Biocontrol Mechanisms of Endophytic Fungi: Review Article Open Access

Adeleke et al.
Egyptian Journal of
Egyptian Journal of Biological Pest Control (2022) 32:46
https://doi.org/10.1186/s41938-022-00547-1 Biological Pest Control
REVIEW ARTICLE Open Access
Biocontrol mechanisms of endophytic fungi

Bartholomew Saanu Adeleke, Modupe Stella Ayilara, Saheed Adekunle Akinola and
Olubukola Oluranti Babalola*
Abstract
Background: Scientific approaches into modern agricultural systems, as opposed to the use of synthetic pesticides
in food production, became important by exploring endophytic fungi capable of protecting plants against pathogens
for maximum crop productivity.
Main body: Diverse endophytic microbes colonizing the internal tissue of plants exhibit beneficial and pathologi-
cal effects on plants. The beneficial endophytic fungi assisted plants in the control of pathogenic endophytic fungi in
plants due to their ability to directly or indirectly promote plant health. Inefficient agricultural practices and envi-
ronmental factors contribute to the disease emergence in plants. Endophytic fungi employed diverse mechanisms
in phytopathogen control by activating and inducing plant resistance through gene expression, synthesis of fungi-
derived metabolite compounds, and hormonal signaling molecules. The mutual coexistence between endophytic
fungi and host plants remains an important mechanism in disease suppression. An in-depth understanding and
selection of beneficial endophytic fungi and interaction between pathogens and host plants are important in manag-
ing challenges associated with the endophyte biocontrol mechanisms.
Conclusion: Research findings on the use of endophytic fungi as bioinoculants are advancing, and understand-
ing endophytic fungi antibiosis action through the synthesis of biocontrol agents (BCAs) can, however, be explored
in integrated plant disease management. Therefore, in this review, the biocontrol mechanism of endophytic fungi
against plant pathogens was highlighted.
Keywords: Biocontrol mechanism, Endophytic microbiome, Fungal diversity, Plant-soil interface, Sustainable
agriculture
Background production using agrochemicals, which are not sustain-

The scientific approaches to the study of plant-fungal able due to negative threats to the ecosystem (Glick et al.
interactions are becoming interesting in modern agri- 2001). To this premise, checkmating these threats to the
culture with prospects to ensure food security and zero ecosystem and exploring potential endophytic microbes
malnutrition among the world populace (Sharma et al. will help achieve a stable ecosystem and grow pathogen-
2021). In recent times, a higher world population index free plants for higher crop productivity (Akanmu et al.
by 2050 has been envisaged with emphasis and recom- 2021).
mendations on the use of biological approaches in tack- Researches focusing on endophytic microbes and
ling food demand pressure, food insecurity, and future exploration as bioinoculants have created many oppor-
food scarcity (Sahu and Mishra 2021). From antiquity, tunities as a substitute for synthetic pesticides usage in
farmers employed diverse approaches to enhance food modern agricultural systems (Orozco-Mosqueda et al.
2021). Nevertheless, information on the endophytic fungi
antibiosis action through the synthesis of biocontrol
*Correspondence: [email protected] agents (BCAs) can, however, be explored in integrated
Food Security and Safety Focus Area, Faculty of Natural and Agricultural
Sciences, North-West University, Private Bag X2046, Mmabatho 2735, plant disease management, which is the focus of this
South Africa review paper.
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Adeleke et al. Egyptian Journal of Biological Pest Control (2022) 32:46 Page 2 of 17
Literarily, the discreet regions in the internal tis- Main body

sue of plants are referred to as the endosphere and Endophytic fungi classification and characteristics
the microorganisms found in these regions are called in the endosphere
endophytic microbes (Dubey et al. 2020). Of most Screening of endophytic fungi against some plant patho-
interesting, microbial endophytes establish mutual- gens has been recently intensified by Abaya et al. (2021),
ism or antagonism association with the host plants, due to their effectiveness as a source biocontrol agent.
depending on their similar or dissimilar genetic make- Endophytic microbes enhance plant growth, diseases
up. The beneficial types that do not express any patho- tolerance and control, and carbon sequestration (Wang
logical effects with unique plant growth-promoting et al. 2022). Endophytic fungi inhabiting various plant
(PGP) attributes, such as phytohormone synthesis, compartments broadly promote plant growth through
nutrient acquisition, secretion of BCAs, and stress different mechanisms classified as direct and indirect
induction mechanism, are referred to as plant growth- mechanisms (Adeleke and Babalola 2022). In the direct
promoting endophytes (Adeleke et al. 2021). mechanism, the endophytes regulate various plant hor-
The endosymbiotic relationship of endophytic fungi mones, such as cytokinin, ethylene, and auxins, enhance
with the host plants aimed to contribute to plant soil nutrient availability, which includes phosphorus
growth and pathogen control depending on their colo- and iron solubilization, siderophore production, and
nization and secretion of biocontrol agents (Reshma nitrogen fixation; whereas in the indirect mechanism,
et al. 2019). Most endophytic microbiomes in the plant the endophytes prevent damage to the plants by releas-
endosphere have been reported to influence plant ing enzymes, antibiotics, hydrogen cyanide, and volatile
phenotypic functions against environmental stresses compounds which inhibit the activities of pathogens,
and control plant pathogens (Yu et al. 2019). Recent and induce systemic resistance (Segaran and Sathiavelu
findings have pointed out the need to elucidate how 2019).
endophytic microbes can be engineered in agricultural Suebrasri et al. (2020) reported the production of plant
biotechnology for plant health sustainability and inte- growth-promoting metabolites, such as the enzymes
gration in crop breeding (Zhang et al. 2020). Depend- (protease, xylanase, amylase, and cellulase), and indole-3-
ing on the plant organ location, some endophytic fungi acetic acid by endophytic fungi, namely; Daldinia
inhabiting below ground level easily change form to eschscholtzii, Diaporthe phaseolorum, Macrophomina
become endophytes due to proximity to the root endo- phaseolina, Trichoderma koningii, and T. erinaceum from
sphere. Interestingly, evidence has shown the dynamic Sunchocke and medicinal plants. It is interesting to note
nature, colonization, and infiltration of endophytic that M. phaseolina, a notorious plant pathogen, can be
fungi from the external root environment (rhizos- beneficial to plants as reported by Suebrasri et al. (2020);
phere) into the internal tissue of plants (endosphere) hence, it could be explored for more plant beneficial
to establish endophytic microbial communities (Yan activities. Also, the strain of T. koningii (ST-KKU1) dis-
et al. 2019). covered by Suebrasri et al. (2020) is also different from
Endophytic microbes directly or indirectly stimu- the strain T. koningii SMF2, which has previously been
late plant growth and sustain plant health based on reported by Xiao-Yan et al. (2006) to be active against
their genes involved in metabolic pathways (Baghel phytopathogens. Sravani et al. (2020) also reported the
et al. 2020). The biocontrol potential of endophytic ability of endophytic fungi from the Hypocreales fam-
microbes can be attributed to their ability to colonize ily to prevent their host plant from insect infection by
plant tissues, produce hydrogen cyanide, and exopol- releasing peramine, which prevented nematodes, insects,
ysaccharide, and stimulate novel genes involved in and other parasites from feeding on them. Also, authors
secretion systems and secondary metabolite secre- reported secretion of indole-like compounds, such as;
tions (Singh et al. 2021). Due to the under-exploration diacetamide and sesquiterpene from endophytic fungi,
of endophytic fungi in plant disease control; research which exerted lethal effects on the other microbes, which
efforts toward harnessing their bioactive secondary could be pathogenic to their host plant. Furthermore,
metabolites as biopesticides and incorporation into endophytic fungi contributed to the enhancement of root
plant disease control remain fundamental and will development patterns in their host plants to increase
help mitigate the effect of synthetic pesticides appli- access to water and other nutrients.
cation on plant growth for improved crop produc- Fungal endophytes are classified according to differ-
tion. Therefore, this review provided an update of the ent criteria. For instance; (i) based on ecology, they are
unique features of endophytic fungi, and the mecha- classified into clavicipitaceous and non-clavicipitaceous
nisms necessitating their roles in plant protection endophytes, (ii) based on the mode of reproduction,
against phytopathogens. they are classified as sexual and asexual endophytes, (iii)
according to transmission, they are classified as vertically their functional profiling (Manzotti et al. 2020). The
and horizontally transmitted endophytes, (iv) according frontline networking of diverse fungal communities
to the source of nutrition, they are classified as biotrophic in the plant-root interface can be influenced by envi-
or necrotrophic endophytes, (v) according to the expres- ronmental factors; biotic, such as pathogens, abiotic,
sion of infection, they are classified as symptomatic and salinity, drought, and high temperature. (Nadeem et al.
asymptomatic endophytes and (vi) based on the body 2014). Root exudate secretion and substrate metabo-
part they attack, they are classified as foliar and root lism; however, serve as key frontline components and
endophytes (Bamisile et al. 2018). The summary of endo- driving factors mediating biodiversity and metabolism
phytic fungi classification was presented in Table 1. of fungal communities’ belowground level (Woźniak
et al. 2019). Based on the nutrient pool in the soil-root
Systemic frontline networking of endophytic fungi interface, this region has been recognized as a ‘hotspot’,
for plant protection which facilitates the establishment of microbial com-
In recent times, research into the plant-microbial interac- munities and colonization of root-associated endo-
tions in the below-and-above plant levels has been con- phytic microbes (Liu et al. 2019).
ceptualized on the endosphere inhabitants. The microbial Critical evaluation of metabolite secretion, which
networking in the root endosphere regions can be influ- facilitates plant–microbe communication, is important
enced by diverse biotic and abiotic factors (Adeleke and to reveal the complex dynamics and type of interac-
Babalola 2021a). The microbial domain tends to show tions that exist between endophytic microbes and the
high biomass below ground level compared to the phyl- host plants (Adeleke and Babalola 2021b). An approach
losphere depending on the prevailing environmental by reductionists stated an impressive production of root
factors (Ananda and Sridhar 2002). Urbina et al. (2018) exudates from plants (Qu et al. 2020). The advancement
reported a higher microbial population in the below plant in endosphere biology through the combined strate-
parts compared to the stem due to the high rhizodepo- gies in understanding plant-fungal interactions can help
sition of organic molecules, which mediated microbial develop a stable approach to fungal biodiversity in plants.
activities below ground. Several beneficial endophytes with bioprospecting in
From the literature, studies on fungal isolation from agriculture have been identified in diverse plant species
the plant environments capable of sustaining plant under different climatic and geographical locations (Jia
growth and health were known with less exploration in et al. 2016). They can be isolated and identified either
phytopathogen control (Bilal et al. 2018). The aforemen- by using direct observation or culture-dependent tech-
tioned might be due to a lack of information on their niques. The direct observation enabled direct visualiza-
transitional networking in the plant-root interface and tion of fungal in plant tissues with the aid of a light and
the type of metabolite produced. Plant roots inhabit- electron microscope, which reflect endophytic fungal
ing fungi have been classified as natural micro-flora in species and those that cannot be cultured on normal
the endo-rhizosphere, whereas those causing diseases in growth media (Nazir and Rahman 2018). However, this
plants were classified to be found dominant in the soil- method can only be used to detect the presence of endo-
root environment (Sylvia and Chellemi 2001). Research phytic fungi by revealing the hyphal structure without the
into the understanding of association that exists among taxonomic grouping, which suggested the need for the
endophyte colonizing plant roots have enabled scien- cultivation-dependent method. In the culture-dependent
tists to deduce their functional traits by in vitro assay technique, endophytic fungi can be isolated from plant
(Vélez et al. 2017). The rhizosphere is regarded as a sub- tissues and subjected to conventional or molecular evalu-
set of root endophytes because they can easily infiltrate ation. The conventional method involves the morpho-
from the external soil environment into the plant roots logical characterization, whereas molecularly, ribosomal
and colonize the region (Ghaffari et al. 2019). The poten- DNA Internal Transcribed Spacer (ITS) sequence analy-
tial of endophytic fungi to induce plant resistance to sis was employed (Nazir and Rahman 2018).
environmental stress adaptors and phytopathogens has Endophytes peculiar to different plants protect them
necessitated more research in their exploration in plant from phytopathogens and promote their growth through
disease management. Additionally, the beneficial plant– different mechanisms. This protection was conferred on
microbe cooperation for increased biomass yield can be the plants to enhance crop productivity and consequently
linked to the diverse functions of these microbes in the food security. The majority of plant endophytic fungi
environment. are active against plant insects; hence, the production of
Insights into the community structure and lifestyle plant insecticides for commercial purposes from these
of the endophytic fungi in some plants by combining endophytes and their metabolites will go a long way in
diverse approaches have been reported to determine improving food security.
Table 1 Classification of fungal endophytes
Mode of classification (MOC) Type-based on MOC Characteristics Plant host Specific endophytic fungi References
Ecology Clavicipitaceous They are associated with sedges Hypocreales family (Grasses and Balansia spp., Acremonium Rodriguez et al. (2009), Khiralla
and grasses; they phylogeneti- sedges) coenophialum, Epichloe spp., et al. (2016), Sravani et al. (2020)
cally belong to the hypocreales Neotyphodium coenophialum,
family. They help to protect the and Epichloe festucae
host plants against insects who
feed on them
Non- Clavicipitaceous They are majorly associated with Non-vascular plants, ferns, and Fusarium culmorum, T. dicco- Llorens et al. (2019)
conifers, ferns, and non-vascular conifers coides, Curvularia protuberate,
plants Colletotrichum spp. and A.
sharonensis,
Mode of reproduction Sexual These fungi reproduce sexually; Angiosperm trees, such as Picea Ph. nodosa, Acer saccharum, Schardl and Craven (2003), Schardl
their stomata which are pro- abies Epichloe spp., Ph. Piceae and Ph. et al. (2004), Tanney et al. (2016), Li
duced during sexual reproduc- Scopiformis, et al. (2017), Bamisile et al. (2018),
Adeleke et al. Egyptian Journal of Biological Pest Control
tion, reduce seed and flower Hume et al. (2020)

production, a condition referred
to as “choke” in grasses
Asexual This group of endophytes shows Ericaceous plants (such as Phialocephala dimorphospora, Ph. Schardl and Craven (2003), Schardl
no symptoms in plants: however, Empetrum nigrum, Calluna scopiformis, Neotyphodium spp., et al. (2004), Tanney et al. (2016), Li
they colonize plant embryos vulgaris, Vacci-nium myrtillus) and Ph. Fortinii, and Ph. sphaeroides et al. (2017), Bamisile et al. (2018),
(2022) 32:46
and thus could be transmitted coniferous trees (such as Picea Hume et al. (2020)
through the plant seeds spp., Pinus spp., and Abies spp.)
Mode of transmission Vertically transmitted It involves the transmission Grasses, e.g., switch grass, Rumex Epichloë spp., A. alternata and C. Hodgson et al. (2014), Gundel et al.
of fungi through the seeds of acetosa, Senecio vulgaris, C. nigra, sphaerospermum (2017)
plants. There are often differ- Plantago lanceolata, Papaver
ences in the proportion of the rhoeas, and Centaurea cyanus
offspring of plants that carry
the seeds, also, there could be
variation in the concentration of
the mycelia and secondary com-
pounds present in the seeds
Horizontally transmitted The majority of the fungi in this Panicum rigidulum Lolium Balansia henningsiana Neotypho- Ren and Clay (2009), Wiewióra
group are natural inhabitants of perenne dium lolii et al. (2015)
the soil from which they get into
the plants. However, they could
be transmitted through the air or
air-borne spores
Sources of nutrition Biotrophs They derive their nutrients from Oryza sativa Aspergillus spp., Magnaporthe Su et al. (2013)
the tissue of living matters oryzae, Penicillium Chrysogenum,
and Hymenochaete sp.
Necrotrophs They kill the host cells and live Syngonium podophyllum Bjerkandera fumosa, Phanero- García-Guzmán et al. (2017)
in them chaetaceae spp., Phlebiopsis
flavidoalba and Hymenochaete
ustulata
Page 4 of 17
Table 1 (continued)
Mode of classification (MOC) Type-based on MOC Characteristics Plant host Specific endophytic fungi References
Expression of infection Symptomatic They cause diseases in plants and Vitis vinifera Acremonium spp., Cadophora Nerva et al. (2019)
show symptoms spp., Coniothyrium spp., Fomitipo-

ria spp., Hypoxylon spp., Paecilo-
myces spp., and Paraphoma spp.
Asymptomatic They cause diseases in plants and Vitis vinifera Leptosphaeriulina spp., Microdip- Nerva et al. (2019)
show no symptoms lodia spp., Scopulariopsis spp.,
Fusarium spp., and Myrothecium
(2022) 32:46
spp.
Body part attacked Foliar They exist on the leaves and Theobroma cacao Colletotrichum tropicale Christian et al. (2017), Christian
stems of plants et al. (2019)
Root They are generally found in the Phoenix dactylifera Aspergillus tubingensis, Corynes- Mahmoud et al. (2017)
root of plants pora cassiicola, Clonostachys
rosea, Penicillium commune, Beau-
veria bassiana, Fusarium solani,
Campanella olivaceonigra, Pho-
mopsis lagerstroemiae, Ilyonectria
radicicola, Aspergillus sclerotiorum
Phomopsis lagerstroemiae, and
Fusarium equiseti
Page 5 of 17
Mechanisms employed by endophytic fungi SA-dependent transcription gene known to induce

in phytopathogen control resistance against rice blast was up-regulated in research
Fungal induced resistance in plant by Shimono et al. (2007) using Harpophora oryzae—an
Endophytic fungi colonize the internal part of both mon- endophytic fungus, to prevent root-necrotization by
ocotyledonous and dicotyledonous plants, help to induce Pyricularia oryzae infection in rice. Efforts to unravel
resistance, and promote plant growth in a diverse num- fungal metabolites associated with plant resistance for
ber of systems (Waqas et al. 2012). Though, most endo- commercial purposes in striving against phytopathogens
phytes solely colonize the root, inducing a system that have been documented (Peng et al. 2021). Many natural
protects other parts of the plant (Adeleke et al. 2021). bioproducts, such as terpenoids, polyketides, steroids,
Induced resistance to plant pathogen is a preventive quinones, flavonoids, alkaloids, and peptides, have been
mechanism that is actively favored by the host plant’s extracted from endophytes, with most reported to have
chemical and physical barriers and induced by both abi- antimicrobial activities against plant pathogens (Latz
otic and biotic factors (Wani et al. 2016). These agents et al. 2018).
(especially fungi) induce exchangeable signals in the host The combined effect of diverse microorganisms in the
plant, so that they activate an acquired response to subse- root endosphere can trigger synergistic effects and the
quent threats from the pathogen(s). An induced response production of BCAs used to control the growth of phy-
is usually triggered by some agents that impel distinctive topathogens (Rojas et al. 2020). Often based on speci-
expression of genes, metabolic changes, and protein syn- ficity, endophyte-induced metabolites can share similar
thesis. The plant’s metabolic swift and change in the eli- pathways to induce metabolism. A familiar instance is
gibility of the plant as a host has led to the reduction in the recent finding that most endophytic fungi produce
disease level (Latz et al. 2018). As mentioned above, both anticancer substances in Taxus brevifolia (El-Bialy and
biotic and abiotic factors can induce host response locally El-Bastawisy 2020). But, many of these endophytes were
or systemically. The activation of defense mechanisms discovered working simultaneously with other organisms
produced by plants for protection against pathogens is as producers (Heinig et al. 2013).
usually referred to as priming (Martinez-Medina et al. To confirm the potency of an antimicrobial substance
2016). Plant-induced resistance is most often linked to produced by an endophyte against pathogens, most
the mobilization potential for cellular defense responses importantly, close contact with the pathogen should be
against nonself. confirmed. Although, it is quite difficult to confirm this
Molitor et al. (2011) reported the mechanisms of S. finding since endophytic fungi are embedded in the plant
indica in inducing plants’ resistance to barley powdery endosphere and the rate of metabolite synthesis may
mildew. Authors inferred that the induced resistance to be hard to quantify. Nonetheless, metabolites induced
powdery mildew by S. indica can be a result of physi- by plant endophyte could be translocated through the
ological responses by reducing pathogen penetration via microorganism to the base of these pathogens within
an increase in local cell death and papillae formation of the plant; whereas, organic compounds secreted can
barley with an up-regulation of HvPR17b (a PR gene) in easily spread to the site of infection (Mejía et al. 2008).
foliage. Other changes in the expression of PR gene in the Meanwhile, it is yet to be confirmed whether the num-
plant root were also noticed. Likewise, genes denoting ber of compounds secreted at the site of infection could
Hsp70, PR1, PR2, and BCI-7 (barley chemically induced be enough to control the invasion of phytopathogens, or
7) are a set of genes instigating protein synthesis, which may be other mechanisms are involved in the plant path-
activate defense reactions as a result of inoculating bar- ogen management.
ley with Blumeria graminis f. sp. hordei (Molitor et al.
2011). These PR-complex are exclusively involved in both Endophytic fungi‑derived compounds activating plant
direct and indirect plant growth promotion and antifun- defenses
gal activities. Just like human responses, nonself/microbial compo-
An investigation conducted on rice roots showed that nents are easily recognized by the plant as specific for-
Harpophora oryzae suppressed the effect of Pyricu- eign substances. Host plants can easily be prepared for
laria oryzae in rice (Su et al. 2013). Also, Polonio et al. potentially harmful microorganisms by inducing defense
(2015) showed the effect of endophyte—Diaporthe citri responses. Although both endophytes and pathogens
on Guaco (Mikania glomerata Spreng.) associated path- are recognized by the host plant, in the same manner,
ogens, such as Fusarium solani and Didymella bryo- the response to both foreign bodies is quite different
niae. Endophytic fungi induced antimicrobial activity (Wani et al. 2016). Invariably, the favorable coexistence
against both pathogens and also increased the growth between endophyte and host plant revealed that fungal-
of the plant (Table 2). Furthermore, OsWRKY4—an induced resistance remains an important mechanism
Table 2 Endophytic fungi and mechanisms of phytopathogen management and disease suppression
Endophyte(s) Host plant Pathogen Mode of action (Biocontrol Inference Reference
process)
Endophytic fungi belonging to Panax notoginseng Mycocentrospora acerina Bioactive compounds produc- Endophytes isolated from P. Zheng et al. (2017)
genera, viz., Aspergillus, Botryo- Fusarium oxysporum, F. solani, tion notoginseng protected plants
tina, Colletrotrichum, Penicillium, Alternaria panax, Phoma against root disease-causing
etc herbarum, organisms
Cladosporium cladosporioides Zygophyllum mandavillei Pseudomonas syringae, Xan- Antimicrobial metabolites, viz., 3-phenylpropionic acid was the Yehia et al. (2020)
thomonas oryzae, Aspergillus 3-phenylpropionic acid, 1-acse- most active compound against
flavus, Fusarium solani tyl-17-methoxy aspidosper- potential fungal and bacterial
midin-20-ol, Isocladosporin, phytopathogens
Cladosporin, etc
Cryptosporiopsis sp., Phialoceph- Picea abies Botrytis cinerea, Phytophthora New antimicrobial metabolites P. sphareoides inhibited all path- Terhonen et al. (2016)
ala sphareoides pini, Heterobasidium parviporum were secreted by both endo- ogens with improved growth of
phytes the plant, while Cryptosporiopsis
sp. gave a stronger inhibitory

effect but retarded the root
growth of Norway spruce
Trichoderma asperellum Lactuca sativa L Curvularia aeria, Corynespora Antagonistic activity with myco- Endophytes inhibited the Baiyee et al. (2019)
cassiicola parasitism growth of highlighted
pathogen(s)
(2022) 32:46
Trichoderma viride Spilanthes pariculata Alternaria solani, Fusarium solani, Antagonistic activity with myco- Endophytes inhibited the Talapatra et al. (2017)
Colletotrichum acutatum parasitism growth of highlighted
pathogen(s)
Penicillium simplicissimum, Gossypium arboretum L Verticillium dahlia Antagonistic activity with myco- Endophytes inhibited the Yuan et al. (2017)
Leptosphaeria sp. parasitism growth of highlighted
pathogen(s)
Diaporthe sp., Leptosphaeria Olea europaea L Colletotrichum acutatum Antagonistic activity with myco- Endophytes inhibited the Landum et al. (2016)
spp., Nigrospora oryzae parasitism growth of highlighted
pathogen(s)
Fomitopsis sp., Fusarium solani, Sophora tonkinensis Gapnep Colletotrichum gloeosporioides Antagonistic activity with myco- Endophytes inhibited the Yao et al. (2017)
Nigrospora sphaerica, Purpureo- parasitism growth of highlighted
cillium lilacinum pathogen(s)
Cornus florida Macrophomina phaseolina, F. Antagonistic activity with myco- Endophytes inhibited the Mmbaga et al. (2018)
solani, F. oxysporum parasitism growth of highlighted
pathogen(s)
Trichoderma citrinoviridae Panax ginseng Botrytis cinerea, Alternaria panax, Antagonistic activity with myco- Endophytes inhibited the Park et al. (2019)
Rhizoctonia solani, Pythium spp. parasitism growth of highlighted
pathogen(s)
Paenibacillus polymyxa Morinda citrifolia L Aspergillus aculeatus Antagonistic activity with myco- Endophytes inhibited the Liu et al. (2018)
parasitism growth of highlighted
pathogen(s)
Page 7 of 17
Table 2 (continued)
process)
Rhizopycnis vagnum Zingiber officinale Rosc Fusarium oxysporum, Sclerotium Antagonistic activity with myco- Endophytes inhibited the Anisha et al. (2018)
rolfsii, Rhizoctonia solani parasitism growth of highlighted
pathogen(s)
Paraconiothyrium variabile Cephalotaxus harringtonia Fusarium oxysporum The Biocontrol process include Metabolites produced from Combès et al. (2012)
the production of metabolites, endophyte showed inhibitory
viz., 13-oxo-9,11-octadecadien- effect against F. oxysporum
oic acid, beauvericin
Induratia coffeana, I. yucatan- Phaseolus vulgaris L Peudocercospora griseola, Sclero- Antagonistic activity with myco- Both species of Induratia control Mota et al. (2021)
ensis tinia sclerotiorum, Colletotrichum parasitism diseases caused by the three
lindermuthianum pathogens on common beans
Hypoxylon anthochroum, Nod- Solanum lycopersicum var. Fusarium oxysporum Volatile organic compounds VOC showed antifungal activity Medina-Romero et al. (2017)
ulisporium spp. cerasiforme synthesized include; Phenylethyl both in planta and in vitro
alcohol, 2-methyl-1-butanol, assessment

ocimene, terpinolene, etc
Fungal genus from Aspergillus, Cannabis sativa L Botrytis cinerea, Trichothecium Antagonistic activity with myco- Endophytes inhibited the Kusari et al. (2013)
Chaetomium, Paecilomyces, roseum parasitism growth of phytopathogens
Penicillium
(2022) 32:46
Diaporthe citri Mikania glomerata Spreng Fusarium solani, Didymella Antagonistic activity with myco- Endophytes induced antimi- Polonio et al. (2015)
bryoniae parasitism crobial activity against both
pathogens
Bipolaris sp., Fusarium sp., Vitis labrusca L Alternaria spp., Glomerella spp., Antibiosis, parasitism, and pro- Biological control agents help in Felber et al. (2016)
Phoma sp., etc Sphaceloma spp. duction of lytic enzymes the control of phytopathogens
Muscodor yucatonensis, Penicil- Monarda citriodora Sclerotina spp., Colletotrichum Antagonistic activity by direct Endophytes showed biocontrol Katoch and Pull (2017)
lium commune, A. oryzae capsica, A. flavus, A. fumigatus contact suspected to have activity against highlighted
bioactive compounds pathogens
Lasiodiplodia theobromae, Piper hispidum Sw Alternaria alternata,
Phoma herbarum, Schizophyllum
commune
Colletotrichum spp., Phyllos- P. herbarum and S. commune Both endophytes had a high Orlandelli et al. (2015)
ticta citricorpa, Moniliophthora produce proteolytic enzymes enzymatic halo and were able
perniciosa to protect the plant against
pathogens
Fusarium sp., Penicillium sp., Camellia oleifera Anthracnose phytopathogenic Mycoparasitism Endophytes inhibited the Yu et al. (2018)
Pichia spp., Postalotiopsis sp., fungus growth of pathogens (especially
Xylaria sp. Oidium sp.)
Aporospora terricola, Aureoba- Vitis labrusca L Fusarium oxysporum f. spp. Antibiosis against plant patho- Both F. flavus and C. gloeospori- Brum et al. (2012)
sidium pullulans, Bjerkandera herbemontis gens oides showed antagonistic
adusta, Colletotrichum bonin- activity against F. oxysporum
ense, C. gloeosporioides, Flavodon
flavus,
Page 8 of 17
Table 2 (continued)

process)
Cladosporium sp., Ophiognomo- Coffea arabica L. cultivar IAPAR- Glomerella spp. (CNPUV 378), Antagonistic effect against Endophytes inhibit the growth Bongiorno et al. (2016)
nia sp., Trichoderma sp., etc 59 Colletotrichum spp., Sclerotinia phytopathogens of pathogens
sclerotiorum
(2022) 32:46
Muscodor coffeanum, M. vitige- Coffea arabica Rhizoctonia solani (LAPS 369), Volatile organic compounds VOC produced by endophytes, Monteiro et al. (2017)
nus, M. yucatanensis, Simplicil- Fusarium oxysporum (LAPS 152), produced which helps to inhibit the
lium sp. Phoma spp. (DFP 01), F. solani growth of pathogens
(LAPS 298), F. verticillioides (CML
1896), Cercospora coffeicola (CML
2984), Pestalotia longisetula (DFP
02), A. ochraceus (SCM 1.15)
Diaporthe citri, Phomopsii spp. Sapindus Saponaria L Fusarium solani, Glomerella spp., Amylase, pectinase, and cel- Phyto-protective properties Santos et al. (2019)
Moniliophthora perniciosa lulase produced of these compounds were
observed against pathogens
Page 9 of 17
used by endophytic microorganisms in disease suppres- of host pathogens (Hossain et al. 2008). Meanwhile, S.
sion (Fig. 1). Normally, specific endophytic fungal com- indica instigated other resistance pathways different
ponents, viz., cell wall, lipids, protein substances, volatile from ethylene/JA in other pathophysiological responses.
compounds, BCA, and some molecules with hormonal In another scenario, SA-dependent pathway is induced
responses, are usually recognized by the host plant. These in a T. asperellum inhabited plant (Yoshioka et al. 2012).
compounds/components are selective endophyte-derived This showed that hormonal responses and interaction
compounds that induce plant defense mechanisms (Latz with the host plant were very complex, withal, numerous
et al. 2018). events; and cross-communication is normally involved in
Components of microbial origin are oftentimes plant-induced responses. Microbial inhabitation changes
referred to as MAMPS–microbe-associated molecular a plant’s normal reflexes and profile, at times, rather
patterns or PAMPS pathogen-associated molecular pat- than affecting only a single hormonal response, it brings
terns, which induce MAMP/PAMP-triggered immuno- together a couple of them. Nevertheless, the coexistence
logical response (Nürnberger and Kemmerling 2009). between microbial strain and the host of the inducing
Components of the fungal cell wall, such as β-glucans agent also determines variations in hormonal responses.
and chitin, are referred to as MAMPs, they are usually Despite the tremendous progress in the studies on signal
recognized by the receptors of plants to trigger immune transfer in induced resistance, there is still a lacuna in
responses. Endophytic secretions, such as peptides and attributing functions to each hormone in signal transduc-
proteins, have been described by most researchers as tion, especially in complex systems. Therefore, there is a
agents that trigger host plant responses (Rojas et al. need to intensify the defense mechanisms in a plant and
2020). Other secreted enzymes, viz., cellulase, xylanases, adopt it as a biomarker to detect an induced resistance.
and chitinases, were produced as a result of infection also
induce plant defense and are easily identified by hosts via Use of plant defense mechanisms to detect induced
their decomposed products (Druzhinina et al. 2011). resistance
Proteins rich in cysteine and fungal effectors are An induced resistance occurs as a result of activated
secreted as a result of endophytic and pathogenic inhabi- plant-defense mechanisms to make the plant less suscep-
tation processes to increase host plant compatibility, by tible to a variety of pathogens. Most of these mechanisms
inducing physiological and defense responses (Ku et al. are activated simultaneously to help strengthen plant
2020). Nonetheless, different studies have shown that physical barriers, and secretion of pathogen-repellants
compounds produced to inhibit the growth of compet- in the form of proteins and enzymes with antimicrobial
ing microorganisms can also instigate resistance (Akinola properties to prevent phytopathogens (Farhangi-Abriz
and Babalola 2021). The above-mentioned products and Ghassemi-Golezani 2019).
obtained from endophytic fungi have the complexity and To study if an endophytic fungus induces a resistance
potential to induce plant defense responses necessary to mechanism against a specific pathogen, two (2) crite-
reduce the menace posed by plant pathogens and other ria are used to test and classify plant responses. Firstly,
soil-related anomalies. the induced responses should control the targeted plant
pathogen(s). There should be proven that fungal-induced
Hormonal signaling and its ability to induce resistance response could effectively eradicate the pathogens, and
Plant hormones are crucial in the transmission of struc- expressions observed therewith should be related to hin-
tural and comprehensively inherent resistance. The sys- dering pathogen infection. Secondly, the elimination pro-
temically induced plant resistance can be classified into cess of phytopathogen should be correlated with Koch’s
two; namely; ISR-induced systemic resistance and SAR– postulate. This can be verified by noticing the defense
systemically acquired resistance. Ethylene and hormonal response expression after introducing the pathogen to
jasmonic acid (JA) are the most important secretion pro- the plant. In essence, adopting the principle of exclu-
duced in ISR, whereas SA–salicylic acid plays a pivotal sion is an acceptable condition to evaluate the effective-
role in systemically acquired resistance (Latz et al. 2018). ness of induced resistance in plant protection. With that
The coherent relationship among hormonal responses is said, excluding a direct in vitro assessment of an induced
induced by the coexistence of ethylene, JA, and SA main- resistance assumes the effect of the induced response to
tains defensive responses in the plant (Latz et al. 2018). the pathogen is unacceptable.
For instance, the defensive response induced by the bio- Another conserved process is the ability to strengthen
logical agents’, viz., Serendipita indica, Penicillium sp., a plant’s structural barriers to resist the easy invasion of
and Trichoderma asperellum, has induced the produc- pathogens and reinforcement of cell wall appositions,
tion of ethylene and JA-dependent systemic resistance which might have been involved (Waller et al. 2005).
that plays important role in preventing the inhabitation According to different studies, this effect occurred as a
Fig. 1 Mechanism of endophytic fungi in plant growth promotion and disease suppression
result of inducing agents. For instance, a study on Tricho- thionins (Sels et al. 2008). In research by Lahlali et al.
derma harzianum (T-22) showed the expression of an (2014), a plant-related protein (PR2) —β-1, 3-glucanase
enzyme (phenylalanine ammonia-lyase—PAL) involved was enhanced in oilseed rape plants infected by Plasmo-
in lignin formation was well enhanced in maize (Shoresh diophora brassicae when an endophytic fungus—Hetero-
et al. 2010), whereas in the case of another strain T. har- conium chaetospira colonized the plant. Also, Combès
zianum (T-203), the cortical and epidermal cell walls of et al. (2012) detected a systemic resistance induced by
cucumber fruit were strengthened and the process was an endophytic fungus—Paraconiothyrium variable on
confirmed to have been induced by intercellular inhabi- Cephalotaxus harringtonia infected by Fusarium oxyspo-
tation of endophytic fungi (Yedidia et al. 1999). Systemic rum, which led to the production of important metabo-
introduction of inducing agents has been very useful in lites such as 13-oxo-9,11-octadecadienoic acid and
the promotion/activation of important proteins and beauvericin that are capably inhibiting the growth and
metabolites in plants, with antimicrobial properties pathogenic effect of F. oxysporum as shown in Table 2.
that are very effective against plant pathogens. Another An induced resistance activated by F. solani (strain
example of a useful metabolite was phytoalexin-type Fs-K)—infested tomato enhanced the expression of thau-
compounds reported by Oliveira et al. (2016). The path- matin-like (PR5) and endo-proteinase (PR7) enzymes
ogen-related (PR) proteins produced as a result of the in the plant (Kavroulakis et al. 2007). More so, Waller
colonization of endophytic fungus also perform other et al. (2008) also hypothesized the up-regulation of pro-
roles, such as stress response and antimicrobial proper- tein HvPR17b was suspected to have antifungal activity
ties. These metabolites include peptides and enzymes, in a barley-infested with endophytic–Serendipita indica
viz., thaumatin-like proteins, lipid transfer proteins, and against Blumeria graminis f. sp. hordei. The synergistic
effect of Fusarium graminearum and S. indica on barley the tree eliminated specific endophytes, creating space
was also reported by Deshmukh and Kogel (2007) with for pathogens to grow.
the reduced expression of pathogen-related genes (PR1b Phyto-pathological control mechanisms involving com-
and PR5), which means PR genes are not always involved/ petitive exclusion incorporate the co-occurrence of other
pronounced in all systems. mechanisms and also require endophytic colonization of
intracellular plant parts where the pathogen might have
attacked. For instance, the treatment of a sterile seed with
Antibiosis activities of endophytic fungi against plant endophytic fungus isolated from a cacao tree reduced the
pathogens effect of Phytophthora spp. on the plant leaves (Arnold
Antibiosis—an antagonistic relationship involving endo- et al. 2003). The colonization of the oilseed roots by an
phytic fungal control of potential plant pathogens using endophytic fungus—Heteroconium chaetospira, nega-
metabolic substances was produced by endophytes. A tively correlated with the symptoms of clubroot disease
purified form of Efe-AfpA mined from an apoplastic fluid (Lahlali et al. 2014). Withal, an increase in the inoculum
of endophyte-inoculated red fescue showed anti-parasitic size of the pathogen reduced the control effect, show-
activity against Sclerotinia homoeocarpa (Ambrose and ing the restraints of competition. Competitive exclusion
Belanger 2012). The same result was also observed in the could be well studied using in planta microscopic assess-
recombinant product of Efe-AfpA expressed gene found ment and quantification of endophyte biomass related to
in Pichia pastoris. In a transcriptome study to detect the phytopathogen management. To evaluate endophyte—
percentage protein (Efe-AfpA) produced from the endo- pathogen in planta interaction, visualization using
phytic relationship between Epichloë festucae and Fes- microscopy is advisable, when investigating pathogen
tuca rubra sp., a 6%—Efe-AfpA was produced from the strains and fungal BCA (Latz et al. 2018). In situ detec-
fungal transcriptome. The product mined from the study tion of metabolite distribution, microorganism involved
was observed to have the same property as the product and genomic evaluation of the role of mined metabolites
secreted in a relationship between Aspergillus spp. and could be determined using molecular 3D cartography-
Penicillium sp. as reported by Tian et al. (2017). mass spectrometry as described by Floros et al. (2017).
The synergy between endophytic funguses—Paraco-
niothyrium strain SSM001 and a yew tree producing
Taxol against wood-decaying fungus was investigated Mycoparasitism
by Rafiqi et al. (2013). Although, the yew tree usually Fungal parasitism involves the direct reliance of a fungus
forms bark cracks that allow easy penetration of patho- on another fungus for nutrients. The process of myco-
gen. Meanwhile, the endophytic fungus was observed parasitism occurs either through necrotrophic or bio-
growing toward these cracks in a way to prevent Taxol trophic relationships. In necro-trophism, parasites live
accumulation and also down-regulated the transcription on the dead cells of the host, while bio-trophism is a situ-
of Taxus genes, viz., DXP reductoisomerase and taxadi- ation, whereby the parasite takes nutrients from a living
ene synthase, that is very crucial for Taxol secretion. An host (Kim and Vujanovic 2016).
in vitro assessment of strain SSM001 endophytic fungus Normally, in planta verification of mycoparasitism is
and Taxol treatment prevented the growth of impor- very hard, since the transfer of nutrients among micro-
tant wood-decaying fungal species, such as: Perennipo- organisms is very tedious to detect. In essence, most
ria subacida, Phaeolus schweinitzii, and Heterobasidion studies claiming mycoparasitism only based their verifi-
annosum, meanwhile, the growth of the endophyte strain cations on circumstantial shreds of evidence. The close
SSM001 was not hindered by Taxol (Soliman et al. 2015). relationship between two fungi is not enough to claim
a mycoparasitism, rather they are referred to as a fungi-
colous relationship. Mycoparasitism may occur directly
Competition or indirectly. In indirect fungal-parasitism, a metabo-
Microbial competition remains an important factor lite produced by the parasite releases nutrients from the
determining plant tissue inhabitation and a probable way host at a distance, while direct contact with the prey is
endophytes inhibit pathogens from colonization (Mar- referred to as direct mycoparasitism (Latz et al. 2018).
tinuz et al. 2012). The endophytic fungus colonizes plant In either case, the parasite secrets some metabolites to
tissues systemically and locally, within or outside the tis- release host nutrients such as toxins, antibiotics, and
sues. Through this method, rapid inhabitation and feed- cell wall degrading enzymes (Kim and Vujanovic 2016).
ing on available nutrients are easily explored, and also For instance, a study by Chamoun et al. (2015) showed
occupy the space that could have been filled by potential the production of specialized compounds in a relation-
pathogens. A study by Mohandoss and Suryanarayanan ship between Manatephorus cucumeris and Stachybotris
(2009) on mango leaves showed that the fumigation of elegans, where S. elegans was preying on T. cucumeris.
A lot of researchers misplace mycoparasitism and anti- interaction put a lacuna on the importance of endophytes
biosis with potential intermingling relatedness making it on disease suppression in the plant. Therefore, evaluating
very hard to differentiate interactions, but this has shown endophyte-pathogen interaction using in vitro assess-
that parasitic relationships among microorganisms could ment will only result in false conclusions. For instance, an
employ several mechanisms to prey on each other. in vitro experiment on Pseudozyma flocculosa- an endo-
Since microbial interactions are easily studied using phytic yeast suggested to inhibit the growth of Blumeria
conventional methods, the inhibitory relationship among graminis in barley via antibiosis, but after adopting cellu-
microorganisms becomes very easy to study using tra- lar microscopy and transcriptomics to study the control
ditional methods, such as microscopy techniques and mechanism, it was concluded that the parasitic relation-
culturing in Petri dishes than in planta screening. Using ship was mycoparasitism not antibiosis (Laur et al. 2018).
simple microscopic methods, mycoparasite is observed (ii) what are the pattern and colonization methods of a
having direct contact with the host either by coiling biological control agent? Understanding the mechanism
around the hyphae of the prey for easy acquisition of involved in BCA infiltration and pattern will help link
nutrients. This relationship was demonstrated in a study intra-and-intercellular endophytic structure to a disease
by Donayre and Dalisay (2016). An endophytic fungus, suppression mechanism (Compant et al. 2005), (iii) what
Geotrichum sp. isolated from Echinochloa glabrescens is the pathosystem involved in the BCA mechanism of
was observed having a direct mycoparasitic relationship action? Understanding the mechanisms of biological con-
with a soil-borne pathogen, Thanatephorus cucumeris. trol agents in phytopathogen management is also very
Likewise, three (3) endophytic fungi isolated from Phrag- important. Conclusively, if a BCA was isolated from an
mites australis were observed penetrating and coiling external surface and identified to be of endophytic origin
around the hyphae of soil-borne pathogens to degrade when applied in planta, does the BCA grow intracellu-
their cytoplasm, meanwhile, other degrading enzymes, larly? It is necessary to confirm the function of the bio-
viz., β-1, 3-glucanase, and extracellular cell wall degrad- logical control agent within the plant tissue, to confirm
ing enzymes, were involved in the process (Cao et al. its potency in phytopathogen control (Busby et al. 2016).
2009).
Conclusions
Challenges associated with endophyte biocontrol Endophytic microbes employ direct and indirect mecha-
mechanisms nism options in plant growth promotion and protection
For an in-depth understanding, utilization, and selec- against pathogens. Exploring endophytic microbes as
tion of endophytic fungi, an assessment of the biology bioinoculants, upon inoculation can cause changes in the
behind the interaction between pathogen, host plant, and plant’s physiological and phenotypic modifications, thus
endophytic fungus is required in addition to the physi- boosting plant tolerance to biotic and abiotic stressors.
ological activities involved in the tie-in. Some impor- The biotechnological importance of valuable metabolites
tant principles are generally acceptable for the study of produced by endophytic fungi, which stimulate antibiosis
endophytic fungus and biological control agents (BCAs). against phytopathogens for plant protection is less to be
These include; (i) activation of plant defense mechanism fully explored in plant disease management.
induced by endophyte, (ii) inhibition via mycoparasitism, The combined application of culture-dependent and
(iii) inhibition through antibiosis, and (iv) competition culture-independent techniques helps in the predictive
for nutrient and space (Latz et al. 2018). Also, the most functional analysis of notable genes involved in phyto-
times and several mechanisms may be activated at the hormone synthesis, secretion systems, biocontrol, and
same time. Nutrient acquisition for plant growth promo- synthesis of cellular components, metabolic pathway, and
tion or modification of the level of plant growth hormone secondary metabolites (SM) from endophytic microbes.
can generally improve plant health and disease suppres- The presence of biocontrol genes in some endophytic
sion (Berthelot et al. 2016). Studying the complex interac- fungi was suggested their ability to control plant diseases.
tion between pathogen, host plant, BCAs and the process Studies have successfully shown the biocontrol activity of
of pathogen inhibition are complicated to study. To bet- endophytic fungi, which promise to be used in the syn-
ter explore the relationship between highlighted factors, thesis of certain novel BCA to confront the challenges
several questions are raised and these include; (i) are the associated with phytopathogens control in plants. Nev-
mechanisms involved in BCAs really assessable within ertheless, how endophyte infiltrates plant endosphere is
the tissue of plant (in planta)? Because most mecha- still a question that demands clarification by researchers.
nisms associated with endophytic metabolite production The SM biosynthesis potential of endophytic fungi is
in plants are usually performed under in vitro condi- characterized by complex biocontrol activity, which can
tions. Putting out one of the factors from the tripartite be explored as valuable bioproducts. Hence, providing
updated information on the plant growth-promoting Adeleke BS, Babalola OO (2021a) The endosphere microbial communities, a
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Abbreviations cue reveals numerous effects on host transcriptome and an abundance
BCA: Biological control agent; MAMPS: Microbe-associated molecular patterns; of highly expressed fungal secreted proteins. PLoS ONE 7:e53214.
PAMP: Pathogen-associated molecular patterns; ISR: Induced systemic resist- https://doi.org/10.1371/journal.pone.0053214
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acid; PAL: Phenylalanine ammonia-lyase. grove species on the west coast of India. Canad J Microbiol 48:871–878.
https://doi.org/10.1139/w02-080
Acknowledgements Anisha C, Jishma P, Bilzamol VS, Radhakrishnan E (2018) Effect of ginger endo-
National Research Foundation of South Africa and The World Academy of phyte Rhizopycnis vagum on rhizome bud formation and protection
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stipend to BSA (UID: 116100). MSA and SAA were grateful to North-West doi.org/10.1016/j.bcab.2018.02.015
University, South Africa, for the postgraduate bursary. OOB recognized NRF for Arnold AE, Mejía LC, Kyllo D, Rojas EI, Maynard Z, Robbins N, Herre EA (2003)
grants (UID: 123634; 132595) that support work in her research group. Fungal endophytes limit pathogen damage in a tropical tree. Proc Natl
Acad Sci 100:15649–15654. https://doi.org/10.1073/pnas.2533483100
Author contributions Baghel V, Thakur JK, Yadav SS, Manna MC, Mandal A, Shirale AO, Sharma P,
BSA and OOB had the idea for the review article and suggested the review Sinha NK, Mohanty M, Singh AB, Patra AK (2020) Phosphorus and potas-
topic. BSA, MSA, and SAA performed the literature search and wrote the first sium solubilization from rock minerals by endophytic Burkholderia sp.
draft. BSA revised and formatted the manuscript. OOB made substantial and strain FDN2-1 in soil and shift in diversity of bacterial endophytes of
technical contributions to the structure of the various manuscript drafts. All corn root tissue with crop growth stage. Geomicrobiol J 37:550–563.
authors read and approved the final revised manuscript. https://doi.org/10.1080/01490451.2020.1734691
Baiyee B, Ito S-i, Sunpapao AJP, Pathology MP (2019) Trichoderma asperellum
Funding T1 mediated antifungal activity and induced defense response against
This research was funded by the National Research Foundation, South Africa leaf spot fungi in lettuce (Lactuca sativa L.). Physiol Mol Plant Pathol
(UID: 123634; 132595). 106:96–101. https://doi.org/10.1016/j.pmpp.2018.12.009
Bamisile BS, Dash CK, Akutse KS, Keppanan R, Wang L (2018) Fungal endo-
Availability of data and materials phytes: beyond herbivore management. Front Microbiol 9:544
Not applicable. Berthelot C, Leyval C, Foulon J, Chalot M, Blaudez D (2016) Plant growth
promotion, metabolite production and metal tolerance of dark septate
endophytes isolated from metal-polluted poplar phytomanagement
Declarations sites. FEMS Microbiol Ecol. https://doi.org/10.1093/femsec/fiw144
Bilal L, Asaf S, Hamayun M, Gul H, Iqbal A, Ullah I, Lee IJ, Hussain A (2018) Plant
Ethics approval and consent to participate
growth promoting endophytic fungi Asprgillus fumigatus TS1 and
Not applicable.
Fusarium proliferatum BRL1 produce gibberellins and regulates plant
endogenous hormones. Symbiosis 76:117–127. https://doi.org/10.
Consent for publication
1007/s13199-018-0545-4
Not applicable.
Bongiorno VA, Rhoden SA, Garcia A, Polonio JC, Azevedo JL, Pereira JO,
Pamphile JA (2016) Genetic diversity of endophytic fungi from Coffea
Competing interests
arabica cv. IAPAR-59 in organic crops. Ann Microbiol 66:855–865.
The authors declare that they have no competing interests.
https://doi.org/10.1007/s13213-015-1168-0
Brum M, Araújo W, Maki C, Azevedo J (2012) Endophytic fungi from Vitis
Received: 4 February 2022 Accepted: 17 April 2022
labrusca L. (‘Niagara Rosada’) and its potential for the biological control
of Fusarium oxysporum. Genet Mol Res 11:4187–4197. https://doi.org/
10.4238/2012.December.6.2
Busby PE, Ridout M, Newcombe G (2016) Fungal endophytes: modifiers of
plant disease. Plant Mol Biol 90:645–655. https://doi.org/10.1007/
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Biocontrol mechanisms of endophytic fungi

Background production using agrochemicals, which are not sustain-

Literarily, the discreet regions in the internal tis- Main body

tion, reduce seed and flower Hume et al. (2020)

show symptoms spp., Coniothyrium spp., Fomitipo-

Mechanisms employed by endophytic fungi SA-dependent transcription gene known to induce

sp. gave a stronger inhibitory

alcohol, 2-methyl-1-butanol, assessment

Endophyte(s) Host plant Pathogen Mode of action (Biocontrol Inference Reference

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