Potato Research

https://doi.org/10.1007/s11540-023-09648-2

Utilization of Plant Growth‑Promoting Rhizobacteria

(PGPR) for Managing Recently Reported Potato Cyst
Nematodes, Globodera spp. in North Himalayan Regions
of India

Vijeta Katoch1 · Shavnam1 · Saurabh Sharma2 · Manisha Negi2

Received: 15 September 2022 / Accepted: 23 July 2023

© The Author(s), under exclusive licence to European Association for Potato Research 2023

Abstract
Globodera rostochiensis (Woll.) and Globodera pallida (Stone), two potato cyst
nematodes (PCNs), are significant pests worldwide. The first PCNs from North Him-
alayan regions of India were detected in 2010 in Shimla, Himachal Pradesh (HP).
Despite a domestic quarantine, succeeding surveys in the mountainous regions of
North India have shown the presence of PCNs in numerous districts of HP, Jammu
and Kashmir and Uttarakhand. The seed plot technique is utilized in North Himala-
yan regions of India to create virus-free seeds and the state agricultural departments
distribute it to farmers in other parts of the nation. Therefore, the entire nation should
be concerned about the emergence of such a significant potato seed location as the
primary sites for large PCN populations. Inorganic pesticide based nematode man-
agement is an effectual way to address this biotic stress but improper use of chemi-
cals can have a negative impact on environment. Therefore, the farming community
must prefer the employment of potential biocontrol agents, this might aid in fend-
ing off regulatory and environmental pressure. Plant growth-promoting rhizobacte-
ria (PGPR) enabled control of PCNs can be environmentally benign. In light of this,
we undertook an extensive and thorough review of the available literature pertain-
ing to PGPRs. Our analysis uncovered that PGPRs can greatly assist in enhancing
plant resilience to various stressors that can impede their growth and yield. However,
achieving these benefits will require successful development and commercialization
efforts. Furthermore, before PGPRs can be widely employed, it is crucial to tackle the
challenges related to their selectivity and restricted range of activity.

Keywords Enzymes · Hyperparasitism · Induced systemic resistance · Phytohormone

* Shavnam
[email protected]
1
Department of Plant Pathology, Dr. YS Parmar University of Horticulture and Forestry, Nauni,
Solan 173230, Himachal Pradesh, India
2
Department of Soil Science and Water Management, Dr. YS Parmar University of Horticulture
and Forestry, Nauni, Solan 173230, Himachal Pradesh, India

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Introduction

Potato (Solanum tuberosum) is cultivated throughout the globe and is one of the
most important crops for human nutrition (CIP 2020). It offers more protein and
minerals than any other type of staple crop and it is abundant in carbohydrates
(Beals 2019). Optimum pest and disease control are essential for the sustainable
agricultural production following the rising demand from a growing global popu-
lation. Plant parasitic cyst nematodes (PCN) namely Globodera pallida (Stone)
and G. rostochiensis (Woll.) cause huge economic losses in potato crops (Aires
et al. 2009). These are soil inhabitors, biotrophic sedentary endoparasites as path-
ogens specific to the Solanaceae family (Sabeh et al. 2019; Gartner et al. 2021).
PCN are temperate species and attains high population densities in cool climates
on the host plant. In warm or tropical areas, the PCN find it difficult to complete
their life cycle because the annual egg mortality rate in warmer geographic areas
is more than 75% as compared to 50% in temperate regions. As a result, temper-
ate regions must be the main focus of PCN investigations.
These nematodes (worms) have a very severe negative impact on tuber produc-
tion and quality (Brodie et al. 1993; Marks and Brodie 1998). The PCN are the
most strictly controlled nematodes because of their capacity to form cysts that are
passively transferred over great distances with contaminated or infected plant root
remnants, tubers and soil debris (Sullivan et al. 2007). According to estimates,
PCN can result in yield losses of 19 to 80%, based on the variety, seasonality and
soil population of PCN (Turner and Evans 1998). Potato yields may fall short
of the tonnes per hectare of the seed planted when PCN numbers are large in
the field (Mai 1977; Whitehead and Turner 1998). According to Contina et al.
(2020), PCN species may reduce potato output by up to 80% if left unchecked,
resulting in significant financial losses for the global potato industry.
For researchers working on sustainable agriculture, controlling PCN has
become a difficult task. In the past, nematodes were controlled with a variety
of synthetic chemical nematicides, including carbamates, organophosphates
and fumigants. The demand for more effective nematode control strategies has
increased as a result of the large decline in pesticide use caused by European
Union (EU) rules (EC No. 1107/2009), which show that pesticides are detrimen-
tal for human health and the environment (Zhang et al. 2017). Chemical nemati-
cides expanded agrarian and economic possibilities by boosting food and fibre
production at the expense of human health and the environment (Akhtar and Sid-
diqui 2009). Plant growth-promoting rhizobacterial (PGPR) variants can operate
as an effective nematode biocontrol agent (BCA) and environmentally friendly
methods and also act as plant growth promoter for increased yield (Mhatre et al.
2018). Recognizing the communities of benevolent microorganisms and their
modes of action for regulating PCNs will give a ground for enhancing the pos-
sible biocontrol strains’ pathogenicity and create cutting-edge biocontrol methods
for PCN management. This review aims in utilization of PGPR for management
of PCN.

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Occurrence

The cyst nematodes are the primary source of crop losses in the potato (Solanum
tuberosum L.) industry (Van Riel and Mulder 1998). These are important quarantine
pests worldwide originated from the Andean Mountains of South America. Hafez
and coworkers (2007) for the first time reported the PCN species G. pallida in USA.
Occurrence and distribution of G. rostochiensis (pathotype Rol) were the most abun-
dant in Northern Ireland (Zaheer et al. 1993), whereas in England and Wales rather
than G. rostochiensis another PCN species, i.e. G. pallida was predominant (Minnis
et al. 2000; Karnkowski et al. 2011). Similarly, in Algeria (Africa), the most diver-
gent PCN species was G. pallida (Tirchi et al. 2016). However, both species were
prevalent in Quebec, Canada, having high economic impact on potato crop (Sun
et al. 2007). In Japan, in the year 1972, PCN was earliest identified in Hokkaido
region (Yamada et al. 1972). It was believed that PCNs were introduced into Europe
during 1850 along with the planting materials introduced for breeding to late blight
resistance varieties; afterwards, their worldwide spread were recorded by the intro-
duction of improved varieties developed in Europe (Evans and Stone 1977).
Their presence in India was first noted in the Nilgiri and Kodaikanal hills of
Tamil Nadu (Fig. 1) in 1961 (Thangaraju 1983). Domestic quarantine was enforced
in India to stop PCN from spreading outside of Tamil Nadu because the seed tubers
are the main means of PCN transmission. However, PCN was afterwards recorded
from Kerala, Himachal Pradesh and Karnataka. Being a quarantine pest, somehow,

Fig. 1  Occurrence of PCN in India

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in 2010, Shimla, Himachal Pradesh (HP) witnessed their first interception from
North India (Ganguly et al. 2010). Despite the domestic quarantine against this
nematode in place, later surveys of 2011–2019 conducted in the Himalayan regions
of North India have shown the presence of PCNs in a number of districts of HP,
Jammu and Kashmir and Uttarakhand (Chandel et al. 2020). In the North Himala-
yan regions, the seed plot technique is frequently utilized to create virus-free seeds
that are sown from October to January. State agricultural departments multiply these
seeds and distribute it to farmers in other parts of the nation. Therefore, the entire
nation should be concerned about the emergence of such a significant potato seed
location as the primary sites for large PCN populations.

Life Cycle

Nematode completes one life cycle with each crop. During maturation, G. ros-
tochiensis female nematode attains yellow colour from white and later develops
a cyst of brown colour; however, female of G. pallida becomes “brown” from
“creamy white” (Wainer and Dinh 2021). In soil, PCN cysts are dormant and a stony
deceased body of female nematode protects the egg inside. Root exudates of a host
plant stimulate the eggs to hatch, leading to spread of infective juvenile second
stage (J2) nematodes in the soil. These get entered into roots by puncturing the cells
around the root tip and move to the interior cortical layers (Perry 2002; Varandas
et al. 2020). During migration of juveniles via the outer root tissues, fast and vig-
orous stylet insertion takes place which causes the cell death due to liberation of
effector proteins, cell wall modification and dissolution proteins (Cotton et al. 2014;
Palomares-Rius et al. 2017). Modification of plant cells by establishing a feeding
site, which is source of nutrients for nematode, results into poor root system and also
reduced productivity (Wainer and Dinh 2021). Afterwards, molting of nematode
takes place from third (J3) and fourth (J4) stages up to maturity. Fully developed
females become sedentary while matured males are movable in nature and leave the
root attracted towards the females, and when they produce sexual pheromones, fer-
tilization takes place (Ali et al. 2015). The female produces 300–500 eggs that she
keeps inside her body after the male fertilizes them in the soil. The female perishes
along with the root, but her skin hardens and changes to create a cyst (Fig. 2) that
protects the eggs (Stanton 1987).

Symptoms

At above ground, PCN infestation symptoms are frequently vague and difficult to
identify. In the farm, poor growth patches that exhibit yellowing, necrosis and with-
ering of leaves are common symptoms (Wainer and Dinh 2021). These species harm
potatoes by piercing and entering inside the tissues of root, which results in nutrient
and water shortage that manifests as chlorosis and withering of the leaves. They may
also cause poor growth, dwarfism and the multiplication of small lateral roots, which
reduce yield. These symptoms are not specifically produced by PCN nematodes

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Fig. 2  Life cycle of PCN

but may be caused by any other plant pathogen such as fungi, bacteria and viruses
including other nematodes. By uprooting the infected plants, young female and with
the naked eye, cysts appear as tiny white, yellow or brown nitwits on the surface of
the root. This method is possible for a short period of time as the female gets mature
into a cyst which can be simply lost with uplifting and it also takes a long time
(EPPO 2017) (Fig. 3).

Plant Growth‑Promoting Rhizobacteria (PGPR)

Various free-living bacteria present in soil, known as plant growth-promoting rhizo-

bacteria (PGPR), invade the rhizosphere and promote plant development, which
raises agricultural crop yields (Kumar et al. 2016). Azospirillum, Pseudomonas
and Bacillus are the most studied root dwelling microbes among the rhizobacterial
genera. The triumph of the PGPR has been employed for a variety of advantageous
reasons, including phytostimulation, biofertilization, bioremediation and biologi-
cal control, all of which depend on their colonization in the root system (Mhatre
et al. 2018). PGPR have also been found to be possible agents for reducing plant
parasite damage (Table 1). Nematodes and their interactions have been intensively
researched in order to effectively manage phytopathogenic nematodes (Tabatabaei
and Saeedizadeh 2017; Rashad et al. 2015). When compared to traditional manage-
ment techniques, recent research has suggested that biocontrol can be a more cost-
effective strategy to deal with PCN. Particularly, PGPR have shown promise as BCA
contenders (Siddiqui and Mahmood 1999; Wani 2015).

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Fig. 3  Skin diseases caused by Globodera pallida (A whole tuber and B second stage juveniles (J2s) in
the cuticle) (Mugniery and Phillips (2007) with permission of © (2007) Elsevier)

Non-pathogenic bacteria called PGPR have indeed been shown to increase

plant growth under both stress-free and stressful situations through both direct
and indirect approaches (Lugtenberg and Kamilova 2009). The direct and indirect
approaches define PGPR being a bio-fertilizer that generates organic chemicals
which in turns enhances soil nutrient absorption/uptake boosting plant development,
creation of antibiotics, chelates of Fe (also known as siderophores) and exterior cell
membrane disintegrating enzymes (such as chitinase and glucanase) that may hydro-
lyze the cell wall of pathogens (Van Loon et al. 1998). According to this scenario,
it has been shown that numerous PGPR, regularly belonging to the genus Agrobac-
terium, Bacillus and Pseudomonas, can lower PCN population size through several
different mechanisms, such as parasitism, the generation of hydrolytic enzymes and
antinematicidal metabolites and by inducing systemic resistance (Lugtenberg and
Kamilova 2009). PGPRs can have both direct and indirect approaches in reducing
the population of PCN.

Direct Approaches

Animosity Between PGPR and PCN

Direct antagonism of PGPR can prevent egg incubation and/or the growth and repro-
duction of PCN by predating and the release of allergens or hydrolytic enzymes.
Ammonifying bacteria release ammonia during the degradation of nitrogenous organic
compounds, which is harmful to nematodes and aids in the control of worm popula-
tions. The population of cyst nematodes was reportedly decreased by Pseudomonas
fluorescens all the way through synthesis of secondary metabolites like DAPG (Sid-
diqui and Shaukat 2003). Additionally, according to Rose et al. (2012), Pseudomonas

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 Table 1  Effects of PGPR application on nematodes
PGPR used Mode of action Observation References
Potato Research

Pseudomonas fluorescens F113 2,4-Diacetylphloroglucinol (DAPG) The improvement in hatch ability and the Cronin et al. (1997)
decrease in juvenile mobility were both driven
by DAPG
Bacillus flexus and B. pumilus The specific activity of the protease and chitinase Observed 98% egg mortality (B. pumilus) and Widianto et al. (2021)
differed significantly among the isolates 96.4% (B. flexus)
P. putida and P. aurantiacea Produced certain metabolites that can prevent Decreased nematode proliferation with P. putida Trifonova et al. (2014)
nematode eggs from hatching 3 and P. aurantiacea 13 by 40.7–42.2% com-
pared to the control, respectively
B. subtilis and P fluorescens Production of antibiotics and protease enzyme B. subtilis and P. fluorescens highest antagonis- Nikpay and khodakaramian (2013)
in case of Bacillus while in Pseudomonas tic activity and caused 78.74% and 76.54%
fluorescens production of cell wall-degrading mortality
enzymes such β1, 4-gluconase and siderophore
as biocontrol agents as well as antibiot-
ics including phenazine 1-carboxylic acid,
fenazine 1-caboxamide, pirolnithrine and
pilotheorine
Agrobacterium radiobacter and Root infection was significantly reduced by Racke and Sikora (1992)
B. sphaericus 24–41% when Agrobacterium radiobacter and
Bacillus sphaericus were present at concen-
trations of 9.7 108 and 3.16 109 cfu ­ml−1,
respectively
Agrobacterium radiobacter G12 Exopollysaccarids (EPS) and lipopolysaccha- Caused a significant decline in nematode pen- Hasky-Gunther et al. (1998)
rides (LPS) etration: 77% with B. sphaericus
Pasteuria penetrans Hyperparasitism Ciancio (2018)
B. sphaericus Induced systemic resistance (ISR) Xiang et al. (2018)
Pasteuria nishizawae Predation Sayre et al. (1991)

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fluorescens reduces the rates at which nematodes proliferate and it performed better in
conjunction with neem cake. Numerous PGPR from the families of Bacillus and Pseu-
domonas generate nematicidal substances. For instance, many strains of Pseudomonas
fluorescens reduce the levels of PCN (Globodera spp.) and root knot nematodes (Meloi-
dogyne incognita) in soil conditions and/or in vitro by producing DAPG, a phenolic
anti-phytopathogenic metabolite (Subedi et al. 2020). In comparison to mutant strains
that disrupted DAPG manufacturing, the wild-type bacteria may inhibit the activity of
adolescent PCNs by 85% (Cronin et al. 1997).

Synthesis of Enzymes (lytic)

The creation of certain enzymes is another way that PGPR works such as dehydroge-
nase chitinases, phenylalanine ammonia lyase, lipases proteases, peroxidase and so on.
Corynebacterium paurometabolous was identified to generate hydrogen sulphide and
chitinase, both of which impede the hatching of worm eggs (Mena and Pimentel 2002).
PGPR-produced hydrolytic enzymes can hinder the production and preservation of cell
membranes and walls, as well as the creation of cellular organelles (Maksimov et al.
2011). Chitinases and distinct proteolytic enzymes generated by Bacillus cereus, Bacil-
lus licheniformis, Bacillus megaterium and Bacillus subtilis, as well as Corynebacte-
rium paurometabolous, are known to contribute for hindering the proliferation of cer-
tain cyst nematodes (Subedi et al. 2020).

Induced Systemic Resistance (ISR)

The improved defense system for plants known as induced resistance is acquired with
the proper stimulation in opposition to a variety of diseases and pests. Induced systemic
resistance (ISR) or systemic acquired resistance (SAR) describes the elevated defensive
reaction induced by a substance subsequent to pathogen infection (Van Loon 2000).
Rhizobacteria have been shown in several studies to decrease the impact of nematodes
by fostering systemic resistance in plants (Ramamoorthy et al. 2001; Pieterse et al.
2002). ISR resembles SAR phenotypically; SAR is the phenomena wherein a plant
develops resistance to successive challenge inoculations by the same pathogen and dif-
ferent diseases and, in some cases, even insects (Van Loon and Bakker 2006). Treat-
ment with biocontrol agent initiates systemic resistance and ISR is induced by PGPRs
depending on jasmonic acid and ethylene as signal. Reitz et al. (2002) revealed that the
development of systemic resistance in roots of the cyst nematode Globodera pallida is
cell membrane lipopolysaccharide of Rhizobium etli strain G12. This demonstrates how
PGPR may contribute to systemic resistance against nematodes by concentrating and
regulating defense enzymes.

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Indirect Approaches

Nitrogen Fixation, Phosphorus and Potassium Solubilization

In addition to being crucial for both photosynthetically and for protein biogenesis,
nitrogen (N) is a crucial major nutrient for plants. Furthermore, it contributes sig-
nificantly to nucleotides in the form of N bases (Elrys et al. 2019, 2021). Due to
frequent N loss, agricultural soils are deficient in N. Since plants cannot utilize air
nitrogen directly therefore, in this situation, PGPR is essential for N fixation and
food replenishment. These N-fixing bacteria are divided into two categories, includ-
ing symbiotic and free-living N-fixers (Gopalakrishnan et al. 2017). As they supply
the host plant with both a source of N and an environment free of nematodes, N-fix-
ing PGPR strains having nematode-fighting activity are significantly crucial for agri-
culture on sustainable basis (Vejan et al. 2016). Utilizing PGPR containing N-fixers
as biological pest controllers for plants parasite nematodes has been confirmed par-
ticularly for sugar beet and PCN by Sikora (1992) and Nagachandrabose (2020).
Both biochemically and physiologically, phosphorus (P) and potassium (K) are
necessary for plant growth (Hasanuzzaman et al. 2018). But, the bulk of miner-
als having P and K may be found in the soil in a fixed state that the plant finds
arduous to utilize (Khanna et al. 2019). PGPR from soil such as Bacillus, Beijer-
inckia, Erwinia and so on, combine chelation, release of organic acid and acidifica-
tion to convert fixed forms of P to available ones, influencing target plant growth
and nutrient availability (Gulati et al. 2010). The microbial species Pseudomonas
syringae, Bacillus megaterium, Bacillus cereus, Pseudomonas cichorii and Bacil-
lus caryophylli are known to have the potential to both solubilize and mineralize
phosphate, resulting in improved phosphate bioavailability. Furthermore, by offering
additional aid, PGPR helps the host plant deal with nematode infestation. Accord-
ing to Seenivasan et al. (2007), Pseudomonas fluorescens and Pseudomonas lilaci-
nus reduced Globodera rostochiensis and Globodera pallida cyst levels in potato.
Some rhizospheric microorganisms convert insoluble K in an available form for
crop development and productivity, much like P-solubilizing bacteria do. For K
solubilization, PGPRs uses a variety of processes, including excreting organic acid,
chelation, reduction, complexolysis, exchange and acidolysis (Meena et al. 2016).
Additionally, the growth and nematicidal activity in tomato (Solanum lycopersicum)
were found to be benefited by the inoculation of K-solubilizing bacteria (El-Hadad
et al. 2011).

Hyperparasitism and Production of Phytohormone

Hyperparasitism involves tropic development of the biological control toward the

target cell, coiling, final attack and membrane or cell wall destruction by enzyme
activity (Mavrodi et al. 2017). Excellent efficacy has been shown by Pasteuria pen-
etrans against G. rostochiensis (Tian et al. 2007).
Many phytohormones are produced by PGPR, including auxins (IAA, IBA
and phenylacetic acid), cytokines (isopentenyl adenosine, isopentenyl adenine
riboside, transzeatin ribose and zeatin), gibberellic acid, abscisic acid, ethylene,

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polyamine, etc. which can have an influence on the metabolism and growth of plants
(Gopalakrishnan et al. 2015). It is believed that phytohormones generated by PGPR
facilitate plant growth and interactions with microorganisms (Chandra et al. 2018).
Additionally, the addition of bacteria that create IAA may promote disease resist-
ance (Chakraborty et al. 2006) and nematode biocontrol is also improved when phy-
tohormones generating PGPR are introduced to the rhizosphere (Khan et al. 2012).
Consequently, any direct impact of microbes on phytohormone creation may influ-
ence the potency of these hormones as phytostimulants. It has been demonstrated
that the class of phytohormones known as strigolactones, which Arabidopsis roots
secrete, actively contributes to host attraction during cyst nematode parasitism
(Martinez et al. 2019).

Generation of Siderophores

Living beings demand iron for a variety of biological processes, including electron
transport, respiration, photosynthesis and as an enzyme cofactor (Proenca et al.
2019). In circumstances with poor iron bioavailability, PGPR have created novel
strategies to attach iron’s insoluble form by creating siderophore of lower weight
(Schwabe et al. 2020). Enterobacter spp., Pseudomonas spp. and Bacillus spp. have
a variety of plant growth traits coupled with nematicidal behavior (El-Sayed et al.
2014). Ruanpanun et al. (2010) revealed siderophores synthesis and anti-nematode
action of Streptomyces sp. Likewise, Azadirachta, Azospirillum, Azotobacter, Bacil-
lus, Enterobacter and Pseudomonas sp. are amongst the PGPR that contribute to the
generation of siderophores, which bring iron into plant cells and encourage growth
(Cornelis 2010).

Complications Related to PGPR Application

Most traditional agrochemicals are broad-spectrum substances that have an impact

on a variety of species. Therefore, a single agrochemical can be used for a variety of
pathogens and species. However, this quality is lacking in PGPR and limiting it from
being utilized more broadly. Another issue related to use of PGPR is even if some
strains of PGPR are offered as biological nematicides in the market, there is still
a basic issue of commercialization to be resolved. PGPR products need to have a
diverse range of uses, a prolonged keeping quality, be secure for using, have a profit-
able market, be easily accessible, be reliable and having a low capital cost in order
to be financially prosperous. The partnership between multidisciplinary scientific
organizations and private enterprises, as well as its expansion to the stakeholders,
will result in the success of PGPR commercialization. Unlike other potent PGPR
species like Pseudomonas spp., Bacillus spp. are the most common species of mar-
keted PGPR because of their resoluteness. They are regarded as good choices since
they can easily create endospores, rapidly proliferate and colonize plants in adverse
situations. However, additional research is needed to evaluate their mode of opera-
tion as BCAs of PCN in order to determine more efficient procedures.

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Conclusions and Future Thrusts

Potato is staple food crop grown worldwide and is infested by plant parasitic nema-
todes, amongst cyst nematode namely Globodera rostochiensis and G. pallida which
causes the huge economic losses up to 80%, when proper management practices
are not followed. Globally, potato cyst nematode is important quarantine pest, and
to prevent the spread of PCN, Government of India kept quarantine guidelines in
Tamil Nadu and Madras. In spite of strict legislative regulations, their prevalence
was recorded in northern hills of India viz., Jammu and Kashmir, Uttarakhand and
Himachal Pradesh. It is imperative to increase production while enhancing crop pro-
tection and soil fertility with an environmentally sustainable method. Recent find-
ings have suggested the potential and sustainable benefit of PGPR as BCAs in con-
trolling PCN. This approach is widely accepted around the world as one of the safest
ways to manage nematodes and encourage plant development. By using genetic
engineering to advance PGPR methodology, consortium applications and other
advancements will guarantee the productivity and stability of agro-ecosystems,
resulting in an agricultural system that is both optimal and sustainable. Through a
direct or indirect mechanism, several strains of PGPR appear to effectively inhibit
PCN infestation and proliferation in plants and fields. PGPRs will be considerably
aided in improving plants’ ability to survive in the face of varied ecological restric-
tions without compromising yield potential if they are successfully developed and
commercialized.
The success of PGPR products’ commercialization depends on future research
into enhancing their growth circumstances, security, all-purpose action, keeping
quality and permanence. The conditions for successful use of effective strains of
PGPR for sustainable crop production include a thorough knowledge of the rhizos-
phere community and colony formation by them. In order to increase the effective-
ness of an integrated nematode management strategy, a use of both molecular and
biotechnological methods with conventional methods will help to better control of
PCN. Furthermore, by overexpressing a variety of anti-phytopathogen features syn-
ergistically for effective PCN management, scientific know-how of genetic engineer-
ing may be employed to augment the biocontrol effectiveness of PGPR.

Declarations
Competing Interests The authors declare no competing interests.

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