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Exploring the efficacy of antagonistic rhizobacteria as native biocontrol

agents against tomato plant diseases

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DOI: 10.1007/s13205-020-02306-1

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3 Biotech (2020) 10:320
https://doi.org/10.1007/s13205-020-02306-1

REVIEW ARTICLE

Exploring the efficacy of antagonistic rhizobacteria as native

biocontrol agents against tomato plant diseases
S. Karthika1 · Sherin Varghese1 · M. S. Jisha1

Received: 7 February 2020 / Accepted: 15 June 2020

© King Abdulaziz City for Science and Technology 2020

Abstract
As the environmental and health concerns alert the necessity to move towards a sustainable agriculture system, biologi-
cal approach using indigenous plant growth-promoting rhizobacteria (PGPR) gains a strong impetus in the field of plant
disease control. In this context, the present review article addresses the usage of rhizospheric antagonistic bacteria as a
suitable alternative to control tomato fungal diseases namely Fusarium wilt and early blight disease. Biological control has
been considered to be an eco-friendly, safe and effective method for disease management. The inherent traits of PGPR to
antagonize a pathogen through various mechanisms has been investigated extensively to utilize them as potent biocontrol
agents (BCA). Hence, the article provides a detailed account on different biocontrol mechanisms displayed by BCA. It is
also suggested that the use of bacterial consortium ensures consistent performance by BCA in field conditions. Likewise,
this review also deals with the opportunities and obstacles faced during commercialization of these antagonistic bacteria as
biocontrol agents in the market.

Keywords Tomato diseases · Plant growth-promoting rhizobacteria · Biocontrol · Bioformulation

Introduction and scientists tend to emphasize sustainable and biological

Tomato (Solanum lycopersicum), a dicotyledonous plant
belonging to Solanaceae family, is considered as the ances-
tor of cultivated tomatoes. It is a short duration crop and
re-enumerative vegetable, appreciated with high nutritive
esteem and antioxidant curative and therapeutic properties
(Nour et al. 2018; Azeez et al. 2019). Hence, it is a versatile
vegetable in the Indian culinary tradition. It exerts several
beneficial effects on health as it is rich in vitamin A, B and
C, minerals, organic acids and a considerable amount of total
sugar (Salim et al. 2017). Moreover, it is also used commer-
cially to produce various food products, such as ketchup,
soup, paste and powder (Paul and Kehinde 2012; Manivan-
nan and Tholkappian 2013). Therefore, it leads to increased
demand in improved quality, high yield, better storage dura-
bility and promising disease and pest management system
for tomato plants. In order to keep a balance between crop
production and demographic food demand, the growers
* M. S. Jisha
jishams@mgu.ac.in
1
School of Biosciences, Mahatma Gandhi University,
Kottayam, Kerala 686560, India
agricultural production (Syed et al. 2018). One of the major
constraints faced during their cultivation is the assault of
pathogens both in fields as well as greenhouse condition
(Sanoubar and Lorenzo 2017; He et al. 2020).
The crop is affected by various pathogenic diseases lead-
ing to a decrease in its nutritional contents and the over-
all economy (Danish et al. 2014). Late blight induced by
Phytophthora infestans is one of the damaging ailments
of tomato bringing about huge financial loss (Singh et al.
2017). Sclerotinia rot brought about by Sclerotinia sclerotio-
rum, is another significant infection influencing the tomato
crop profitability. Several studies on wilt, crown and root rot
diseases in tomato caused by Fusarium species have been
reported (Laurence et al. 2014; McGovern 2015). Root knot
caused by the nematode Meloidogyne sp. is another destruc-
tive and widespread disease in tomato (Zhou et al. 2016). It
not only influences the crop yield directly but also makes
the plants increasingly prone to fungal and bacterial infec-
tions (Ashraf and Khan 2010). Bacterial leaf spot is a com-
mon bacterial ailment of tomato caused by Xanthomonas
campestris. It is exceptionally ruinous in both greenhouses
as well as in field conditions, causing 10–50% yield loss
(Singh et al. 2017). Ralstonia solanacearum is a soil-borne

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320 Page 2 of 17
plant pathogen that causes bacterial wilt in tomatoes and
hampers their yield (Huang et al. 2013). Clavibacter michi-
ganensis infection systemically causes wilting and canker
on the stem, while blister-like spots are developed in locally
infected leaves causing a substantial economic loss (Agrawal
et al. 2012). Among the fungal diseases encountered by
tomato plants, wilt disease and early blight disease caused
by Fusarium oxysporum. f. lycopersici (FOL) and Alternaria
solani (AS), respectively, are known to be the most deleteri-
ous ones (Pawar et al. 2016).
Mere cultural sanitation is not much effective since the
pathogens are pandemic. Although current agricultural prac-
tices have been subject to the utilization of agrochemicals as
a solid technique for harvest insurance, expanded chemical
utility has caused a few negative impacts including resist-
ance development among pathogens to the applied agents
and their non-target natural effects (Ghazanfar et al. 2016).
These reasons have prompted a quest for substitutes for
chemical inputs. Biological control is considered as a suc-
cessful option for lessening the seriousness of the diseases
(Linu and Jisha 2017).
The utilization of rhizobacteria that colonize the under-
lying foundations of harvest plants and smother soilborne
ailments is turning into an elective choice as compared to the
utilization of chemical fungicides as referenced before. The
utilization of plant growth-promoting rhizobacteria (PGPR)
as soil inoculants for control of soilborne infections, can
be a suitable organic option. Rhizobacteria with biocontrol
viability frequently give long haul security from soilborne
pathogens at the root surface since they have the ability to
quickly colonize the rhizosphere and spread down the root
from a solitary seed treatment or soak application into the
soil (Haldar and Sanghamitra 2015).
Currently, a large number of bacterial strains have been
isolated and identified for their advancement as biocontrol
agents against tomato diseases. Punja et al. (2016) utilized
Bacillus subtilis strain under greenhouse conditions to con-
trol the postharvest fruit infection. B. subtilis strains were
likewise used by Kilani-Feki et al. (2016) for the suppression
of Botrytis cinerea, the causative agent of tomato fruit rot.
Gowtham et al. (2016) used ten rhizobacterial strains to deal
with the Fusarium wilt in tomato and observed that two dis-
tinct strains Bacillus amyloliquefaciens and Ochrobacttrum
intermedium significantly repressed the incidence of wilt and
furthermore enhanced the vigour index of seedlings. Singh
et al. (2017) have depicted the efficacy of Pseudomonas fluo-
rescens strains against different diseases, such as damping-
off, root rot, stem canker and leaf blight of tomato. Higher
accumulation of phenolic compounds in leaf tissues induced
by P. aeruginosa against A. solani makes it an effective
BCA against this pathogen (Hariprasad et al. 2013). Abo-
Elyousr et al. (2019) reported that the PGPR (B. subtilis, B.
amyloliquefaciens, P. fluorescens and P. aeruginosa) and
13
3 Biotech (2020) 10:320
their formulations could successfully control the wilt and
canker disease of tomato caused by Clavibacter michigan-
ensis subsp. michiganensis, in vitro and under greenhouse
conditions. Attia et al. (2020) investigated a new strategy by
using PGPR in the induction of systemic resistance of plants
diseases. They were confirmed to be efficient in the reduc-
tion of early blight disease-causing A. solani which infected
tomato plants following the assessment of invitro antago-
nistic activity. The isolated bacterial strains namely Bacillus
subtilis SBMP4, Lysinibacillus fusiformis NBRC15717 and
Achromobacter xylosoxidans NBRC15126 could increase
the induction of systemic resistance in the tomato plant and
thereby reduce the percent disease severity by 13.0%. They
also recorded a protection percentage of 84.3% when com-
pared to non-treated plants.
Incorporation of microorganisms for disease control/plant
growth promotion in cropping systems and the elimination
of chemical utility is reliant on effective selection, screen-
ing and safety analysis of potential PGPR strains. Moreover,
information on targeting diseases, cost of mass-scale produc-
tion and registration procedures also need to be amended
likewise to raise the market status of these biocontrol enti-
ties. This review is an attempt to explore the role of antago-
nistic rhizobacteria in biological control of tomato plant dis-
eases namely Fusarium wilt and early blight. Moreover, the
hurdles and benefits of commercializing these microbes in
the agriculture sector are also included in this review article.
Fusarium wilt and early blight diseases
of tomato
Fusarium wilt is considered as the deadliest disease con-
fronted by tomato plants throughout the world, particu-
larly in the uplands (Fig. 1a). The causal agent is Fusarium
oxysporum f. lycopersici (FOL) (Sacc.). They are well-
established soil-borne pathogens ubiquitous in all soil types.
They are saprophytic and are able to survive in the soil for
a prolonged period (Mj et al. 2017). At least 32 countries
have reported this disease, among which warm climate
countries have been severely affected (Bawa 2016). Srinivas
et al. (2019) reviewed that Fusarium wilt is characterized by
60–70% of fruit yield loss with wilted plants containing yel-
lowed leaves. FOL invades plant’s vascular system and binds
inside xylem vessel, which results in impaired water trans-
portation, finally leading to wilting of the foliage (Boukerma
et al. 2017) (Fig. 1b). The symptoms of the disease appear
as slight vein clearing. The infection develops yellowing of
leaves which is restricted to one side of the plant or a single
shoot followed by defoliation of the older leaves. The brown-
ing of vascular tissue is the characteristic feature, which is
used as an identification tool for the disease (Ajilogba and
Babalola 2013). Since the pathogen inhabits in the soil by
3 Biotech (2020) 10:320
Fig. 1  Symptoms of Fusarium wilt disease caused by Fusarium oxysporum. f sp. lycopersici. a Field view of infected tomato plants; note the
yellowing of the oldest leaves. b Discolouration of vascular tissue in Fusarium infected tomato plants
producing resistant spore structures, it is a challenging task
to manage this disease (Sundaramoorthy and Balabaskar
2013; Hussain et al. 2016).
‘Early blight’ (EB) of tomato is another fatal disease
which occurs worldwide (Hadimani and Kulkarni 2016).
Alternaria solani (Ellis & Martin) Sorauer, is the causa-
tive organism which has been recognized as a serious
foliar pathogen of tomato (Patel et al. 2011; Ghazanfar
et al. 2016). Around 79% yield loss in tomato have been
reported in India, Canada, the United States and Nige-
ria due to the early blight damage (Maurya et al. 2015).
This disease results in yield reduction both in quantity
and quality. Under humid conditions followed by warm
and wet weather, tomato plants are susceptible to this dis-
ease (Sadana and Nidhi 2016). The disease mainly affects
the leaves, stems, flowers and tomato fruit which leads
to reduced plant health and vigour (Thakkar and Saraf
Fig. 2  Symptoms of early blight disease caused by Alternaria solani
in tomato plants. a Development of concentric rings in tomato leaves
due to early blight disease. b Early blight on tomato stem showing
Page 3 of 17 320
2015) (Fig. 2a–c). Moreover, severe infection leads to
the complete death of the plant. Sparse foliage results in
fruit sunscald, which in turn disposes the fruit quality.
Alternaria solani reproduces asexually through the pro-
duction of a reproductive structure called conidia. This
resistant structure can thrive under adverse conditions and
germinates during favourable environmental conditions.
Conidial germination requires a temperature of 8–32 °C.
The germ tube produced, invades the host tissue through
stomatal openings or wounds resulting in the establish-
ment of the infection. Lesions appear after 2–3 days of
infection and sporulation occurs after 3–5 days. The spores
or conidia disperses to other healthy plant parts through
rain splashes and wind thereby resuming the disease cycle.
This fungus leads a polycyclic disease cycle when excess
moisture occurs through rain, mist, fog, irrigation etc.
(Patel et al. 2011).
elongated concentric rings. c Early blight disease on tomato fruit
showing black spore masses
13
320 Page 4 of 17
Control measures
The ruinous diseases namely early blight and Fusarium wilt
should be effectively treated to improve the market yield
and diminish the economic loss (Hassanein et al. 2010). The
emergence of new phytopathogenic races is another threat
to deal with (Khiareddine and Riad 2015). Currently there
are different disease control strategies which comprises of
pathogen exclusion and eradication through cultural, chemi-
cal (quarantine measures and eradication) control, use of
resistant varieties, botanicals and also by native biocontrol
agents. Cultural practices include perpetuation of a patho-
gen-free condition in the field, by long crop rotation, sani-
tation, removal of affected plant debris from the field etc.
Beyond pathogen exclusion and eradication, it is arduous
to control these diseases because the soil pathogenic fungi
produce resting structures which persist for a longer period
of time. The major steps involved in the control measures of
the above-mentioned diseases are described here.
Chemical control
Today, agriculture is intensely dependent on the use of
chemical pesticides for disease control. They are routinely
and frequently applied to control fungal pathogens (Cwalina-
Ambroziak and Ryszard 2012). Application of protective
and systemic fungicides inhibit spore germination and thus
reduces disease incidence. Benomyl, carbendazim, prochlo-
raz, bromuconazole (Amini and Sidovich 2010), Nativo
(Akhtar et al. 2017) are fungicides of FOL. Fungicides
used against early blight disease include Mancozeb (Desta
and Yesuf 2015) copper oxychloride, chlorothalonil (Sahu
et al. 2013), carbendazim, antracol (Kumar et al. 2017)
Thiophanate–methyl etc. (Gharasheed 2016). Moreover,
these fungicides are applied to the field in every 7–10 days
without considering epidemiology. Eventually, the soil as
well as the crop receives an excessive amount of fungicide.
The strong demand of the public and scientific community
for eco-friendly management tends to shift from the use of
chemical fungicides. Although other disease management
practices like cultural sanitation and usage resistant breed
plants prevail, they do not exhibit complete diminishment
of these diseases. The apparition of fungicide resistance in
phytopathogen and usage of three to four fungicides in order
to avoid any fall or ineffectiveness can be a major concern.
Disease‑resistant varieties
During the nineteenth century, plants resistant to diseases
were recognized and their breeding was successfully done.
Disease resistant varieties checks the pathogen attack
13
3 Biotech (2020) 10:320
thereby restricting disease incidence. It is an economical
and environment friendly method. Likewise, it eliminates the
additional effort to prevent pathogen attack. Recent develop-
ments in molecular markers and molecular assisted selection
technology (MAs) fostered the advances in tomato genet-
ics and breeding (Foolad and Panthee 2012). Adhikari et al.
(2017) reviewed that the use of resistant varieties is an effec-
tive control measure of EB of tomato. There are a few breed-
ing experiments in the pipeline to develop resistant varieties,
however, the absence of qualitative genes and markers along
with the association of undesirable traits are the major chal-
lenges to be tackled. Hence there is no commercial cultivar
with sufficient resistance to EB. Apart from this, FOL resist-
ant varieties are available in the market (Hanson et al. 2016).
The resistant gene I-1 (S. Pimpinellifolium “PI79532”), I-2
(S. lycopersicum × S. pimpinellifolium hybrid “PI126915”),
I-3 (S. pennellii “LA716”) and I-7 (S. pennellii “PI414773”)
from wild tomato has been reported to confer wilt resistance
against different races of FOL. (Catanzariti et al. 2015; Lee
et al. 2015).
Plant extracts or metabolites as biocontrol agents
Botanical fungicides are another appealing option with a
negligible negative impact on the environment. They are
plant-derived compounds with pesticidal activity. Botani-
cal fungicides are selective and biodegradable with low
residues. Hence, these herbal products can be fostered in
organic and sustainable agriculture (Yoon et al. 2013). There
are various reports which recommend the use of botanical
fungicides. The chloroform extract of Piper betle. L. was
observed to be strongly inhibitory to FOL population when
amended in the soil (Manoj et al. 2010). Yeole et al. (2016),
reported that out of the seven plant extracts screened against
the same fungus, Syzyjium asromaticum methanolic extract
showed encouraging results with 100% inhibition to spores.
Furthermore, reduced disease incidence and lipid peroxida-
tion were found in tomato seedlings treated with aqueous
neem and willow extract from Egypt (Farag et al. 2011). The
use of plant extracts to check pathogen growth have also for-
tified these days since it does not give rise to environmental
and health hazards (Mamgain et al. 2014). Plant extracts of
Azadirachta indica, Allium sativum, Parthenium lysteropho-
rus, Datura stramonium were reported as potential growth
inhibitors of A solani (Raza et al. 2016). Ravikumar and
Garampalli (2013) have communicated the capabilities of
aqueous extract of Crotalaria trichotoma (16.6%), Azari-
dacta indica (10%), Capsicum annum (7.1%), Datura metel
(6.6%), Polyalthia longifolia (6.3%) and Citrus aurantifolia
(5.5%) as an antifungal agent against EB. Moreover, Majo-
rana syriaca and Hibiscus sabdariffa extracts have shown
strong inhibitory action to this fungus when compared to
standard fungicides (Goussous et al. 2010). Few literature
3 Biotech (2020) 10:320
reports suggest the importance of neem extract as a powerful
phytofungicide. Hassanein et al. (2010) has supported it with
the report of effective suppression of FOL and AS with dif-
ferent concentration of neem extracts. Despite these benefits,
they have limitations as well. The major challenge is that,
a huge amount of plant source needs to be exploited which
poses a threat to their population. The synthesis and purifica-
tion of these active compounds is a tedious process. When
it comes to the field, easy decomposition and short shelf
life are the other challenging factors. Moreover, the effect
of most of the botanical fungicides are moderate thereby
demanding repeated applications.
Microbes as biocontrol agents
It is desirable to control plant disease with high specificity
towards the target pathogen and with low mass production
cost (Kumar et al. 2011). There are some microorganisms
present in nature that can antagonize pests. These organisms
can be exploited as biopesticides. They include biofungi-
cide, bioherbicide and bioinsecticide. Usually, these kinds
of microbes are seen in close association with the host plant.
Hence, investigations to utilize the beneficial microorgan-
isms as biocontrol agents gain significance to subside the
fatal effects of such plant diseases (Linu and Jisha 2017).
(Fig. 3) describes the desirable characteristics of a biocontrol
agent which is in demand in the agriculture sector.
Biological control has been used over two millennia and
since the end of the nineteenth century, it has been com-
monly used in pest management. Khan et al. (2012) used
Paenibacillus lentimorbus strains for repressing early blight
disease in tomato brought about by Alternaria solani. Spe-
cific strains, such as B. subtilis, B. amyloliquefaciens, B. pas-
teurii, B. pumilus, B. mycoides and B. cereus brought about
significant reduction in various diseases by inducing sys-
temic resistance (Bouizgarne 2013). Endophytic actinomy-
cetes were utilized for the biocontrol of Rhizoctonia solani
causing damping-off in tomato. These strains significantly
Fig. 3  Schematic diagram showing desirable characteristics of a bio-
control agent (Carmona-Hernandez et al. 2019)
Page 5 of 17 320
restrained the pathogen development and enhanced the
growth parameters of tomato (Goudjal et al. 2014). Mj
et al. (2017) suggested that, successful biological manage-
ment of Fusarium wilt of tomato can be carried out using
vermicompost biofortified with selected biological control
agents (BCAs) i.e. Trichoderma harzianum, Pseudomonas
fluorescens and Bacillus subtilis. The application of Burk-
holderia gladioli pv. agaricicola strain 12322 could enhance
disease protection and improve the consistency of biological
control against tomato wilt disease caused by Verticillium
dahlia (Elshafie et al. 2017). Lian et al. (2017) proved that,
Streptomyces pratensis LMM15 could be a potential bio-
control agent for controlling tomato gray mold because the
incidence of tomato grey mold decreased by 46.35% in asso-
ciation with an increase in proline content and malondial-
dehyde (MDA) and changes in the defence-related enzymes
on tomato leaves were observed when the strain was sprayed
on the tomato leaves 24 h prior to inoculation with patho-
gens. Arenas et al. (2018) evaluated the efficacy of Mexican
strains of Trichoderma spp. and its antagonistic effect on F.
oxysporum on tomato seedlings. It was observed that the
Trichoderma harzianum strain presented the highest growth
rate with a mean of 1.25 cm/day, proving to be the most
aggressive strain to control F. oxysporum with a develop-
ment rate of 3.80 mm/day.
The types of biological control can be classified into natu-
ral, conservation, inoculative (classical) and augmentative
biocontrol. Pest reduction in natural biocontrol has been
occurring since evolution using natural enemies, whereas
conservation biocontrol involves human actions to stimulate
and protect the performance of the same. Inoculative (clas-
sical) mode is the first type of biocontrol being practised
widely, in which natural enemies are released into new areas,
where the pest was accidentally introduced. Massive rearing
of natural enemies in biofactories and its release into the
market for immediate pest control made augmented biocon-
trol more desirable (van Lenteren 2012). This approach is
considered to be an environment-friendly and food-hygien-
ically-safe plant protection method (Sandheep et al. 2012).
Furthermore, understanding the mechanisms of biological
control through the interactions between the biocontrol
agents and the pathogens may help in improving and devel-
oping biocontrol strategies.
Plant growth‑promoting rhizobacteria:
tapping for BCA
The rhizosphere is the narrow region of soil surrounding
the living plant roots that are directly influenced by root
secretions and associated soil microorganisms. It is a zone
of mutual cooperation between plant, soil and microorgan-
isms, such as biochemical interactions and exchange of
13
320 Page 6 of 17
signal molecules, nutrient transformation, genetic exchange,
the release of root exudates, etc. (Haldar and Sanghamitra
2015). Hence, it is known as a unique niche for microbial
activities and diversities (Venant et al. 2011). The diverse
group of bacteria colonizing the rhizospheric habitat are
called rhizobacteria. They are potent microbial competitors
in the root zone. They influence plant growth directly or
indirectly (Tank and Meenu 2010; Beneduzi et al. 2012).
The direct mechanism involves the beneficial activities
done by PGPR that support the plant growth directly. These
mechanisms promote plant growth, but the ways in which
it influences will vary species to species as well as strain to
strain (Kundan et al. 2015). The PGPR nurture the plant by
transforming the nutrients present in the soil through bio-
geochemical cycling. They also facilitate transport of these
nutrients into the plant which aids plant growth directly.
They enhance plant growth directly by producing plant hor-
mones, (Glick 2014; Miransari and Smith 2014; Damam
et al. 2016), fixing atmosphere nitrogen (Singh et al. 2015),
solubilizing minerals, such as phosphorous (Kannapiran and
Sri 2011; Stephen and Jisha 2011; Sagervanshi et al. 2012;
Karpagam and Nagalakshmi 2014; Mahantesh et al. 2015;
Sanjoth and Sudheer 2016), synthesizing siderophores that
may solubilise and sequester iron (Radzki et al. 2013) and
avail nutrients to plants beyond controlling soil-borne plant
pathogens (Akhtar et al. 2012; Nadeem et al. 2014). In addi-
tion to plant growth promotion, it also has a significant role
in resisting the phytopathogenic microorganisms (Son et al.
2014). Chen et al. (2013) discovered that Bacillus subtilis
exhibited above 50% biocontrol efficacy on tomato plants
against the plant pathogen Ralstonia solanacearum under
greenhouse conditions through robust biofilm formation.
Abdallah et al. (2016) inoculated seven distinctive endo-
phytic strains isolated from the native Nicotiana glauca
plants and discovered 88–94% significant reduction in yel-
lowing and wilt symptoms and 95–97.5% in vascular brown-
ing of tomato plants. Antifungal activity of several Bacil-
lus sp. against plant pathogenic fungi were accounted for,
elevating them to be utilized as promising candidates for the
biological control (Attia et al. 2020). You et al. (2016) dis-
covered Trichoderma-mediated growth inhibition of Botrytis
cinerea and their application in soils promoted growth of
tomato. A combination of biocontrol isolates Mitsuaria sp.
TWR114 and nonpathogenic Ralstonia sp. TCR112 exerted
a synergistic suppressive effect resulting in enhanced bio-
control efficacy against tomato bacterial wilt (Marian et al.
2019). Maung et al. (2017) demonstrated the effectiveness
of Bacillus amyloliquefaciens Y1 not only in the control of
Fusarium wilt disease, but also for the enhancement of plant
growth in cultivated tomato. Elsayed et al. (2020) revealed
that the strains, Bacillus velezensis B63 and Pseudomonas
fluorescens P142 were promising candidates for biocontrol
of bacterial wilt caused by Ralstonia solanacearum under
13
3 Biotech (2020) 10:320
field conditions through significantly lowered R. solan-
acearum densities in tomato shoots and in the rhizosphere.
Biocontrol mechanisms (indirect mechanisms)
PGPB as BCA has certain advantages over other disease
control methods such as they are eco-friendly and non-toxic
indigenous microorganism and its application is sustaina-
ble to both environment and human health. The important
mechanisms involved in the antagonism by BCA involves
antibiosis, competition, induced systemic resistance of the
host plant, hydrolytic enzyme production, HCN production
and siderophore production (Bhattacharyya and Jha 2012)
(Fig. 4). In addition, they can alleviate various stress condi-
tions, such as salinity, drought, flood, heavy-metal toxic-
ity, etc. in plants, thus empowering them to survive in such
stress conditions (Heidari and Amir 2012). Even though sev-
eral free-living rhizobacteria are considered as plant growth
beneficial rhizobacteria, all the strains under the same spe-
cies do not possess the same metabolic capacities to enhance
plant growth. It is significant to know the potentialities of
rhizosphere microbiota besides its mechanism of action
involved in sustainable crop production (Bhattacharyya and
Jha 2012).
The antagonistic traits exhibited by the PGPR over vari-
ous pathogens augment possibilities for their use as bio-
control agents (Pathak et al. 2017; Tariq et al. 2017). Well-
known BCAs in suppressing tomato disease include strains
of Pseudomonas (Toua et al. 2013), members of the genera
Arthrobacter, Azoarcus, Azospirillum, Bacillus, Burkholde-
ria, Enterobacter, Gluconacetobacter, Herbaspirillum, Kleb-
siella, Paenibacillus, Pseudomonas and Serratia (Mahesh-
wari 2011). Various literatures suggest that consortium of
Fig. 4  Different mechanisms by which plant growth-promoting rhizo-
bacteria (PGPR) accomplish biocontrol against phytopathogens
3 Biotech (2020) 10:320
BCA helps to reduce disease incidence through synergistic
action. It has been reported that Bacillus subtilis, Bacillus
megatarium, Bacillus polymyxa, Pseudomonas fluorescens
and Trichoderma harzianum in combination with dinitrogen
fixers of Azotobacter sp and Azospirillum sp were identified
as successful biocontrol agents against soil-borne pathogens
F oxysporum and Verticillium dahliae causing fungal wilt
disease of tomato (Saad et al. 2016). Serratia plymuthica,
Bacillus coagulans, Paenibacillus macerans, Bacillus
pumilis and Pantoea agglomerans were tested for inhibi-
tory effects on Alternaria solani, early blight pathogen on
tomato (Yazici et al. 2011).
Antibiosis
Antibiotics are chemical substances produced by microor-
ganisms against microorganisms. Interaction between the
organisms leads to the production of these chemical sub-
stances in order to survive in predation, competition etc.
(Ulloa-Ogaz et al. 2015). Bacterial antagonists impose sup-
pression of phytopathogens by extracellular secretion of
these metabolites that have inhibitory property, even at low
concentration (Goswami et al. 2016). In fact, it is an effi-
cient and most effectively studied characteristic of biocontrol
(Ramadan et al. 2016).
The well-characterized antibiotics for biological control
are 2,4 diacetylphologlucinol (DAPG), phenazine, pyrrol-
nitrin, pyoluteorin, tensin, tropolone, oomycin A, cyclic
lipopeptides and HCN (Babalola 2010). The majority of
Bacillus antibiotics shows activity against plant pathogenic
fungi Alternaria solani and Fusarium oxysporum (Mak-
simov et al. 2011). HCN production mediated biocontrol
has been proved by B. subtilis against FOL (Akintokun and
Taiwo 2016). However, B. subtilis was reviewed as a poten-
tial BCA, capable of inhibiting the growth of fungal patho-
gen due to their ability to produce a vast array of antibiotics,
such as zwittermicin, bacillomycin, fengycin, bacilysin and
difficidin (Mangalanayaki et al. 2016). Jain and Das (2016),
reviewed the positive correlation between HCN production
and plant protection in tomato wilt disease. Bacillus isolate
TNAM5 was found to be effective in suppression of FOL
by the production of diffusible and volatile antifungal com-
pound ammonia and HCN (Prashar et al. 2013). Similarly,
it was observed that B. subtilis inhibited AS by the secretion
of antifungal metabolites (Bellishree et al. 2015). A similar
result was observed by Phichai (2014), against Alternaria
spp. wherein B. subtilis could secrete several antifungal
metabolites such as subtilin, bacitracin, bacillin and bacillo-
mycin which had an inhibitory effect on the fungal pathogen.
However, too much dependence on antibiotic-producing
bacteria as a BCA may be a disadvantage since there is a
problem of antibiotic resistance. In order to overcome this
threat, researchers are using potent biocontrol strains that
Page 7 of 17 320
could synthesize one or more antibiotics such as Bacillus
and Pseudomonads (Glick 2012; Beneduzi et al. 2012).
The biological control of plant phytopathogens by endo-
phytes was reported in the late 50s where, a Micromono-
spora isolate from tomato showed antagonistic activity
against Fusarium oxysporum f.sp. lycopersici (Manikprabhu
and Li, 2016). Bacterial endophytes such as Bacillus and
Streptomyces species (Frank et al. 2017) both exhibited sec-
ondary metabolites showing antimicrobial activity against
plant pathogens. As a result, Bacillus spp. endophytes were
proposed for crop management (Aloo et al. 2018). Simi-
larly, Streptomyces spp. endophytes are widely reported as
phytopathogens biocontrol agents. For example, Kennedia
nigriscans-endophytic Streptomyces sp. strain NRRL 30562
was recently reported to produce antibiotics as munumbicins
A, B, C and D active against plant pathogenic bacteria and
fungi (Castillo et al. 2002). Mohamad et al. (2018) evaluated
the antimicrobial activity of endophytic bacterial popula-
tions from Chinese traditional medicinal plant licorice and
characterized their bioactive secondary metabolites against
Verticillium dahlia. The results revealed that the genus
Bacillus, particularly B. atrophaeus and B. mojavensis, were
the most effective biocontrol agents with most strains exhib-
iting broad antibacterial and antifungal activities. Around
13 compounds were produced during co-cultivation with
V. dahlia which included putative compounds possessing
antimicrobial activity such as 1,2-benzenedicarboxylic acid,
bis (2-methylpropyl) ester; 9,12-octadecadienoic acid (Z,Z)-,
methyl ester; 9- octadecenoic acid, methyl ester, (E)- and
decanedioic acid, bis(2-ethylhexyl) ester.
Competition
The BCA should be able to withstand and multiply in a natu-
ral environment since effective colonization and enhanced
competition play a significant role in biocontrol (Raguchan-
der et al. 2011). For the successful establishment on rhizos-
phere, microorganisms must effectively compete for nutrient
availability and niche. Competition among pathogenic and
non-pathogenic microbes is an essential matter in biocon-
trol (Heydari and Mohammad 2010). It has been known
that plant associate microbes give protection to the plant
by accelerated rhizosphere colonization than the pathogen.
The biocontrol agents deplete the limited available substrates
making it unavailable to the pathogens. Simultaneously, they
produce metabolic compounds that are detrimental to the
pathogens (Trapet et al. 2016; Khilyas et al. 2016; Tabassum
et al. 2017).
According to Khan et al. (2012), the antagonism depicted
by Paenibacillus lentimorbus against AS was enforced due
to strong competition between them for niche and nutrient
utilization. Lugtenberg et al. (2001) revealed that the tomato
foot and root rot pathogen FOL reside deep inside the soil.
13
320 Page 8 of 17
Hence, an efficient BCA must be able to protect deeper root
parts by effective colonization to reach the growing tips and
suppress the pathogen. Pseudomonads are reported as highly
efficient competitors for root exudates among the rhizobac-
terial communities (Barea et al. 2005). Some of the root
exudates include antimicrobial compounds which provides a
suitable ecological niche for PGPR which in turn can detox-
ify them. This implies that PGPR competence is majorly
reliant on their ability to take advantage of the specific envi-
ronmental condition or to get adapted to variable circum-
stances (Compant et al. 2005). Kuiper et al. (2001) dem-
onstrated that the competitive colonization ability of root
colonizing Pseudomonads was considerably dependent on
their increased uptake of a tomato root exudate, putrescine.
In addition, Prasanna et al. (2013) reported that competitive
colonization of Anabaena variabilis RPAN59 and A. laxa
RPAN8 were positively correlated with the reduced fun-
gal population of FOL. Biocontrol Bacillus strains namely
3F-II, 3F-VII, 13F-III, 15F-III inhibited EB in tomato and
the enzymatic profile indicated that they were potential eco-
logical competitors (Pane and Zaccardelli 2015).
Apart from competitive root colonization, another impor-
tant mechanism of pathogen suppression via nutrient compe-
tition by PGPR involves the secretion of compounds, such as
siderophores that efficiently sequester iron and deprive the
pathogen from this important element (Singh et al. 2017).
Several siderophore-producing rhizobacteria were evaluated
and reported as biocontrol agents including species of Pseu-
domonas (Weller 2007), Bacillus and Enterobacter (Solanki
et al. 2014). Heidarzadeh and Baghaee-Ravari (2015) proved
that siderophore production by B. pumilus played a crucial
role in suppressing Fusarium wilt disease by depleting iron.
Induced systemic resistance
Plant beneficial bacteria in the rhizosphere interact with the
host plant to stimulate the defence against various patho-
gens. Induced resistance (IR) is the enhanced physiologi-
cal state of defence, elicited by broad-spectrum biotic and
abiotic stimuli. Induced resistance is classified into Induced
Systemic Resistance (ISR) and Systemic acquired resistance
(SAR). When plants innate defence mechanisms are evoked
due to biotic challenges (Choudhary and Johri 2009), it is
known as ISR. In SAR, plants render more resistance to
uninfected plant parts while encountering broad spectrum
of pathogens (Pieterse et al. 2014). SAR occurs through
salicylic acid-mediated signalling pathway whereas ISR
occurs predominantly through jasmonate- or ethylene-sen-
sitive pathway (Fu and Dong 2013). Plant growth beneficial
rhizobacteria colonizing plant roots can evoke resistance
against a broad spectrum of plant diseases. This strategy
has gained attention over the recent years. Few literatures
have suggested that, some strains of Pseudomonas, Bacillus,
13
3 Biotech (2020) 10:320
Serratia and Trichoderma elicit ISR against different patho-
gens (Mandal and Ramesh 2011). BCA demonstrates dis-
ease suppression caused by fungal, bacterial, viral and in
some cases insects and nematodes by eliciting such induced
resistance.
It is evident that the Bacillus genus is an ideal biocontrol
agent and the mostly exploited biopesticide to control plant
diseases (Ongena et al. 2007). Pseudomonas also induces
systemic resistance in plants. It was observed that Pseu-
domonas fluorescens protected tomato plants from the path-
ogens F. oxysporum and Phytophthora infestans by means
of systemic resistance (Santoyo et al. 2012). Ongena et al.
(2007) provided strong evidence of ISR mediated biocontrol
of tomato gray mold disease by the use of potential B. sub-
tilis, an efficient lipopeptide producer. Pseudomonas gladi-
oli triggered defence molecules against Alternaria solani
through induced systemic resistance in tomato (Jagadeesh
and Jagadeesh 2009). Khan et al. (2012) assessed the ability
of Paenibacillus lentimorbus to control early blight disease
in tomato by inducing host resistance. Induction of sys-
temic resistance has been established by the application of
vermicompost biofortified with bioagents—Bacillus subti-
lis and Pseudomonas fluorescens against F. oxysporum by
the accumulation of PAL, PO, PPO free phenol and SOD
in tomato plants (Mj et al. 2017). B. cereus, Streptomyces
cereus and S. marcescens were proved to be effective in
host’s resistance against wilt disease by the activation of
defence enzymes peroxidises (POX), polyphenol oxidases
(PPO), glucanases (GLU), chitinases (CHI), phenylalanine
ammonia-lyases (PAL) and lipoxygenases (LOX) (Gledson
et al. 2014). P. fluorescens has also been reported as a strong
BCA against the pathogen by inducing key enzymes POX,
PPO and superoxide dismutase (SOD), β-1,3 glucanases
(Dorjey et al. 2017). Reduced disease severity coupled
with enhanced enzyme production elicited by B. subtilis, B.
atrophaeus and Burkholderia cepacia mixtures indicated its
mode of action for the vascular suppression through direct
biocontrol and ISR (Shanmugam and Kanoujia 2011).
P. putida and P. syringe stimulated a systemic response
against A. solani by inducing high rates of enzyme activity
of PAL, PO, PPO as well as the accumulation of phenolics
(Ahmed et al. 2011). Similarly, Chowdappa et al. (2013)
reported that B. subtilis OTPB1 isolate was able to inhibit A.
solani by enhanced systemic resistance. The defence-related
enzymes, PO, PPO and SOD were significantly higher in
inoculated tomato seedlings, compared to uninoculated
control. Indigenous isolate P. fluorescens TK3 was recom-
mended as the best biocontrol agent for early blight patho-
gen (Moges et al. 2012). The study reviewed that disease
suppression of Pseudomonad was by induction of systemic
resistance. Exopolysaccharides (EPS) are polysaccharides
synthesized by bacteria which are secreted in the exter-
nal environment. EPS helps to endure drought condition,
3 Biotech (2020) 10:320
protection from stress (Qurashi and Anjum 2012) and
defence against phytopathogens (Tewari and Naveen 2014).
The significant role of bacterial EPS as an elicitor for the
induction of systemic resistance has already been reported
in different crop species. It is reported that application of
Bacillus EPS obtained from natural biofloc at 200 ppm could
effectively reduce the wilt disease incidence (Thenmozhi
and Dinakar 2014).
Hydrolytic enzyme production
Production of hydrolytic enzymes is an efficient mechanism
to eradicate the pathogen growth through lysis of pathogenic
cell wall. Different polymeric compounds present in fungal
cell wall, such as cellulose, hemicelluloses, chitin, proteins
and DNA can be degraded by producing lytic enzymes such
as cellulase, chitinase, protease, etc. The cell wall degrada-
tion of fungal pathogens by the secretion of these enzymes
is an antifungal activity aiding in the biocontrol of phy-
topathogens (Jadhav et al. 2017; Mabood et al. 2014; Aeron
et al. 2011). The soil-borne fluorescent Pseudomonas has
gained more attention in this regard because they can pro-
duce different kinds of cell wall degrading enzymes, such
as chitinase, protease/elastase and β-1,3 glucanase (Kapoor
et al. 2012). This mechanism helps to parasitize the phy-
topathogen directly.
Streptomyces sp. is a promising BCA which possess cel-
lulolytic, chitinolytic and xylanolytic activity. They are iden-
tified as plant growth promoting and disease control agents
in tomato (Da et al. 2008). Bacillus subtilis, B. cereus and B.
thuringensis were reported to produce hydrolytic enzymes
for the biocontrol of phytopathogen F. oxysporum. They
produced swelling and curling in the hyphae which further
caused the hyphal tip to burst (Jadhav and Sayyed 2016).
Chitinase, β-1,3 glucanase and β-1,4 glucanase produced by
Pseudomonas fluorescens and Bacillus subtilis were found to
inhibit pathogenic fungi causing root rot disease in tomato
crop (El-Gamal et al. 2016). Prasanna et al. (2013) reported
that B. subtilis produced hydrolytic enzymes β-1,4 glucanase
and chitosanase which were fungicidal to FOL. Evaluation
of extracellular lytic enzymes viz. chitinase, β-1,3 glucanase,
protease and cellulase from indigenous Bacillus sp. isolated
from tomato rhizosphere provided potential bioresource for
the benefit of the agricultural industry (Praveen et al. 2012).
Seed bacterization and soil application of B. atrophaeus
S2BC-2 reduced disease incidence in plants challenged with
FOL and AS. Maximum induction of chitinase and β-1,3
glucanase were found in the leaf and root sample analysis
(Shanmugam and Kanoujia 2011).
Application of PGPR has extended to remediate the con-
taminated soil enabling the plant to survive such stress con-
ditions. The metal resistant PGPB have also been reported
to produce enzymes that lyse the cell of the fungal pathogen
Page 9 of 17 320
(Ma et al. 2010). Besides the above-mentioned biocontrol
mechanisms of several rhizobacterial strains, the effective-
ness of these strains depends upon the host plant and soil
characteristics. Moreover, their inherent capacities and
rhizospheric competence also plays a major role in exhibit-
ing their biocontrol traits (Gamalero et al. 2010).
HCN production
Biogenic cyanogenesis (HCN) has been found in rhizobac-
teria, by de novo synthesis (Rijavec and Lapanje 2017).
This volatile antimicrobial compound is known for its role
in disease suppression (Jain and Das 2016). HCN is likely to
inhibit electron transport chain and energy supply to the cell
eventually leading to cell death. It is also known to inhibit
the action of cytochrome oxidase (Heydari and Mohammad
2010). This deleterious property allows PGPR to gain a
competitive advantage over fungal pathogens and it can be
further exploited in biocontrol of plant diseases (Ramette
et al. 2003; Hayat et al. 2010). Sehrawat and Sindhu (2019)
reviewed that rhizosphere inhabiting bacteria such as Bacil-
lus and Pseudomonas produces HCN as their secondary
metabolite. Furthermore, members of the genus Chromobac-
terium, Burkholderia, certain Rhizobia and Cyanobacteria
were also reported to show bacterial cyanogenesis (Ahemad
and Kibret 2014). However, fluorescent Pseudomonas is pre-
dominantly reported as HCN producer (Mishra and Arora
2018). Most studies shows the activity of HCN producing
bacteria towards fungal pathogens (Siddiqui and Shaukat
2002; Ramette et al. 2006). The study of Islam et al. (2019)
reported 44.99% disease inhibition of Fusarium wilt dis-
ease treated with Brevundimonas olei Prd2 which showed
HCN production in vitro. Additionally, Lachisa and Dabassa
(2016) reported that the growth reduction of FOL is due to
the production of HCN by the isolate Pseudomonas sp. RhB-
12. Furthermore, Someya et al. (2006) confirmed that the
HCN-producing Pseudomonas fluorescens strain LRB3W1
played a key role in the inhibition of disease development
by FOL.
Siderophore production
Iron is an essential micronutrient for all forms of life and is
present in the soil in the form of F ­ e3+ or ferric ion predomi-
nantly, which is sparingly soluble. Hence, it is not readily
assimilated by plants (Colombo et al. 2014). Plants can use
siderophores produced by microorganisms for iron uptake
(Shirley et al. 2011). Rhizobacteria produces siderophores
that are low molecular weight iron-chelating compounds
with great affinity and selectivity to bind and form a com-
plex Fe(III) (Ferreira et al. 2019). It acts like a ligand that
facilitates the sequestration and transportation of iron into
the cell. This particular trait has drawn attention in the recent
13
320 Page 10 of 17
years and the identification of such siderophore produc-
ing PGPR along with its in vivo testing has been reported
(Ghavami et al. 2017; Liu et al. 2017; Sabate et al. 2017).
Siderophores are stable complexes and different kinds of
siderophores are reported including hydroxamates, phe-
nolcatecholates and carboxylates. (Kundan et al. 2015).
Siderophore mediated iron transfer system is better studied
in gram-negative PGPR transport systems rather than gram-
positive transport systems. Among 500 different types of
siderophores known, 270 have been structurally character-
ized (Hider and Kong 2010). Siderophores have been sug-
gested to be an eco-friendly alternative to harmful pesticides
(Schenk et al. 2012).
Because siderophores produced by PGPR show greater
affinity for iron as compared to fungal pathogens, they
have a competitive advantage for efficiently suppressing
phytopathogen proliferation (Schippers et al. 1987). Even-
tually, the fungal pathogen under iron scarcity are unable
to proliferate and are excluded from the ecological niche.
Hence, siderophore production is a desirable characteristic
of PGPR as a biocontrol agent. Among the various bacterial
siderophores, those produced by Pseudomonas are known
to be with higher affinity (Beneduzi et al. 2012). There are
various reports on biocontrol efficiency of siderophore pro-
ducing PGPR. Arya et al. (2018) reported that siderophore
producing Pseudomonas strains SPs9 and SPs20 suppressed
Fusarium wilt of tomato under field conditions. Streptomy-
ces strain SNL2 with siderophore producing ability was also
reported which reduced wilt disease incidence by 88.5%
(Goudjal et al. 2016). Segarra et al. (2010) also reported
that Trichoderma asperellum producing siderophores
restrict FOL. Similarly, Pseudomonas aeruginosa strains
JO and JO7, capable of producing siderophore inhibited AS
and FOL (Paramanandham et al. 2017). Bacillus subtilis
SBMP4, Lysinibacillus fusiformis NBRC 15717 and Achro-
mobacter xylosoxidans NBRC15126 produced hydroxamate
siderophore which had a vital role in suppression of early
blight in tomato (Attia et al. 2020).
Consortium of BCA
Inconsistent performance of BCA is the primary barrier
for its commercialization. A single organism may fail dur-
ing adverse environmental conditions, so a combination
of more than one BCA is appreciated. There is a chance
for increased, neutral and decreased biocontrol efficiency
when more than one BCA is used together. Moreover, they
can easily colonize the rhizosphere. It is reported that, con-
sortium of BCA helps to reduce disease incidence through
synergistic action. Kannan and Sureendar (2009) proved
the efficiency of consortial treatment in growth promotion
and wilt resistance of tomato. Unfortunately, the interac-
tions involved were not clearly determined and hence they
13
3 Biotech (2020) 10:320
need further clarification. A synergistic study among biocon-
trol agents Bacillus subtilis and Pseudomonas fluorescens
displayed increased plant growth promotion more than its
biocontrol activity on early blight disease (Suleiman et al.
2017). A thorough understanding of biocontrol mechanisms
is therefore essential for employing this method in plant dis-
ease control.
These drawbacks have led to a slow progress in the suc-
cessful development and marketing of bio-formulations.
However, the use of a mixture of BCA for biocontrol were
highly encouraged in the recent decades because it helped
to defeat the efficacy problems. Thilagavathi et al. (2007)
reported that a combination of BCA Pseudomonas fluores-
cens pf1 and Trichoderma viride tv1 improves the manage-
ment of dry root rot in green gram. The activity of defence
enzymes was significantly greater in treated plants receiving
consortium treatments.
Commercialisation of BCAs
Commercialization of BCA is a multistep process that
involves isolation and screening of potential antagonistic
bacteria from natural ecosystems, testing the efficiency of
the isolate in the field, mass production, formulation, toxic-
ity studies, delivery, compatibility, registration and release
(Junaid et al. 2013). Recent reports claims that bioformula-
tions were prepared from BCAs such as Bacillus cereus,
Pseudomonas rhodesiae, Pseudomonas chlororaphis, Pseu-
domonas fluorescens, Bacillus coagulans which acts as both
nutrient and growth enhancer to selected crops (Kalita et al.
2015; Jorjani et al. 2011). Bioformulations are defined as
the preparation of microorganism(s) that may be a partial
or complete substitute for chemical fertilization/ pesticides
(Arora et al. 2010). In this approach, the degree of success
depends upon the consistent broad-spectrum action, eco-
nomical and viable market demand, safety, stability, longer
shelf life, low capital costs and easy availability of carrier
materials. Bioformulations consists of an active ingredient
i.e. a viable organism which may be a microbe/spore and
an inert carrier material that supports the active ingredi-
ent/cells (Mishra and Naveen 2016). Selection of a superior
carrier material is essential as it is expected to meet the pre-
requisites, such as high water holding capacity, high water
retention capacity, no heat production while wetting condi-
tion prevails, sterility, chemically and physically uniform in
nature, biodegradable, non-polluting and nearly neutral pH
that supports bacterial viability (Singh et al. 2015). Avail-
able carrier materials are talc, sawdust, fuller’s earth, rice
husk, sugar cane, bagasse, charcoal and wheat bran (Singh
et al. 2014).
Before developing a biocontrol agent into a commer-
cial product, an intensive knowledge on several factors
such as phytopathogen species, type of hosts it attacks,
3 Biotech (2020) 10:320
epidemiology of the disease, resistance of phytopathogen
and environmental conditions under which the BCA will
be used must be known (Carmona-Hernandez et al. 2019).
Talc based formulation of B. subtilis PSIRB2 and P. aerugi-
nosa 2apa were found to significantly exhibit disease protec-
tion traits against fusarium wilt and early blight diseases of
tomato (Gowtham et al. 2017). Similarly, talc-based formu-
lation of P. fluorescens delivered through farmyard manure
and vermicompost were tested against Fusarium oxysporum
and Alternaria solani (Kaur et al. 2016).
Unfortunately, the successful establishment of such bio-
logicals in agriculture is hampered by the inadequate knowl-
edge about the plant–microbe interactions, though there
were some progress in the past decade (Ravensberg 2015).
For a fruitful PGPR inoculum, the chosen strain ought to
colonize the plant roots, should be able to persevere in the
rhizosphere and ought not to be a potential hazard for the
environment and human wellbeing. Certain bacterial strains
display promising impacts in advancing plant development
in lab conditions yet fail in field trials because of their insuf-
ficiency in colonizing the rhizosphere and plant roots. These
strains are unable to adjust with the plant microenvironment
and consequently, incapable to rival with the current local
bacterial network for foundation and perseverance in the
soil.
Henceforth, investigating the components of PGPB would
uncover new experiences to structure systems for improving
the efficacy of biocontrol agents.
Conclusion
The present review indicates that plant growth-promoting
rhizobacteria (PGPR) not only triggers different biological
promotional effects on tomato plant growth parameters, but
also helps in protecting the plant from several pathogens
by acting as biocontrol agents. To replace chemical pesti-
cides with the system of a biopesticide, it is ought to be
progressively more effective and economical. Before com-
mercialization, molecular analysis can result in stabilizing
the effects of PGPR in biological control and possible risk
assessments. Effective usage of PGPR for disease reduction
or crop protection within the future will demand a rational
selection of organism as well as technical enhancements in
upscaling and formulation techniques. Genetic engineering
of PGPR may intensify the expression of plant growth and
health-promoting genomic products. Therefore, a single
bacterial strain or a consortium with variable attributes will
mitigate the pathogen assault and promote plant growth to
encourage the producers. It has become clear that PGPR
strains employ several mechanisms to act as an effective
biocontrol agent, although studies ought to be centered on
the relative contribution of every mechanism responsible for
Page 11 of 17 320
its significant biocontrol activity. A thorough understanding
of the plant–microbe interactions and processes as well as
exploiting the microbial ecology in the soil and rhizosphere
would help us reveal the multiple facets of disease suppres-
sion by these biocontrol agents. Moreover, carefully con-
trolled field trials of tomato plants inoculated with PGPR
inoculants is obligatory for maximum commercial exploi-
tation of these strains. In conclusion, the success of micro-
bial inoculant-producing industries, notably those utilizing
PGPRs, will depend on measures including product market-
ing and extensive research. In addition, for better fermenta-
tion and formulation processes, the optimization of PGPR
strains will be required to introduce them in the agriculture
industry.
Acknowledgements We would like to thank School of Biosciences,
Mahatma Gandhi University for providing necessary facilities. The
first author is greatly thankful to the University Grants Commission,
Government of India for the financial support in the form of Senior
Research Fellowship, Vide Sr. No.2061530793.
Author contributions SK and SV conceptualized and designed the
manuscript; MSJ reviewed and approved the manuscript.
Funding The first author is very much thankful to the University
Grants Commission, Government of India for the financial support in
the form of SRF, Vide Sr. No.2061530793.
Compliance with ethical standards
Conflict of interest The authors declare that there are no conflicts of
interest regarding the publication of this manuscript.
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