through their roles as plant
et al. [42] designed a microbial community associated growth promoters and biocontrol agents. Their multi-
with Vellozia epidendroides and Barbacenia macran- faceted interactions with plants and associated micro-
tha plants to fulfill specific microbiome functions, such organisms enhance nutrient uptake, improve resistance
as nitrogen fixation, phosphorus solubilization, and sid- to abiotic stress, and suppress pathogen proliferation.
erophore production, to better understand and enhance The ability of Trichoderma to modulate the dynam-
plant–microbe interactions. Shi et al. [123] constructed ics of the plant microbiome underscores its potential
18 SynComs based on their ability to antagonize Fusar- as a biotechnological tool to improve crop productivity
ium spp. that cause potato dry rot disease, showing and resilience. This review highlights the importance of
promising results as biocontrol agents. understanding the mechanisms underlying Trichoderma
Synthetic communities designed to control Fusar- interactions in various plant-associated environments,
ium wilt in banana plants were effective against the such as the rhizosphere, endosphere, and phyllo-
pathogen using a combination of various beneficial sphere, as well as its role in the emerging concept of the
microorganisms, including Trichoderma and plant pathobiome.
growth-promoting bacteria (PGPB) [101]. This demon- Despite progress made in understanding Tricho-
strates the biocontrol potential of modulating the diver- derma interactions within the plant microbiome, several
sity of microorganisms associated with plants of interest. research gaps remain. Future studies should focus on elu-
T. simmonsii, in combination with Serendipita indica, cidating the molecular mechanisms underlying the ben-
showed biocontrol capacity against Phytophthora cap- eficial effects of Trichoderma, particularly in relation to
sici on bell pepper plants [111], whereas a consortium its interactions with diverse plant species under varying
formed by T. harzianum and Bacillus subtilis improved environmental conditions. Additionally, exploring the
potato yield and increased plant resistance against com- genetic and genomic differences between endophytic
mon scab disease [143]. and non-endophytic Trichoderma strains could provide
Therefore, modulating microbial communities with insight into their ecological roles and adaptability.
Trichoderma could be a part of agricultural management There is a pressing need for research on the synergis-
techniques aiming to achieve optimal combinations of tic effects of Trichoderma when combined with other
microorganisms to benefit the plant [47, 99]. The use of beneficial microorganisms, as this could lead to the
specific Trichoderma species according to particular agri- development of more effective microbial consortia for
cultural needs aims to optimize beneficial interactions agricultural applications. Investigating the interactions
with plants [99] and/or develop more efficient inoculants. of Trichoderma with the pathobiome is also crucial for
The presence of different Trichoderma species in understanding how to manage plant diseases in a holistic
the rhizosphere appears to correlate with changes in manner.
Guzmán-Guzmán et al. BMC Microbiology (2025) 25:434 Page 13 of 17

8. Ali S, Duan J, Charles TC, Glick BR. A bioinformatics approach to the determi-
Furthermore, the integration of Trichoderma into nation of genes involved in endophytic behavior in Burkholderia spp. J Theor
microbiome engineering approaches holds promise in Biol. 2014;343:193–8. ​h​t​t​p​​s​:​/​​/​d​o​i​​.​o​​r​g​/​​1​0​.​​1​0​1​6​​/​j​​.​j​t​b​i​.​2​0​1​3​.​1​0​.​0​0​7.
enhancing plant health and productivity. Future research 9. Andrade-Hoyos P, Silva-Rojas HV, Romero-Arenas O. Endophytic tricho-
derma species isolated from persea americana and cinnamomum verum
should aim to optimize the application methods and roots reduce symptoms caused by phytophthora cinnamomi in avocado.
formulations of Trichoderma inoculants to maximize Plants. 2020;9:1–17. ​h​t​t​p​​s​:​/​​/​d​o​i​​.​o​​r​g​/​​1​0​.​​3​3​9​0​​/​p​​l​a​n​t​s​9​0​9​1​2​2​0.
their benefits under field conditions. By addressing 10. Athinuwat D, Ruangwong OU, Harishchandra DL, Pitija K, Sunpapao A.
Biological Control Activities of Rhizosphere Fungus Trichoderma virens T1-02
these research gaps, we can unlock the full potential of in Suppressing Flower Blight of Flamingo Flower (Anthurium andraeanum
Trichoderma as a cornerstone for sustainable agriculture, Lind.). J Fungi. 2024;10. ​h​t​t​p​​s​:​/​​/​d​o​i​​.​o​​r​g​/​​1​0​.​​3​3​9​0​​/​j​​o​f​1​0​0​1​0​0​6​6.
paving the way for innovative solutions to meet the chal- 11. Awad-Allah EFA, Shams AHM, Helaly AA, Ragheb EIM. Effective applications
of Trichoderma spp. as biofertilizers and biocontrol agents mitigate tomato
lenges of food security and environmental sustainability. fusarium wilt disease. Agriculture. 2022;12:1950. ​h​t​t​p​​s​:​/​​/​d​o​i​​.​o​​r​g​/​​1​0​.​​3​3​9​0​​/​a​​g​r​i​​c​
u​l​​t​u​r​e​​1​2​​1​1​1​9​5​0.
Authors’ contributions 12. Baazeem A, Almanea A, Manikandan P, Alorabi M, Vijayaraghavan P, Abdel-
PG and GS drafted the first version of the work with significant input and Hadi A. In vitro antibacterial, antifungal, nematocidal and growth promoting
revisions from HE. All authors have approved the final version. activities of Trichoderma hamatum FB10 and its secondary metabolites. J
Fungi. 2021;7:331. ​h​t​t​p​​s​:​/​​/​d​o​i​​.​o​​r​g​/​​1​0​.​​3​3​9​0​​/​j​​o​f​7​0​5​0​3​3​1.
Funding 13. Bala IA, Șesan TE, Oancea A, Craciunescu O, Ghiurea M, Răut I, Trică B, Nicolae
G.S. thanks CONAHCYT-Mexico, CIC-UMSNH, and ICTI-Michoacan for their CA, Constantinescu-Aruxandei D, Oancea F. Influence of foliar treatment with
support of his research projects. suspensions rich in Trichoderma chlamydospores on Momordica charantia
physiology, yield, and quality. Horticulturae. 2024;10:371. ​h​t​t​p​​s​:​/​​/​d​o​i​​.​o​​r​g​/​​1​0​.​​3​
Data availability 3​9​0​​/​h​​o​r​t​​i​c​u​​l​t​u​r​​a​e​​1​0​0​4​0​3​7​1.
No datasets were generated or analysed during the current study. 14. Bandara AY, Kang S. Trichoderma application methods differentially affect
the tomato growth, rhizomicrobiome, and rhizosphere soil suppressiveness
Declarations against Fusarium oxysporum. Front Microbiol. 2024;15:1366690. ​h​t​t​p​​s​:​/​​/​d​o​i​​.​o​​r​
g​/​​1​0​.​​3​3​8​9​​/​f​​m​i​c​b​.​2​0​2​4​.​1​3​6​6​6​9​0.
Ethics approval and consent to participate 15. Baron NC, Rigobelo EC. Endophytic fungi: a tool for plant growth promotion
Not applicable. and sustainable agriculture. Mycology. 2022. ​h​t​t​p​​s​:​/​​/​d​o​i​​.​o​​r​g​/​​1​0​.​​1​0​8​0​​/​2​​1​5​0​​1​2​
0​​3​.​2​0​​2​1​​.​1​9​4​5​6​9​9.
Consent for publication 16. Bashir I, War AF, Rafiq I, Reshi ZA, Rashid I, Shouche YS. Phyllosphere microbi-
Not applicable. ome: diversity and functions. Microbiol Res. 2022. ​h​t​t​p​​s​:​/​​/​d​o​i​​.​o​​r​g​/​​1​0​.​​1​0​1​6​​/​j​​.​m​i​​
c​r​e​​s​.​2​0​​2​1​​.​1​2​6​8​8​8.
Competing interests 17. Behiry S, Soliman SA, Massoud MA, Abdelbary M, Kordy AM, Abdelkhalek
The authors declare no competing interests. A, Heflish A. Trichoderma pubescens elicit induced systemic resistance in
tomato challenged by Rhizoctonia solani. J Fungi. 2023;9:167. ​h​t​t​p​​s​:​/​​/​d​o​i​​.​o​​r​g​/​​
1​0​.​​3​3​9​0​​/​j​​o​f​9​0​2​0​1​6​7.
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