Current Microbiology (2024) 81:222

https://doi.org/10.1007/s00284-024-03755-0

REVIEW ARTICLE

Microbial Consortia: Promising Tool as Plant Bioinoculants

for Agricultural Sustainability
Rajeshwari Negi1 · Babita Sharma2 · Tawseefa Jan3 · Tanvir Kaur1 · Sohini Chowdhury4 · Monit Kapoor5 ·
Sangram Singh6 · Anu Kumar7 · Ashutosh Kumar Rai8 · Sarvesh Rustagi9 · Sheikh Shreaz10 · Divjot Kour2 ·
Naseer Ahmed3 · Krishan Kumar3 · Ajar Nath Yadav1

Received: 30 November 2023 / Accepted: 31 May 2024

© The Author(s), under exclusive licence to Springer Science+Business Media, LLC, part of Springer Nature 2024

Abstract
In the present scenario, growing population demands more food, resulting in the need for sustainable agriculture. Numerous
approaches are explored in response to dangers and obstacles to sustainable agriculture. A viable approach is to be exploiting
microbial consortium, which generate diverse biostimulants with growth-promoting characteristics for plants. These bioinocu-
lants play an indispensable role in optimizing nutrient uptake efficiency mitigating environmental stress. Plant productivity
is mostly determined by the microbial associations that exist at the rhizospheric region of plants. The engineered consortium
with multifunctional attributes can be effectively employed to improve crop growth efficacy. A number of approaches have
been employed to identify the efficient consortia for plant growth and enhanced crop productivity. Various plant growth-
promoting (PGP) microbes with host growth-supporting characteristics were investigated to see if they might work cohesively
and provide a cumulative effect for improved growth and crop yield. The effective microbial consortia should be assessed
using compatibility tests, pot experimentation techniques, generation time, a novel and quick plant bioassay, and sensitivity
to external stimuli (temperature, pH). The mixture of two or more microbial strains found in the root microbiome stimulates
plant growth and development. The present review deals with mechanism, formulation, inoculation process, commercializa-
tion, and applications of microbial consortia as plant bioinoculants for agricultural sustainability.

Abbreviations PGP Plant growth promotion

ACC​ 1-aminocyclopropane-1-carboxylate PGPR Plant growth-promoting rhizobacteria
K Potassium PSMs Phosphorus-solubilizing microbes
N Nitrogen Zn Zinc
P Phosphorus

6
* Ajar Nath Yadav Department of Biochemistry, Dr. Ram Manohar Lohia Avadh
ajarbiotech@gmail.com University, Faizabad, Uttar Pradesh, India
7
1 University Institute of Biotechnology, Chandigarh University,
Department of Genetics, Plant Breeding and Biotechnology,
Mohali 140413, Punjab, India
Dr. Khem Singh Gill Akal College of Agriculture, Eternal
8
University, Baru Sahib, Sirmaur 173101, Himachal Pradesh, Department of Biochemistry, College of Medicine,
India Imam Abdulrahman Bin Faisal University, Dammam,
2 Kingdom of Saudi Arabia
Department of Microbiology, Akal College of Basic
9
Science, Eternal University, Baru Sahib, Sirmaur 173101, Department of Food Technology, School of Applied and Life
Himachal Pradesh, India Sciences, Uttaranchal University, Dehradun, Uttarakhand,
3 India
Department of Food Technology, Dr. Khem Singh Gill Akal
10
College of Agriculture, Eternal University, Baru Sahib, Environment and Life Sciences Research Center, Kuwait
Sirmaur 173101, Himachal Pradesh, India Institute for Scientific Research, PO Box 24885, 13109 Safat,
4 Kuwait
Chitkara Center for Research and Development, Chitkara
University, Baddi, Himachal Pradesh, India
5
Centre of Research Impact and Outcome, Chitkara
University, Rajpura, Punjab, India

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222 Page 2 of 14 R. Negi et al.

Introduction promotes plant development through direct or indirect

methods [4]. They play a vital role for crop production and
Addressing global food insecurity is a persistent challenge protection for sustainable agriculture. They boost diversity
that is expected to exacerbate due to climate change, rapid and interaction with other beneficial microbes, enhance
population expansion, and a shortage of arable land. The soil fertility, boost crop productivity, prevent the growth
significant and anticipated growth in global population will and infectious activity of possible diseases, and overall
place substantial pressure on food security. In an attempt to preserve the sustainability of the systems. The real aspect
meet the rising demand for food, there has been an unbal- of these interactions in the environment, where a variety of
anced utilization of agrochemicals, including fertilizers microbial species can occur, is overlooked despite the fact
and pesticides for crop production. In light of the grow- that this has improved our understanding of plant–microbe
ing population and dwindling natural resources, there is a interactions [5].
pressing requirement to enhance agricultural productivity The term microbial consortium is a group of two or more
in a sustainable and environmental friendly manner. Tradi- beneficial microorganism that work together and contain
tional agricultural methods heavily rely on chemical ferti- diverse array of microbes that exhibited PGP through a
lizers to boost productivity, but this practice poses risks to variety of mechanisms and enhance activity against other
agro-ecosystems, including contamination of the food chain, pathogens [6]. Various studies have reported that inoculation
deterioration of soil quality, and water pollution [1]. Plant of microbial consortium performs better task than individ-
growth frequently takes place in various challenging envi- ual microbial strain [7]. Numerous studies have shown that
ronments, including soils with varying degrees of acidity the effect of biofertilizer containing two or more microbial
and alkalinity. The pH of the soil, controlled by hydrogen ion strains also referred to as co-inoculation or consortium is
concentration, plays a crucial role in regulating the chemical more advantageous than the application of a single microbial
properties of nutrient colloidal solutions for plants. When strain. The PGP microbial formulation used as biofertilizer
soil pH surpasses certain thresholds, it can lead to various generally uses a single microbial strain. However, incon-
stresses in plants, including hydrogen ion toxicity and imbal- sistent outcomes were observed when this biofertilizer was
ances in nutrients, resulting in toxicities or deficiencies. In applied to the soil, because it is difficult for alone strain
agriculture, one approach to address these issues has been to break down complex substances like cellulose, execute
to breed plants for stress tolerance and combine these efforts two-step or multi-step reactions, and maintain stability in
with appropriate agronomic practices to manage such chal- an unstable environment. Microbial consortia are a great
lenges [2]. It is crucial to make transition toward sustainable alternative to improve plant growth, increase crop productiv-
agricultural practices, such as the utilization of plant growth ity, soil properties and biodiversity, and reduce the chemical
promotion (PGP) microbes as an alternative to traditional inputs [8]. The mixture of two or more microbial strains
chemical fertilizers to address these challenges. This shift found in the root microbiome stimulates plant growth and
can contribute to the restoration of agro-ecosystems. development. The present review deals with mechanism,
All plants require a variety of minerals from their sur- formulation, inoculation process, commercialization, and
roundings in order for their vegetative and reproductive applications of microbial consortia as plant bioinoculants
tissues to grow and develop appropriately. These miner- for agricultural sustainability.
als have a variety of uses, such as being ionized species
that balance charges in cellular compartments, cofactors
in enzyme activities, osmotic solutes required to maintain Mechanism of Plant Growth Promotion
the right water potential, and structural elements of macro- by Microbial Consortia
molecules [3]. Based on the proportional amounts required
for plant growth, minerals can be classified into two types. Plant growth-stimulating microbes facilitate the plant
The macronutrients are sulfur (S), calcium (Ca), magne- growth by two types of mechanisms either direct and
sium (Mg), phosphorous (P), nitrogen (N), and potassium or indirect mechanisms. Direct mechanism of microbes
(K). In plants, these elements are often present in amounts stimulates the plant growth directly through production
higher than 0.1% of dry tissue weight. Iron (Fe), zinc (Zn), of various plant growth stimulators, such as auxin, cyto-
manganese (Mn), copper (Cu), boron (B), chlorine (Cl), kinin, and gibberellin; iron sequestration through the
molybdenum (Mo), and nickel (Ni) are among the cur- production of siderophores; and availing the macro- and
rently recognized micronutrients; these are often found at micronutrients through fixation, chelation, and solubiliza-
concentrations less than 0.01% of dry tissue weight. tion. The indirect mechanism facilitates the plant growth
The PGP microorganisms refers to any type of microbe, by protecting the plants from biotic stress exerted by path-
including bacteria, fungi, algae, and actinomycetes, that ogens (bacteria, fungi, virus, and oomycetes) and pest
(insects) [9]. Production of cell wall-degrading enzymes
Microbial Consortia: Promising Tool as Plant Bioinoculants for Agricultural Sustainability Page 3 of 14 222

(chitinase, amylase, protease, xylanase, and pectinase), Absorption of Nutrients from Soil
antibiotics hydrogen cyanide, siderophore, and 1-amino-
cyclopropane-1-carboxylate (ACC) deaminase activity The appropriate growth of plants requires different types of
[10, 11] (Fig. 1). macro- (nitrogen, potassium, and phosphorus) and micro-
nutrients (iron and zinc). Plants assimilate these nutrients
from soil and in soil various forms of nutrients, i.e., soluble,
and insoluble forms. In between both the forms, plants could

Fig. 1  Mechanism of microbial consortia for enhancement of crop productivity for agricultural sustainabilty
222 Page 4 of 14 R. Negi et al.

absorb only soluble form of nutrients. Most of the soil in the [21] reported N-fixing and K-solubilizing bacterial strains,
world lacks the soluble form of nutrients which is one of the namely Acinetobacter guillouiae (EU-B2RT.R1) and Acine-
main reasons for the plant growth depletion and low produc- tobacter calcoaceticus (EU- LRNA-72) inoculated in onion.
tivity. Microbes from soil and plant niches could convert The result revealed that co-inoculation of these strains posi-
the insoluble form of nutrients to soluble form via fixation tively impacted shoot length, root length, biomass, phenolic,
and solubilization mechanism. Fixation process avails the flavanoids, total soluble sugars, and chlorophyll content.
nitrogen nutrients, whereas solubilization process coverts
insoluble form to soluble form of phosphorus, potassium, Phosphorus Solubilization
and zinc [12–14].
Phosphorus (P) is one of the indispensable macronutrients
Nitrogen Fixation required for the various key metabolic processes in plants
life. Energy generation, macromolecules biosynthesis, cell
Nitrogen is essential macronutrients as it is the most impor- division, photosynthesis, signal transduction, member integ-
tant structural nutrient of the cell components (DNA, pro- rity respiration, and fixation of nitrogen are the various sig-
tein, and RNA) of plant [15]. Plant absorbs nitrogen from nificant processes in which phosphorus nutrient plays an
soil in the reduced form, i.e., ammonia, and this form of important role. Plants acquire P from the soil present in the
nutrient is in low concentration. Although, nitrogen is the top layer but the amount for plant uptake is only 0.1% of the
most abundant nutrient in the atmosphere in the form of total available amount (50 to 3000 mg ­kg−1), due to cations
dinitrogen gas which is unavailable for plant use. To fulfill precipitation, absorption, immobilization, and organic form
the plant requirement of nitrogen, urea was being utilized interconversion [22]. Being an essential nutrient farmer has
by the farmers which was prepared synthetically through totally relied on phosphatic fertilizers and its excessive uti-
Haber–Bosch process. The continue practice of urea in the lization had precipitated the soil and accumulated the heavy
fields has resulted in the serious damages to the environment metals. The precipitation and heavy metal accumulation
and the humans [16]. caused various determinantal effects on fertility soil, animal,
As a substitute, microbes with nitrogen fixation capabil- and consumers health [23]. Looking at the health hazards
ity could be used for reducing the dinitrogen into ammonia of using chemically processed fertilizers, an eco-friendly
through the process of biological nitrogen fixation. Microbes approach exigency is very important. Phosphorus-solubi-
with the attribute of nitrogen fixation have been reported lizing microbes (PSMs) have been known as an appropriate
for having specialized gene named nif which codes for a approach without any cons [24].
complex system of enzyme nitrogenase [17]. The complex Phosphorus-solubilizing microbes solubilize the P with
system of enzyme nitrogenase was elucidated into two dif- the help of solubilization mechanisms and they are converted
ferent metalloenzyme components consisting of iron protein in to soluble form with the help of several processes, namely,
dinitrogenase reductase (provides electron with high reduc- production of protons, organic acids, inorganic acid, sidero-
ing power) and a metal cofactor dinitrogenase (uses elec- phores, hydrogen sulfide (­ H2S), and extracellular enzymes
trons to reduce the dinitrogen into ammonia). On the basis [12]. Organic acid production releases the soluble form
of metal cofactor, three types of nitrogen-fixing systems of P by lowering the pH and complexation the metal ion.
were identified, namely Mo-nitrogenase (most abundantly Microbes produces various organic acids like malic acid,
found), V-nitrogenase, and Fe-nitrogenase, which are present lactic acid, lactic acid, gluconic acid, oxalic acid, tartaric
in among various genera of bacteria [18]. acid, and 2-ketogluconic acid which helps in releasing the P
In a report, nitrogen-fixing bacterial strain EU-A3SNfb [25]. The production of inorganic acids (hydrochloric acid,
Rahnella sp. was combine with P and K-solubilizer, namely carbonic acid, nitric acid, sulfuric acid, and H ­ 2S) solubi-
Bacillus tropicus EU-ARP-44 and B. megaterium EU- lizes P with low efficiency as compared to organic acid.
ARK-23, respectively. The combined effect of these strains Protons production also lower the pH of the soil aids the
significantly exhibited growth and physiological parameter dissolution of phosphorus. The production of exopolysac-
of Aegilops kotschyi and wheat crop under greenhouse and charides releases the soluble form by forming the complex
field conditions [19]. In an another report, a microbial mix- with metal ions present in soil and this extrapolated as a
ture of Erwinia sp. EU-B2SNL1 (N-fixer), Chryseobacte- mean solubilization. Microbes could also solubilize the P
rium arthrosphaerae EU-LWNA-37 (P-solubilizer), and through enzymatic actions of phytases and phosphonatases
Pseudomonas gessardii EU-MRK-19 (K-solubilizer) were and C–P lyases [14].
inoculated on barley. The result showed that the combined Numerous microorganisms from diverse habitats have
effect of these strains increased physiological parameters been reported for solubilizing the P nutrient and found to
and overall growth and was found to be more efficient as enhance the growth of the host plants upon inoculation
compared to a single inoculum [20]. A study by Kour et al. [26]. In a study Devi et al. [27] reported that phosphorus
Microbial Consortia: Promising Tool as Plant Bioinoculants for Agricultural Sustainability Page 5 of 14 222

solubilizer and nitrogen-fixing bacterial strain identified as Zinc Solubilization

Pseudomonas gessardii and Erwinia rhapontici were evalu-
ated on Amaranthus crops. The study showed that co-inoc- Zinc is the most significant micronutrient among all other
ulation of both of strain significantly enhances plant growth which plays crucial role in regulation of cofactors that acti-
and physiological parameter such as content of chlorophyll, vates wide range of enzymes. Additionally, this nutrient also
carotenoids, sugar, phenolics, and flavonoids. In a different helps in the maintaining the cellular membrane integrity,
report, three efficient bacterial strains identified as Bacillus synthesis of proteins, regulating of auxin biosynthesis, and
subtilis (P-solubilizer), Bacillus amyloliquefaciens (K-sol- pollen production. Plants uptake zinc from the soil in form
ubilizer), and Pseudomonas extremorientalis (N-fixer) were of zinc ions and the concentration of Zn ion is very less.
evaluated on pearl millet as single inoculum and as bacterial Zinc sulfate, zinc oxide and zinc carbonate are in very high
consortium. The study showed that combine inoculation of concentration which cannot be assimilated by the plants, due
consortia promote physiological parameter including content to which plants have zinc deficiency which cause problem,
of sugar, phenolics, flavonoids, chlorophyll, carotenoids, and such as yellowing of the young plants, necrosis, shortened
overall growth as compared to single inoculum [28] internode, sterility of pollens, and spikelet [35]. Zinc solubi-
lization mechanism exhibited by zinc-solubilizing microbes
Potassium Solubilization could alleviate the deficiency of the nutrient by converting
the insoluble form to soluble form. Microbes could solu-
Potassium is the third essential macronutrient for plants bilize the insoluble form of zinc either producing various
which plays role in physiological and metabolic processes, organic acids such as 2-ketogluconic acid and 5-ketoglu-
such as growth, photosynthesis, accumulation of sugar, conic acid which lowers of the pH or by metabolites that
and regulates the rate of photosynthetic carbon assimila- form a complex with zinc ion. In literature, various zinc-sol-
tion [29]. Additionally, K plays roles like root growth, high ubilizing microbes from diverse niches have been reported
yield seed development, metabolism of organic acids, fats, which also helps in the plant growth promotion [36].
carbohydrates, nitrogenase compounds, biosynthesis of pro- In another investigation, zinc-solubilizing bacteria Strep-
teins, stomata regulation, water uptake and could mediate tomyces venezuelae and Klebsiella aerogenes were found to
resistance against biotic stresses caused by various patho- enhance Zn biofortification in the wheat upon inoculation
gens [30]. Plant assimilates the K from soil in the soluble with ZnO nanoparticles and improving its growth [37]. In
form, i.e., ­K+ ion which is present in limited amount and a finding, potential zinc-solubilizing bacterial strain identi-
about 80–90% of K is exist insoluble form including biotite, fied as Bacillus sp. SH-10 and Bacillus cereus SH-17 were
illite, orthoclase, muscovite, zeolite, glauconite, chlorite, evaluated on rice as individually or as combine mixture. The
and vermiculite [31]. To fulfill the limitation of K, potash mixture of bacterial strain improve yield and Zn content as
fertilizer is widely used which have proven to be hazardous compare to chemical fertilizer [38].
to consumers and environment. To overcome this constraint,
potassium-solubilizing microbes are one of the eco-friendly
approaches which could solubilize the insoluble form and Formulation and Inoculation Process
release the ­K+ ion. Likewise, solubilization P, insoluble form of Microbial Consortia
of K is dissolute by the production of organic acids, extracel-
lular enzymes, and exopolysaccharides [32]. The primary objective when contemplating plant inoculation
In a report, three bacterial strains namely Halomonas with PGP microbes is to identify the most efficient bacterial
aquamarina (K-solubilizer), Erwinia persicina (P-solubi- strain or microbial consortia that exhibits multifarious plant
lizer), and Pseudomonas extremorientalis (N-fixer) were growth-promoting attributes and can achieve the desired
evaluated on chilli. The study showed the combined effect outcome for the specific target crop. Subsequently, the next
of these strains significantly improved plant growth param- step involves creating a tailored inoculant formulation for
eters such as root length, shoot length, fresh weight, dry the target crop, devising a practical application method,
weight, and physiological parameters, including content of and considering the limitations of the growers. The main
phenolics, flavonoids, sugar, chlorophyll, and carotenoids practical features of inoculants expected by the grower are
as compared to single inoculum [33]. In a different report, that the inoculant has to be compatible with routine field
bacterial strains identified as Bacillus sp. (K-solubilizer), practices. Secondarily, important features of microbial con-
Pseudomonas marginalis (P-solubilizer), and Stenotropho- sortia are compatibility with the seeding equipment at the
monas rhizophila (N-fixer) were inoculated on foxtail crop. time of seeding, ability to work under different field condi-
The study revealed that co-mixture of these strains exhibited tions, types of soil, tolerance of abuse during storage, ability
growth of plant as compared to single inoculums, chemical to help prolong survival of the inoculated microbe for the
fertilizer, and untreated control [34]. time needed by the plant, shelf life that lasts more than one
222 Page 6 of 14 R. Negi et al.

season, reproducible results in the field, and human, animal, applications, including seed inoculation with powder formu-
and plant safety by eliminating the use of hazardous materi- lations, mixing water and peat powder, dry fertilizers mixed
als [39]. with the seeds, soil application and seedling root dip, and
The formulation of a microbial consortium involves the suspending the biofertilizer in water along with seeds [46].
following steps: preparing the inoculums; adding additives;
choosing a carrier; sterilizing the carrier material; upscal- Soil Inoculation
ing; performing quality control processes; and packaging
the product appropriately with the best distribution chan- When microorganisms are put directly into the soil, a pro-
nels. Understanding the complexities of microbial interac- cess known as soil inoculation, they face competition from
tions in their natural environments is the biggest challenge in native microbes that have already adapted to the local envi-
designing the consortium. As a result, it will become easier ronment and outnumber the microbial inoculums [8]. It is
to design consortia when you have a solid understanding possible to significantly influence the soil’s microbiological
of metabolic pathways, compatibility of microbes and their balance and improve the environment for plant growth and
limitations [40]. The process of discovering and character- protection using inoculants of mixed cultures of beneficial
izing consortia involves a bottom-up selection approach. microorganisms [47]. Granules or liquid inoculants can be
This approach encompasses the collection of microbial added to the seedbed to inoculate the soil. The possibil-
cultures and the analysis of their properties through cul- ity that part of the inoculants will be lost during seeding
ture-dependent screening techniques [41]. Screening tests machinery and seeding is decreased when the soil is inocu-
primarily depend on specific microbial functions, such as lated. Small seeds are more benefited by soil inoculation
nitrogen fixation, antibiotic production, siderophore produc- than by seed-coat inoculation because they can be exposed
tion, phosphate solubilization, plant hormone production, to higher quantities of inoculants [39]. A microbial consor-
and ACC deaminase activity. tium solution is introduced to the soil during soil soaking as
The most promising microbial consortium is then closely as feasible to the host root. This is essential because
assessed in greenhouse conditions, and further testing is it is in the rhizosphere that the PGP microbes will be able to
carried out in the field following a bottom-up approach. It carry out various essential tasks for supporting plant growth
is worth noting that while many microbial strains may dem- promotion, such as solubilization of phosphate, potassium,
onstrate success in laboratory and greenhouse settings, they and zinc, and synthesis of siderophores and phytohormones
may encounter challenges when it comes to enhancing the [48]. In an investigation, [49] reported that by introducing
suboptimal plant microbiome in the field. Laboratory test- inoculums through soil, Bacillus improves plant growth.
ing may offer limited insights. For instance, a Pseudomonas The plant growth-promoting (PGP) microbes inoculated
strain that exhibited antagonistic activity against Phytoph- through soil improved the growth of Ranunculus asiaticus
thora infestans when developed in co-culture with another and enhanced nutrient efficiency and water absorption [50].
Pseudomonas strain, lost its biocontrol activity under field
conditions [42]. Top-down strategies enable the study of Root Inoculation
microbiome properties at a molecular level, allowing for
the selection of PGP-consortium candidates based on this The process of root inoculation involves submerging roots
molecular information. This has become possible through in a microbial solution. After being inoculated, the seedling
the direct identification of core and satellite microbiota in is set up on a growing medium that is appropriate for it.
environmental samples, relying on single amplicon variants Because inoculation may be done on seedlings of similar
obtained through high-throughput sequencing of nucleic sizes, this method allows for the standardization of plant
acids [43]. Formulations are essential to maintain the long- size. Through putting the inoculum in direct contact with
term viability of microbial cells during storage and to pro- the host roots, this inoculation approach also improves root
vide an adequate number of viable cells for field-grown colonization [16]. The various mechanisms employed by
plants. Unfortunately, there is a shortage of formulations microbes to co-operate and compete on root suggest that
available for many microbes, particularly Gram-negative microbe–microbe interactions play fundamental roles in
bacteria [44]. shaping and structuring microbial networks in environment.
There are only few different ways to distribute PGP The root of most the plant is associated with mycorrhiza.
microbes as microbial consortium in the field. Farmers are The mutualism with mycorrhizas significantly enhance the
not keen on purchasing specialized machinery to be used active surface of roots, thereby facilitating exploration of a
for microbial-based products. Therefore, prepared microbial larger soil volume for nutrients and water uptake and also
inoculation should be easily applied using common farming increase the translocation between the roots and shoots of the
equipment and simple procedures [45]. Formulated micro- host plant [51]. A study concluded that induction of induced
bial consortia as biofertilizers can be used for a variety of systemic resistance (ISR) against the fungus Colletotrichum
Microbial Consortia: Promising Tool as Plant Bioinoculants for Agricultural Sustainability Page 7 of 14 222

graminicola was reported when Pseudomonas putida was with six regional centers [60]. Worldwide, there are over 700
applied as a root inoculant in maize plants [52]. In a similar products available in the market and more than 200 biopes-
study showed that Burkholderia sp. improved Vitis vinifera ticide active ingredients are registered. In 2008, there were
resistance to low temperatures, altered its metabolism of car- just 15 biopesticides authorized for use in India under the
bohydrates, and enhanced plant growth and yield through IA (1968), accounting for a mere 4.2% of the total pesticide
root inoculation Fernandez et al. [53]. market. Nevertheless, growth at a pace of 10% is expected
to occur over the next several years [61]. The government
Seed Inoculation measures in favor of sustainable and environmentally
friendly agriculture have a significant impact on Asia. In
Co-culture inoculation treatment with seed is cost-effective India, there are about 100 public and commercial enterprises
and accessible method for field applications [54]. The inocu- that produce biofertilizer; the following is a list of a few of
lants are mixed with the seeds manually, with the use of these companies and the important products they produce.
huge dough or cement mixers, inexpensive rotating drums, The Ministry of Agriculture passed a new decree on the
or mechanical tumbling gadgets. Beneficial microorganisms control of biofertilizer production and marketing standards
can inoculate seeds to assist and prevent infections and to concerning different kinds of microorganisms. The product
colonize the roots of the seeds when they are planted in should fulfill seven quality parameters, like physical form,
soil [55]. The primary benefit of using the microbial con- minimum count of viable cells, contamination level, pH,
sortia seed inoculation technique is that it transports bac- particle size in the case of carrier-based materials, maxi-
teria directly to the rhizosphere, where they can bind with mum moisture percent by weight of carrier-based products,
plants [55]. When seeds are inoculated in leguminous plants, and efficiency character [51]. In bacterial bioproducts, the
rhizobia proliferate in the rhizosphere, where they colonize minimum viable cells to be maintained are 5 × ­107 CFU/g−1
further, form nodules, and fix nitrogen to maximize produc- for solid carrier or 1 × ­108 CFU ­mL−1 for liquid carrier. For
tivity and yield [56]. Although seed inoculants can interact products containing mycorrhizal fungi, at least 100 viable
with fungicides used for seed treatment, they also establish propagules must be present per gram of product. Nitrogen-
the plant before the pests do and can strengthen microbial fixing efficiency of biofertilizer product should be capable
defenses. In contrast, mature plants require the suppres- of fixing at least 10 mg N ­g−1 of sucrose consumed and for
sion of an established microbiome in order to develop an phosphate solubilization product a zone of solubilization of
entirely distinct one [57, 58]. There are many drawbacks to at least 5 mm in a media. AMF products should provide 80
seed inoculation. Only a little quantity of inoculant can be infection points in roots ­g−1 of inoculum [62].
coated on each seed, especially small ones. This could be a
constraining factor since most PGP microbe may require a
threshold of bacteria for successful inoculation. The sow- Applications of Microbial Consortia
ing device has the potential to remove an inoculant that is in Agriculture
not securely bonded with pelleting [39]. In a study by Kaur
et al. [20] coated barley seed with microbial consortium with PGP microbe as microbial consortium enhances agro-
sugar solution by 1:1 ratio of before sowing. In a similar nomic efficiency by reducing production costs and envi-
study, Negi et al. [19] treated Aegilops kotschyi and wheat ronmental pollution. This is achieved because effective
seed with microbial mixture and sugar solution in ratio of PGP microbes decrease the need for chemical fertilizers
1:1 before sowing. [63]. Different microbial communities, including fungi,
bacteria, actinomycetes, and yeasts, serve as inoculants,
primarily promoting plant growth through various mech-
Commercialization of Microbial Consortia anisms. These include nitrogen fixation, phosphate, and
potassium solubilization, exopolysaccharide secretion,
Commercialization of microbial consortium started in 1895 biocontrol activities, organic matter decomposition, and
when Nobbe and Hiltner introduced the rhizobia product siderophore production [63]. Microbes are employed to
under the “Nitragin” name. N. V. Joshi initiated the market- release soil nutrients for crops without harming soil fer-
ing of Rhizobium in India to promote the growth of legu- tility, ensuring an environmentally sustainable approach.
minous plants [59]. Throughout its ninth five-year plan, the Prior research has demonstrated their beneficial impacts
Ministry of Agriculture launched the National Project on on the growth and yield of diverse crops, soil types, and
Development, which aimed to popularize and promote the even under biotic and abiotic stress conditions. Moreover,
production of biofertilizers while establishing standards these microbes have also proven effective as biocontrol
for various biofertilizers, training, and exploitation. The agents against a range of plant pathogens [1]. Microbial
National Biofertilizer Development Center was established consortia have demonstrated their potential as sustainable
222 Page 8 of 14 R. Negi et al.

enhancers of plant growth and as aids in coping with vari- Limitations and Challenges
ous environmental stresses. Given the extensive, long-term
evolutionary interactions between plants and microbes, it An essential component of any effective co-cultivation
is highly likely that there are still many undiscovered ben- system is strain compatibility [73]. In addition to being
efits that can be harnessed from PGP microbes [2]. able to grow effectively in the same growing conditions
PGP microbe are free-living bacteria that play a direct media, pH, temperature, and oxygen requirement. The co-
role in promoting plant growth and enhancing root systems cultivation constituent stains must also be able to avoid
in plants [64]. Various pieces of literature have revealed producing harmful substances that might seriously impair
that these bacteria enhance plant growth and crop yields the other microbial community members [74]. Microbial
through their plant growth-promoting activities. The lit- strains from the same species can be used to meet these
erature shows that a microbial consortium has a positive requirements because their growth rates and requirements
impact on PGP activities. Numerous reports indicate that for growing environments are identical [75]. However
a wide array of microorganisms thrive in their challenging when several strains from various species are employed
environments, engaging in interactions with other microor- to build the synthetic microbial consortium, issues occurs
ganisms, both intra- and interspecifically. In natural envi- since different species have varied requirements for media
ronments, the majority, around 99%, of microorganisms and have considerably different growth rates. Success-
exist in the form of microbial consortia [65]. Numerous ful synthetic microbial consortia not only carry out the
studies have demonstrated that individual microorgan- desired functions but also sustain cell growth in a stable
isms can have a positive impact on plant growth. How- and robust way. More stable relationships among consor-
ever, in natural settings, it is evident that multiple species tium members are formed when they highly depend on
within microbial consortia can perform a broader range each other. Microbial interactions that lead to the inter-
of beneficial functions for ecosystems compared to single dependence and stable relationships include cross-feeding,
microorganisms. The interaction between PGP microbes detoxification, and biofilm formation, which are important
and plants synergistically contributes to the overall ben- consortium design principles [76].
efits for the plant microbiome [66]. Plants further support It is not intended for the industrial fermentation pro-
the growth of PGP microbe by producing various storage cess used in bioproduction for co-cultivation partners to
substances and root exudates, which serve as sources of use the same growth resources as this will lead to unsta-
nutrition for these beneficial microbes [67].Microbial con- ble co-cultivation and competitive exclusion. Nutritional
sortia are abundant in various natural settings, such as bio- divergence or syntrophy techniques have been used in
films, food products, soils, and wastewater. These consor- co-cultivation systems to overcome this problem. These
tia are prevalent in soil and exhibit superior performance techniques provide effective energy and carbon chan-
compared to individual microorganisms in accomplishing neling, which contribute to the formation of dynamic and
multiple functions. When employed as inoculants, they symbiotic microbial interactions within the consortium
demonstrate the capacity to thrive in dynamic environ- [77]. It is challenging to use this strategy, though, because
ments, as they can occupy a wider resource niche within every organism has different nutritional needs and prefer-
the soil when working together rather than individually. ences. Consequently, it is preferable to have cross-feeding
This enables them to compete more effectively with the or nutritional divergence within a co-cultivation since it
native soil microorganisms [68]. Furthermore, considering enables the removal or reduction of a microbial species
the intricate interactions between soil microbiomes and from the consortium and permits coexistence [78].
plants, utilizing microbial consortia appears to be a more The most significant obstacle to bioproduction in micro-
practical strategy compared to using single microorgan- bial consortia is maintaining the population ratio at the
isms as inoculants [69]. desired level during co-cultivation. The population com-
In an investigation, microbial mixture of Bacillus megate- position of co-cultivations can vary significantly because
rium, Arthrobacter chlorophenolicus, Enterobacter sp., and of a number of reasons, including substrate competition,
P. aeruginosa increased grain yield by 75.80% and 40.09% variations in doubling times, and hazardous by-products
under greenhouse and natural conditions, respectively [70]. generated by consortium members. As the culture vol-
In an another investigation, microbial mixture of Bacillus ume increases, the stability of the culture population ratio
cereus, Lysobacter antibioticus, and Lysobacter capsici declines, which could result in system heterogeneity [79].
increase the yield by 2909.8 kg/666.67 ­m2 [71]. In a finding, To prevent one strain from eradicating the other, there are
microbial consortium of Enterobacter spp. ZW32, Ochro- strategies to maintain the strain-to-strain ratios among
bactrum sp. SSR, and Enterobacter spp. was inoculated on the co-cultivation members. Although it is frequently
wheat crop. The study revealed that mixture of microbial seen that the sub-population ratio varies or fluctuates over
strain significantly enhance 15% grain yield [72] (Table 1).
Table 1  Effect of microbial consortium on different crops
S.N Microbial Consortium Crop Inoculated Effect References

1 Bacillus megaterium + Ensifer adhaerens + Pseudomonas fluores- Wheat Decreasing electrolyte leakage and increasing chlorophyll contents, Khan et al. [81]
cens relative water contents, and positive impact on shoot length, root
length, shoot fresh weight, and root fresh weight
2 Bacillus sp. + Azospirillum brasilense + Azosprillum lipoferum Wheat Decreases electrolyte leakage and enhanced the chlorophyll content, Akhtar et al. [82]
RWC, proline content, amino acid, and antioxidant enzymes
3 Pseudomonas aeruginosa + Trichoderma harzianum + Bacillus Pea Increase in plant length, total biomass, number of leaves, nodules Jain et al. [83]
subtilis and secondary roots, total chlorophyll, and carotenoid content
4 Pseudomonas extremorientalis + Bacillus subtilis + Bacillus Pearl millet Increase length and biomass of root/shoot, chlorophyll, carotenoids, Kaur et al. [28]
amyloliquefaciens total soluble sugar content, phenolics, and flavonoids
5 Siccibacter colletis + Enterobacter huaxiensis + Pantoea sp. Faba bean Increase root and shoot dried biomasses and chlorophyll content, P Chamkhi et al. [84]
uptake
6 Rahnella sp. + Bacillus tropicus + B. megaterium Wheat Enhance growth as well as physiological parameter including shoot/ Negi et al. [19]
root length, fresh/dry weight and chlorophyll, carotenoids, total
soluble sugar content, phenolic and flavonoids content
7 Erwinia sp. + Chryseobacterium arthrosphaerae + Pseudomonas Barley Increase root/shoot length and biomass, chlorophyll, carotenoids, Kaur et al. [20]
gessardii phenolics, flavonoids, and soluble sugar content
8 Erwinia persicina + Halomonas aquamarina + Pseudomonas Chilli Increased shoot/root biomass and length; number of leaves, Devi et al. [33]
extremorientalis branches, and fruits per plant and content of chlorophyll, total
soluble sugar, phenolics, and flavonoids
9 Acinetobacter guillouiae + Acinetobacter calcoaceticus Onion Positively impacted shoot length, root length, biomass, phenolic, Kour et al. [21]
flavanoids, total soluble sugars, and chlorophyll content
10 Stenotrophomonas rhizophila + Pseudomonas marginalis + Bacil- Foxtail millet Improved the growth and physiological parameters Kaur et al. [34]
Microbial Consortia: Promising Tool as Plant Bioinoculants for Agricultural Sustainability

lus sp.
11 Azotobacter sp. + Pseudomonas sp. Malt Barley Highest and significant effect on grain yield, harvest index, biologi- Tadesse, Melak [85]
cal yield, plant height, and thousand seed weight
12 Pseudomonas gessardii + Erwinia rhapontici Rajgira Enhance shoot/root length and biomass and chlorophyll content, Devi et al. [27]
total sugar content, phenolics, and flavonoids content
13 Bacillus thuringiensis + Bacillus horikoshii + Pseudomonas trivialis Sweet pepper Increase plant length, root length, fresh weight, and biomass of the Devi et al. [86]
plant and chlorophyll, carotenoids, flavonoids, phenolics, and total
soluble sugar content
14 Pseudomonas aeruginosa + Trichoderma harzianum + Bacillus Pea Increase in plant length, total biomass, number of leaves, nodules Jain et al. [83]
subtilis and secondary roots, total chlorophyll and carotenoid content, and
yield
15 Azospirillum + Azotobacter chroococcum + Bacillus megaterium Ashwagandha Significantly increased plant height, root length, and alkaloid Rajasekar, Elango [87]
+ Pseudomonas fluorescens content
16 Azotobacter chroococcum + Priestia megaterium + Pseudomonas sp. Pigeon pea Significant increase in plant attributes such as shoot length, root Srivastava, Sharma [88]
length, fresh weight, and dry weight
17 Bacillus licheniformis + Bacillus sp. + Pseudomonas aerugi- Sunflower Increased the plant growth and yield and reduced the SNV disease Srinivasan, Mathivanan [89]
Page 9 of 14

nosa + Streptomyces fradiae incidence

222
 222 Page 10 of 14 R. Negi et al.

the cultivation period, fine-tuning the inoculation ratio

Molina-Romero et al. [94]

between co-cultivation partners has a significant impact
on overall productivity. Additionally, research on mutu-

Pellegrini et al. [91]

alistic growth has been done to keep the engineered co-
Kapadia et al. [90]

Kumar et al. [70]

Kumar et al. [92]

Dary et al. [93]

cultivations’ intended composition of the population [80].
References

Conclusions

Significant increase in the length of the aerial part of the plant, root
Increased plant height, total chlorophylls, crop yields, and bulb dry

Increased grain yield, maximum increments in vigor index, nodule

Increases plant yield and nitrogen, and decreases plant metal accu-

length, total length, stem diameter, circumference, dry weight of

Increase in leaf, shoot, root dry weight, leaf number, shoot length,

Showed significant increase in plant height, grain yield, and straw

A new approach to alteration in metabolic pathway balanc-

ing is provided by microbial biosynthesis via co-cultivation
engineering. It increases the range of options for fine-tuning
root length, secondary roots, and chlorophyll content

the aerial part of the plant, and the biovolume index intricate metabolic pathways and can be tailored for effective
synthesis of various bioproducts. Compared to mono-culti-
vation systems, co-cultivation engineering offers a number
of benefits, including increased productivity, robustness,
number, and root and shoot biomass

modularity, and tolerance (toxic intermediates and waste

created by one partner are consumed or degraded by the
other partner). Co-cultivation fermentations have the poten-
tial to improve production efficiency and enable the use of
less expensive substrates. Furthermore, without sacrific-
ing output or quality, artificial consortiums provide a solu-
tion to solve the problems associated with the functional
expression of complicated biosynthetic pathway enzymes.
mulation

They may also lessen the work involved in reconstructing

matter

yield
Crop Inoculated Effect

recombinant biosynthesis pathways. Co-cultivation of sev-

eral populations is more difficult since it is unknown how
individual strains would behave when grown together using
standard techniques, and it may also be harder to regulate as
Fenugreek

Lupines

the number of constituent strains or species increases. But

Tomato

Wheat

Wheat
Onion

recent advances in co-cultivation engineering have signifi-

cantly increased our comprehension of how microbes inter-
act in communities. Synthetic biology techniques must be
Bacillus megaterium + Arthrobacter chlorophenolicus + Enterobac-
Bacillus sp. + Delftia sp. + Enterobacter sp. + Achromobacter sp.

Bacillus paralicheniformis + Bacillus subtilis + Bacillus megate-

used to overcome certain inherent obstacles, even though

Bradyrhizobium sp. + Pseudomonas sp. + Ochrobactrum cytisi
cus + Herbaspirillum seropedicae + Burkholderia ambifaria

the potential of microbial consortia seems extremely prom-

Azospirillum brasilense + Gluconacetobacter diazotrophi-

ising. In the near future, it is expected that co-cultivations,

or polycultures made up of several specialized members
will be created and used to address the demand for increas-
ingly complex biochemical pathways. The creation of novel
Ensifer meliloti + Rhizobium leguminosarum

microbial consortia with distinct roles should be explored

through the use of metabolic engineering and synthetic biol-
ter sp. + Pseudomonas aeruginosa

ogy techniques. Since functional genomics (metabolomics,

proteomics, and transcriptomics) is still expensive, new
rium + Bacillus cabrialesii

methods must be developed to investigate community pro-

files and interactions among microbes in consortium culture.
S.N Microbial Consortium
Table 1  (continued)

Author Contribution R.N. contributed to Writing of the original draft,

B.S. contributed to Writing of the manuscript, T.J. contributed to
Writing of the manuscript, T.K. contributed to Writing of the man-
uscript, S.C., M.K., S.S., A.K., A.K.R., S.R., S.S., D.K., N.A., and
K.K. contributed to Reviewing of the manuscript, A.N.Y. contributed
to Conceptualization.
18

19

20

21

22

23
Microbial Consortia: Promising Tool as Plant Bioinoculants for Agricultural Sustainability Page 11 of 14 222

Declarations mays L.). J Appl Biol Biotechnol 9:41–50. https://​doi.​org/​10.​

7324/​JABB.​2021.​9306
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