Biologia

https://doi.org/10.1007/s11756-024-01848-6

REVIEW

Microbial biofertilizers: A paradigm shift towards agricultural

sustainability
Rajeshwari Negi1 · Neelam Yadav2 · Ajar Nath Yadav1,3

Received: 17 September 2024 / Accepted: 25 November 2024

© Plant Science and Biodiversity Centre, Slovak Academy of Sciences (SAS), Institute of Zoology, Slovak Academy of Sciences (SAS), Institute of Molecular
Biology, Slovak Academy of Sciences (SAS) 2025

Abstract
The world has changed exponentially from the time of famine to the existent, when food is produced globally to feed a
population that is expanding at an exponential rate. The intensification of agriculture through the introduction of mechanical,
industrial, and economic inputs has been a hallmark of major agricultural revolutions, which have enabled this transforma-
tion. However, the explosion of agricultural inputs consisting of fertilizers, pesticides, and irrigation brought about by this
quick development in agriculture has also resulted in long-term environmental crises. These challenges have highlighted
the pressing need to safeguard our communal spaces, especially the environment, by means of a participatory strategy that
engages nations everywhere, irrespective of their level of development. One notable effort in this area is Zero Budget Natu-
ral Farming (ZBNF), which emphasizes the value of utilizing the complementary effects of plant and animal products to
improve soil fertility, encourage the growth of beneficial microbes, and improve development of crops. The development of
self-sustaining agro-ecosystems is the ultimate goal. Consequently, the world is depending on the use of microbial formu-
lations in agriculture to address the "5F" crisis; food, feed, fuel, fertilizer, and finance. Despite the fact that there are now
many studies being conducted in this area, the market need for effective microbial formulations outweigh the supply. Many
different microbes have been considered so far for their potential as plant stimulants, but there are still innumerable soil
microorganisms that need to be found in order to play a useful role in the formulation companies. The present review deals
with improving the broad range of mechanism of microbial formulations, delivery methods, challenges and biosafety issue
and assessment for agricultural sustainability that support the sustainable development goals (SDGs).

Keywords Agriculture · Biofertilizers · Formulations · Green Revolution · Microorganisms · Sustainability · Zero hunger

Abbreviations K Potassium
PGP Plant growth promoting Zn Zinc
PVP Polyvinyl pyrrolidone ZSB Zinc solubilizing biofertilizers
N2 Nitrogen Se Selenium
IAA Indole-3-acetic acid Fe Iron
P Phosphorus AMF Arbuscular mycorrhizal fungi
PSB Phosphorus solubilizing bacteria BSL Biosafety level
EHSI Environmental and human safety index

* Ajar Nath Yadav

ajarbiotech@gmail.com Introduction
1
Department of Genetics, Plant Breeding and Biotechnology,
Dr. Khem Singh Gill Akal College of Agriculture, Eternal India accounts the second-largest agricultural area in the
University, Baru Sahib, Sirmaur, Himachal Pradesh, India world with 157.35 million hectares under cultivation over
2
Centre of Research Impact and Outcome, Chitkara University 20 agro-climatic zones (Eliazer Nelson et al. 2019). Thus,
Institute of Engineering and Technology, Chitkara University, agriculture plays a significant role since, 58% of rural house-
Rajpura, Punjab, India holds depend on the agriculture sector. During the year of
3
Chitkara Centre for Research and Development, Chitkara 1947 to 1960 India had such low food production and there
University, Baddi, Himachal Pradesh, India

Vol.:(0123456789)
 Biologia

was a chance famine. Thus, the green revolution was started as nitrogen fixers, nutrient solubilizer, biocontrol agents, and
in the 1960s in order to boost food production to feed mil- bioremediation (Singh and Kumar 2023). Preparing micro-
lions and reduce extreme poverty and malnourishment in bial formulations has several main objectives: to establish
the nation (Gómez et al. 2013). In 1968, William S. Gaud a suitable environment for the bioinoculants to function; to
of the United States Agency for International Development offer long-term physical and chemical protection to prevent a
(USAID) coined the term "Green Revolution" to describe sharp decline in cell viability during storage; to encourage the
the introduction of new scientific approaches and policies inoculants' competition with the native soil microbiota; and to
that were carried out in developing countries between the lessen losses caused by local micro fauna depredation (Kes-
1940s and 1960s with assistance from industrialized nations wani et al. 2016). However, another objective is to supply a
to increase crop productivity and the yield. The term "Green sufficient amount of living bioinoculant cells that are available
Revolution" refers to a series of creative scientific research for interaction with plants and the soil microbiota. It has been
and technology transfer projects that took place between noted that while direct application of plant-beneficial micro-
1950 and the late 1960s (Parayil 2003). These projects, organisms on a small or greenhouse scale is acceptable, the
which started to gain attention in the late 1960s, improved microorganism’s viability problems worsen on an extensive
agricultural production that was sustainable everywhere, or field scale. Indeed, in order to get a favorable response from
especially in developing nations. Agrochemicals and chemi- the prepared product, an adequate amount of microbial cells
cal fertilizers were used in conjunction with high-yielding (at least ­106–107) must be acquired (Chojnacka et al. 2020).
cereal types as part of the fundamentally new strategy that In recent years, formulation technologies have focused
was adopted (Negi et al. 2023). more on cell immobilization, since gel-cell immobilization
India's production of food grains is now self-sufficient is a technological solution that can better ensure the result-
due to the Green Revolution, but the environment health ing product's quality and consistency (Khan et al. 2023b).
and our food chain have been harmed by the excessive and A necessary step is the delivery of bioformulations that can
inappropriate use of agrochemicals (Srivastava et al. 2016). be accomplished by treating plants or seeds or by directly
The Green Revolution expanded dissemination affects both inoculating the soil. Demand for biofertilizers has surged
agricultural biodiversity as well as wild biodiversity (Dhi- due to growing concerns about the insufficient uptake of
man et al. 2024). The majority of the developing world's chemical fertilizers by plants and their negative effects on
population depends on agriculture for a living, and ferti- ecosystems, as well as a global increase in concern about
lizers and pesticides are employed extensively in this sec- pollution, greenhouse gas accumulation, and a greater focus
tor to increase output by shielding food grain crops from on plant-based food production (Sahu and Brahmaprakash
any dangers (Sharma et al. 2024b). The majority of plants 2016). Biofertilizers are becoming more and more popular
absorb 10% to 40% of the applied fertilizer, with the remain- among farmers that want to grow their crops organically
ing 60% to 90% being unused (Chojnacka et al. 2020). To and appropriately. Many biofertilizers have been effectively
maintain agricultural production and a healthy environment, marketed to date for a range of crops and ecological condi-
microbial inoculants are crucial components of integrated tions (Badiyal et al. 2024). However, ignorance of how to
nutrient management systems. It is now widely acknowl- choose and administer biofertilizers effectively is a major
edged that we must take action to protect the environment barrier to their widespread adoption in agroecosystem (Rai
and nourish the earth through the use of environmentally et al. 2023). The lack of understanding undermines farmers
friendly, sustainable organic farming methods (Darnhofer reliance on biofertilizers. The scientifically sound techniques
et al. 2010). A significant step towards sustainability will be for choosing and implementing the appropriate microbial
to engage in organic agriculture and follow in the footsteps bioformulations based on their native environment and crops
of our ancestors. must thus be widely propagated throughout farming commu-
Microbial bioformulations provide an alternate method nities. The present review deals to explore the broad range
of employing advantageous soil and plant microorganisms of mechanism of microbial formulation, delivery methods,
to promote healthy plant development and yield (Kour et al. challenges and biosafty issue and risk assessment that sup-
2024b). The utilization of bio-formulated goods, particularly port sustainable agriculture development (Fig. 1).
biofertilizers, has gained widespread acceptance as a substi-
tute for agrochemicals (Rachappanavar et al. 2024).
Therefore, the term bio-formulation can be represented as Characterization of novel plant growth
the development of material that contains living microbial promoting microbiome
strains and uses appropriate carrier materials for their pro-
ductive usage in agriculture (Negi et al. 2022a). The primary The sheer number of microorganisms found in plants,
components of bioformulation are prospective microorgan- both below and above ground, has produced mutualis-
isms with plant growth promoting (PGP) characteristics, such tic interactions that benefit plant health. Three groups
Biologia

Fig. 1  Microbial bioformulation strategies for development of novel biofertilizers for agricultural sustainability

can be distinguished from the microbes responsible for Endophytes are found inside plant tissues comprising
plant invasions: microbes found in the soil around roots seeds, leaves, and stems. Surface sterilization is most widely
(Rhizospheric microbes), inside plant tissues (Endo- used technique for isolation of endophytes. Plant material
phytic microbes) and aerial parts of plant (Phyllospheric must be cleaned and treated with 70% ethanol, 2.5% sodium
microbes). Enrichment techniques that employ traditional hypochlorite, and sterile water in order to undergo surface
serial dilution and pour plating method can be used to iso- sterilization. On the appropriate growing medium, bacteria
late the culturable microorganisms present in rhizospheric and endophytic fungi can be isolated (Abdalla and McGaw
soil. Employing the serial dilution method, one gram of 2018). Numerous rich and varied communities of bacteria,
soil sample of soil is need to be dissolved in sterile water fungi, and viruses can be found flourishing on the phyl-
until an overall 10 mL suspension. Then, 1 mL of the losphere, or aerial surface of plants. The leaf imprinting
10 mL suspension need to be transferred to another test method can be employed to isolate the epiphytic microbes
tube that has 9 mL of sterile water to obtain ­10–2 serially (Holland et al. 2000). Pressing the leaves onto the agar
diluted suspension. Similar suspensions can be used for plates. The leaf should be removed from the plates after
­10–3, ­10–4 and onward dilutions. The 100 µL suspension imprinting for 30 to 45 min and incubated in the BOD incu-
need to spread onto agar plates having growth medium and bator at 5 to 30 °C for 6–7 days.
inoculated plates needed to be incubate for 2–7 days. The Bergey's Manual of Determinative Bacteriology could be
pure cultures need to preserved for future uses (Verma used to identify and characterize the microbes on the basis of
et al. 2013). their physical and biochemical traits (Buchanan and Gibbons
 Biologia

1974). Identification based on molecular biology is thought Lesueur 2013). Biofertilizer formulations can be produced
to be faster and more accurate than identification based on in liquid, granular, or dry powder form by utilizing vari-
both physical and biological characteristics. For molecular ous carriers such as peat, clay, or liquid media that promote
identification, (pA) and (pH) primer may be used for 16S the growth of associated microbes and facilitate effective
rRNA gene (bacterial) and ITS for18S rRNA gene (fungal) distribution (Fig. 2). Among them, a unique cell protectant
amplification under standard conditions (Sonowal et al. including sucrose, arabic gum, glycerol, and polyvinylpyr-
2024). The amplified PCR product should be sequenced rolidone found in liquid biofertilizers can promote the devel-
using an automated Sanger's di-deoxy nucleotide sequenc- opment of dormant spores, extending the fertilizers shelf life
ing technique. The identification of microbes should be done (Saif et al. 2021). In comparison to other forms of biofer-
using BLASTn techniques, and sequence submitted to NCBI tilizers, liquid biofertilizers are advised to be used at lower
GenBank and accession number assigned to the strain (Mir- dosages. Choosing an appropriate carrier material for the
sam et al. 2022). The ClustalW program should be used to inoculums is also crucial because every carrier has pros and
further align the sequences and MEGA 4.0 software could cons that impact the final product's quality. A carrier serves
be used to create a phylogenetic tree using the neighbor- as a vehicle to deliver the microorganisms from the produc-
joining method and evolutionary distance data (Tamura et al. tion process to the field (Nagpal et al. 2021). Because it
2007). offers the microorganisms a temporary protective niche both
in the soil and in the package, the carrier (weight or volume)
represents a significant percentage of the biofertilizer and is
Preparation of bioformulations: steps crucial to the delivery of viable cells. The appropriate car-
and standards riers should be: nontoxic; highly capable of absorbing mois-
ture; highly capable of holding water; long lasting shelf life;
In order to meet biofertilizer requirements, the production of and lump-free and easy to handle (Sylia et al. 2021; Anand
biofertilizers is a multifaceted procedure that goes through et al. 2022b; Yadav et al. 2023; Khan et al. 2023a). The
several stages of evaluation and quality control. The surviv- carrier encapsulation of the inoculants ensures long term
ability of microorganisms is one of the main challenges. The storage, convenient handling, and the biofertilizers efficacy
formulation of biofertilizers must allow microbes to survive (Sahu and Brahmaprakash 2016). Make sure the carrier
and persist to enhance soil fertility even after a considerable has been fully sterilized using either autoclaving or gamma
amount of time has passed since packaging (Herrmann and radiation before adding inoculants to it. When developing

Fig. 2  Microbial biofertilizer types, inoculation methods and roles of different biofertilzer for agricultural sustainability
Biologia

biofertilizers, an adjuvant is also included in addition to car- Wettable powders It's among the more traditional forms of
riers (Sahu and Brahmaprakash 2016). The microbial cells bioformulation. According to the international units, wet-
are preserved and adhered to by this adjuvant. table powder should have the following composition by
weight: 50% powder, 15%–45% filler, 3%–5% surfactant,
and 1%–10% dispersant (Brar et al. 2006). These formula-
Types of bioformulations tions are developed with a range of raw materials, compris-
ing sand and wheat bran mixtures, sawdust and molasses,
Solid bioformulations molasses manufactured from corn cobs and sand, organic
cakes, sand and cow dung, compost, fly ash, and charcoal.
Dried versions of solid formulations are available for pur- These formulations can be introduced to the liquid carrier
chase. Compared to other formulation, they have a longer right before application, or they can be extremely miscible
shelf life and are convenient to transport and store (Patil with water (Hazra et al. 2017). Comparatively speaking
et al. 2013). Granules that are wettable or water-dispersible to other varieties, this bioformulation has a reasonably
and dust are all components of the solid formulation. Disper- long shelf life because of its nearly nonexistent moisture
sants, wetting agents, and binders are combined to develop- content.
ing them (Balasubramanian and Karthickumar 2017).

Dried powder Dry powdered bioformulation is made from Liquid bioformulations

soil, organic, inorganic, or inert components (Mishra and
Arora 2016). These formulations are time-tested and com- The liquid bioformulations have been produced with min-
prise an extremely finely ground combination of active eral oil, organic oil, broth cultures, oil in water, or suspen-
ingredients. Merely 10% of the particles size falls between sions based on polymers. It contains 10 – 40% microbes,
50 and 100 μm. The uppermost natural layer of the soil, 1 – 3% suspender, 35 – 65% carrier liquid, 1 – 5% dis-
called peat, is typically used to make the dried bioformula- persant, and 3 – 8% surfactant. It's simple to apply liquid
tion (Khan et al. 2023b). When organic stuff falls to the biofertilizers to both soil and seeds. Compared to the other
ground over time, it breaks down naturally and forms peat. two categories of bioformulation, there is a greater con-
Microorganisms and plants both receive nourishment from centration of cells (Bharti et al. 2017). Because farmers
peat. Peat is used as a carrier in most formulations because may employ it in fewer quantities, it is less exclusive while
of its high natural content and water retention property. It maintaining the same level of effectiveness (Negi et al.
is inexpensive, easily sterilizable, and very absorbent. The 2024a, 2024c). In addition to having a longer shelf life
removal of hazardous elements from peat is necessary dur- of one to two years, liquid formulations are less likely to
ing sterilization to prevent microbial development and main- become contaminated, produce large quantities and work
tain the quality of the bioformulation (Nagpal et al. 2021). well in the field, don't require sticky materials, are sim-
ple to use and manage with contemporary equipment, can
Granules The granular bioformulation comprises dry parti- tolerate high temperatures, promote microbial growth, are
cles with active ingredients (5 – 20%), binder, and granular simple to apply to seeds and soil, and offer more protec-
carrier. Granules are coarse particles (size 100 – 1000 µm), tion against environmental conditions (Yadav 2023). Air
non-dusty, and without risk of inhalation (Ijaz et al. 2019). seeders and other farming equipment are compatible with
The granules consist of wheat and corn meal granules, glu- liquid formulations. Liquid bioformulation contains arabic
ten, cornmeal baits, cottonseed flour, gelatin, peat prill, pesta gum, sucrose, glycerol, trehalose, polyvinyl pyrrolidone
granules, tiny pebbles, and sodium alginate (Sharma et al. (PVP), polyethylene glycol as cell protectant, that help in
2023). After applying an adhesive, they are combined with improving the survival of microorganisms, and maintain-
inoculums that have been powdered. These compositions ing high number (John et al. 2011). Suspension concen-
may be handled more easily and produce less dust. They trations can be produced from liquid formulations. The
can be separated into two main categories based on granule production of these formulations involves the combination
size including coarse particles size about 100–1000 μm and of solid active ingredients, most of which have low water
micro-granules (size 100–600 μm) (Mishra and Arora 2016). solubility. Before using these formulations, they must be
The active ingredients in these formulations dissolve imme- diluted with water or another carrier. Improved solubil-
diately into the soil, allowing them to be non-caking, free- ity and storage are achieved by adding various surfactants
flowing. Granular bioformulations are successful, but have and additives (Mishra and Arora 2016). Over the couple
major constraints, such as the deactivation of active ingre- of decades, they have become considerably more crucial
dients in the presence of UV radiation (Xavier et al. 2004). to farmers.
 Biologia

Encapsulated bioformulations of plant as compared to un-inoculated control (Pham et al.

2017). A study concluded that, three efficient N­ 2-fixing bac-
To sustain the survival of the cells within the beads, the teria identified as Sphingomonas trueperi, Psychrobacillus
microbial cells in this formulation are coated, trapped, or psychrodurans and Enterobacter oryzae were evaluated on
enclosed in a polymeric material (Saberi Riseh et al. 2021). maize and wheat plants. The result showed that, inoculation
This causes gases, nutrients, and metabolites to be released of these strains improves surface area, growth parameters,
into the beads so they can pass through. The macro-encap- and nutrient content in both of the crop (Xu et al. 2018). A
sulation involves beads of millimeters to centimeters in study documented that, application of N ­ 2-fixing bacterium
size, while micro-encapsulation is 1–1000 µm in size (de enhance maize growth (Ke et al. 2019). According to a study
Vos et al. 2009). A variety of polymers, including gelatin, by, Singh et al. (2020b), two N ­ 2-fixing bacteria identified
cellulose, starch, and others, are frequently used to encase as Bacillus megaterium and B. mycoides improves N con-
bioactive molecules. The natures of the active components tent, and growth of sugarcane crop. A study revealed that,
are determined prior to encapsulation, and stabilization is efficient ­N2-fixing bacterium identified as Bradyrhizobium
accomplished by physio-chemical (gelation or coacerva- yuanmingense enhance nodule and plant biomass of Desmo-
tion), physical (solidification or evaporation), or chemical dium incanum (Toniutti et al. 2021). A study by (Negi et al.
(polymerization) processes (Ayyaril et al. 2023). To provide 2022a) reported that, application of efficient ­N2-fixing bac-
additional protection, some dyes can be applied to the active terium identified as Rahnella sp. improves growth and physi-
ingredients. Research has shown that bioformulation made ological parameters of Aegilops kotschyi. A study concluded
by encapsulation typically have longer shelf lives because that an endophytic ­N2-fixing bacterium was isolated and
the covering layer shields the bioactive ingredient inside identified as Rahnella aquatilis from a cereal crop, and the
from external stressors (Shashirekha et al. 2024). inoculation of this bacterial strain enhances wheat growth
(Triticum aestivum L.) (Rana et al. 2024).

Microbial biofertilizer with diverse plant Symbiotic nitrogen‑fixing biofertilizers

growth promoting attributes
Plants and ­N2-fixing microbes can work synergistically to
Nitrogen‑fixing biofertilizers fix nitrogen in biofertilizers. These biofertilizers take advan-
tage of α-proteobacteria, a family of microorganisms that
The atmosphere is approximately 78 percent nitrogen, yet infect and bond with the roots of leguminous plants (Yadav
plants are unable to absorb or use nitrogen in its molecular et al. 2021). The primary components of this substance
form. Nitrogen (­ N2) in this form is not utilized by plants for consist of Bradyrhizobium, Mesorhizobium, Azorhizobium,
their metabolic processes (Yadav 2021). The only way that Allorhizobium, Rhizobium, and Sinorhizobium. Rhizobium
plants can absorb nitrogen is if N ­ 2-fixing microbes trans- is a bacterium that coexists with its host plant in symbiotic
form atmospheric nitrogen into forms that plants can use, relationships and resides in the root nodules of legumes. Iron
such as ammonia, ammonium ions, or nitrate (Mahanty et al. and molybdenum serve as cofactors for nitrogenase, which
2017). An enzyme known as dinitrogenase helps transform is the enzyme that Rhizobium uses to convert atmospheric
gaseous nitrogen into ammonia during biological nitrogen nitrogen to ammonia (Mahanty et al. 2017). Frankia also
fixation. Nitrogen availability is improved by applying these fixes nitrogen symbiotically, usually forming a relationship
­N2-fixing microbes (Negi et al. 2024b). There exist three with alder trees. In rice fields, Anabaena azollae is one of
distinct categories of N ­ 2-fixing biofertilizers: symbiotic the most promising nitrogen biofertilizer. As an alternative
­N2-fixing biofertilizers, free-living ­N2-fixing biofertilizers, to commercial nitrogen fertilizer, Azolla was taken into con-
and associative symbiotic N ­ 2-fixing biofertilizers (Patel and sideration by several researchers (Kour et al. 2024c). A study
Patel 2023). conducted that, when compared to rice fields without Azolla,
A study concluded that ­N2-fixing bacterium identified Azolla increases rice yield by 10% to 25% (Kour et al. 2020).
as Azotobacter vinelandii was isolated from rhizospheric
region of rice and application of this bacterium on same Free‑living nitrogen‑fixing biofertilizer
crop improve growth (Sahoo et al. 2014). According to a
report, ­N2-fixing bacterium Herbaspirillum seropedicae was Microorganisms (both free-living and endophytic) such
evaluated on maize crop and result showed significant as Azotobacter, Cyanobacteria, Azospirillum are found in
enhancement in growth and nutrient content (Alves et al. non-symbiotic or free-living biofertilizers. The primary
2015). Another study evaluated the effect of N ­ 2-fixing bac- free-living ­N2-fixing bacteria found in both neutral and
terium namely Pseudomonas stutzeri on rice crop. The result alkaline soils are Azotobacter, which are members of the
showed that inoculation of bacterial strain improves growth Azotobacteriaceae family (Bhattacharyya and Jha 2012). It
Biologia

fixes atmospheric nitrogen in non-leguminous crops, such as solubilize the phosphate by both the organic approach and
vegetables, cotton and rice. According to Noar and Bruno- the proton-extrusion mechanism.
Bárcena (2018), the bioinoculation of Azotobacter improves A study reported that rhizobacteria, namely Enterobacter
wheat crop growth, protein content, grain production, and sp., Serratia sp., Acinetobacter sp., and Enterococcus sp.,
biomass. Numerous Azotobacter species have been recog- demonstrated P-solubilizing ability and were used as bioin-
nized to produce phytohormones, vitamin B complex, and oculants in peanut (Arachis hypogaea). The study showed
other bioactive compounds that serve as biocontrol agents to that the application of bacterial strains improved plant
prevent root infections. Primitive microorganisms known as growth (Anzuay et al. 2015). Another study reported that
Cyanobacteria were initially found in rice fields and used as P-solubilizing bacterial strains, namely Pseudomonas beteli
biofertilizers. Since then, numerous variations of Cyanobac- and Pseudomonas lini, enhanced the growth of wheat plants
teria have been discovered in a variety of different environ- (Ehsan et al. 2016). Another study concluded that the appli-
ments. Nostoc, Calothrix, Anabaena, and Aulosira are the cation of P solubilizing bacterial strains identified as Acine-
main photosynthetic, free-living nitrogen fixers known as tobacter sp. and A. baumannii improved tomato plant growth
Cyanobacteria (Noar and Bruno-Bárcena 2018). Cyanobac- (Zhang et al. 2017). Another study revealed that application
teria improve soil fertility by secreting several compounds of Bacillus sp. improve growth and yield of rapeseed (Valetti
that promote plant growth, such as vitamins, amino acids, et al. 2018). A study by Rezakhani et al. (2019) reported that
phytohormones. two P-solubilizing bacterial species, Bacillus simplex and
Pseudomonas sp., enhanced the length and biomass of wheat
Associative symbiotic nitrogen‑fixing biofertilizers plants. A study reported that, application of P-solubilizing
microbes as biofertilizer namely Pseudomonas sp., Sphingo-
Azospirillum is associated with various grasses; it is primar- bacterium suaedae, Bacillus pimilus and B. cereus enhance
ily investigated under associative symbiotic nitrogen fixers maize growth (Benbrik et al. 2020). Another study by Kaur
(Kumar et al. 2022). There are currently 17 species of Azos- et al. (2021) reported that, inoculation of P-solubilizing bac-
pirillum known, and the two most extensively researched teria namely Pseudomonas gessardii improve the growth of
are Azospirillum lipoferum and A. brasilense. These species barley (Hordeum vulgare L.). Another study concluded that,
have been sorted out from both the soil and the aerial regions co-inoculation of P-solubilizing bacteria Pseudomanas ges-
of plants that are capable of fixing nitrogen. These microbes sardi and Erwinia rhapontici enhance growth of amaranth
not only fix nitrogen but also generate gibberellins, cyto- crop (Devi et al. 2022). A study concluded that, P-solubiliz-
kinins, and indole-3-acetic acid (IAA) (Groppa et al. 2012). ing bacterium identified as Rhizobium sp. increased the soil
According to reports, Azospirillum also promotes alterations P availability in maize plant (Beltran-Medina et al. 2023).
in cell wall flexibility and osmotic adaptations, which aid In an investigation, application of Serratia marcescens,
plants in surviving under stressful circumstances. Pseudomonas azotoformans, Staphylococcus aureus, and
Pseudomonas sp. improve growth wheat (Gabba et al. 2024).
Phosphorus solubilizing biofertilizers
Potassium solubilizing biofertilizers
Phosphorus (P) is an essential element that is vital to plant
growth and development, with plants containing approxi- The second most prevalent and significant nutrient for plants
mately 0.2% of phosphorus and currently the nutrient that is after phosphorus and nitrogen is potassium (K). K is abun-
least transportable and available to plants in most soil con- dant in the soil compared to other elements, but only 1–2
ditions (Kour et al. 2020). The transformation of phosphate percent of it is available to plants; the rest is present as min-
from its insoluble forms to its soluble forms requires micro- eral K, making it unavailable to plants (Singh et al. 2020a).
organisms. Phosphorus solubilizing bacteria (PSB) convert Thus, the soil solution needs to have its K levels continuously
insoluble phosphates like H ­ PO4 and H ­ 2PO4 into soluble replenished. It is essential to the development and growth of
forms through a variety of mechanisms, including chela- plants. K also plays a part in regulating stomata movement,
tion, ion exchange events, and the synthesis of organic acids starch synthesis, and root growth, among other functions
(Vassilev et al. 2015). PSB make up 1–50% of microbial (Raji and Thangavelu 2021). Low potassium levels lead to
populations, while fungi contribute only 0.1–0.5% to these poor root development, sluggish growth rates, minimal seed
activities. The cation bound to the phosphate can be chelated production, increased susceptibility to diseases, and reduced
by the PSB and transformed into a soluble form usable by yields (Rajawat et al. 2020). K mostly exists as silicate min-
plants, such as organic acids containing hydroxyl (gluconic) erals in the soil continuum that are inaccessible to plants and
and carboxyl (ketogluconic) groups (Soumare et al. 2021). are only released when the minerals weather or become solu-
The produced acids also reduce the pH of the soil, allowing ble (Brindavathy 2022). K is released from many insoluble
the plants to access the bound phosphate. Microorganisms soil compounds by rhizospheric microorganisms. Numerous
 Biologia

studies found that adding K-solubilizing microorganisms to important role in crop productivity due to the global soil Zn
plant seedlings or seeds themselves increased germination deficiency (Kour et al. 2024a).
rate, growth, yield, seedling vigor, and K uptake (Devi et al. A study concluded that, application of Zn-solubilizing
2024a). Numerous bacterial and fungal strains have been Bacillus sp., and Pseudomonas sp enhance growth of maize
shown to solubilize the insoluble K into soluble forms by plant (Goteti et al. 2013). Another study by Wang et al.
a variety of mechanism, including the generation of acids, (2014) concluded that, application of efficient Zn-solubi-
chelation, acidolysis, complexolysis, and exchange processes lizer Burkholderia sp., Enterobacter sp. Variovorax sp., and
(Verma et al. 2017). Sphingomonas sp. improve growth of Japonica rice (Wang
A study documented that, K-solubilizing bacterial species et al. 2014). According to a report by Hussain et al. (2015),
identified as Pseudomonas nitroreducens, Bacillus subtilis, Zn-solubilizing Bacillus sp. inoculated on maize crop
and Burkholderia cepacia improved growth and soil fertility resulted enhancement in growth and physiological param-
of tea plant (Bagyalakshmi et al. 2017). In a report, applica- eters (Hussain et al. 2015). Application of ZSB namely
tion of K-solubilizing bacteria increase growth and yield in Acinetobacter sp. improve growth and yield of rice crop
ryegrass (Xiao et al. 2017). A study revealed that applica- (Gandhi and Muralidharan 2016). A study reported that,
tion of K- solubilizing Pantoea agglomerans, Pseudomonas bacterial genera identified as Azotobacter, Pseudomonas,
orientalis, and Rahnella aquatilis enhance growth and yield Azospirillum,and Rhizobium increase Zn uptake in wheat
of rice plant (Khanghahi et al. 2018). In a different report, crop (Naz et al. 2016). According to a study by Ullah et al.
bacterial species namely Acinetobacter pittii, Cupriavidus (2020), application of ZSB namely Enterobacter sp. improve
oxalaticus, Ochrobactrum cicero and Rhizobium pusense zinc efficiency and productivity of desi chickpea. Two poten-
were showing K- solubilizing ability and application of tial Zn-solubilizing efficient strains identified as Acinetobac-
these strains in paddy plant enhance growth and physi- ter baumannii and Burkholderia cepacia increase Zn uptake
ological parameters (Ashfaq et al. 2020). According to a and growth promotion of maize (Zea mays L.) (Upadhyay
study, K- solubilizing bacterial strain namely Bacillus cereus et al. 2021). A study concluded that Bacillus aryabhattai
was inoculation on rosemary plant and a result showed that improve Zn content in rice plants (Oryza sativa) (Prathap
application of B. cereus significant enhance growth and et al. 2022). A study reported that, application of ZSB iden-
physiological parameters as compared to untreated control tified as Brevibacillus brevis enhances growth of wheat Triti-
(Sharma et al. 2024a). A study by (Kour et al. 2023) reported cum aestivum (Devi et al. 2024c).
that, inoculation of K-solubilizing bacterial strain identi-
fied as Acinetobacter calcoaceticus improves the growth of Selenium solubilizing biofertilizers
onion. A study documented that silicate solubilizing bacteria
identified as Bacillus albus improve growth of tomato plant Selenium (Se) is another micronutrient for plant growth and
(Shukla et al. 2023). A study by Kaur et al. (2024) reported a target for biofortification in crops. It plays multiple physi-
that, application of K-solubilizing bacterium identified as ological roles in a variety of biological systems (Ye et al.
Pseudomonas gessardii improves the growth of eggplant 2020). Se is an essential micronutrient for both human and
(Solanum melongena L.). plants, but taking too much of it can be hazardous while
giving plants a number of positive benefits. The soil con-
Zinc solubilizing biofertilizers tains 0.4 mg/kg of Se. Selenate and selenites are the two
most prevalent inorganic forms of selenium that are found in
Zinc (Zn) is one of the vital elements needed in tissues for soils (Lanza and Dos Reis 2021). A variety of environmen-
plant development and reproduction, although exists in tis- tal factors including pH, ionic strength, ion concentration,
sues at relatively low amounts (5–100 mg/kg) (Sharma et al. and others, selenium forms are frequently more intensely
2023). Zn deficiency is a prevalent occurrence in soil due retained in soil colloids (Sharma et al. 2015). Se shields
to intense farming practices, excessive fertilizer application the plant from low-level abiotic stressors like desiccation,
applied unevenly, and poor soil health (Khan et al. 2022). drought, cold and salinity. There are various oxidation states
If the underlying causes are ignored, it is predicted that by of Se, such as selenium, selenide ­(Se2−), selenite ­(SeO32−),
2025, the percentage of people deficient in Zn might rise and selenium ­(SeO42−). The primary factors influencing the
from 42 to 63%. A Zn deficit in plants affects grain yield, types of Se in soil were pH and redox potential, which were
root development, pollen generation, water uptake and trans- determined by the amount of organic matter and clay in the
port, and delayed shoot growth, diminished leaves, chlorosis, soil (El-Ramady et al. 2014).
weakened membrane integrity, and heightened susceptibil- In naturally occurring soil conditions, selenite and
ity to fungus, heat, and light (Rana et al. 2023). Human selenate are the main organic forms. In contrast to selenite
Zn insufficiency can also result from eating grains low in found in natural and acidic soils, which is easily adsorbed
zinc. Zinc solubilizing biofertilizers (ZSB) have an equally onto oxyhydroxides, selenite discovered in alkaline soils
Biologia

has higher mobility (Lopes et al. 2017). The soil contains (Syed et al. 2023). A study concluded that, endophytic
selenium in its soluble form. Plant cells have been shown to siderophores producing bacterium identified as Spinacia
transport information across their plasma membranes (Wang Oleracea enhance nutrient uptake (Misra et al. 2024).
et al. 2022). It might absorb via the sulfur assimilation routes
to become Se-amino acid. Se-containing biofertilizers, such
as selenite or selenate, can be sprayed over the soil as a Phytohormones producing biofertilizers
foliar spray to enhance the amount of Se in the plant's edible
sections and produce Se-biofortified crops (Hossain et al. Plant hormones, also known as phytohormones, is released
2021). A study reported that Se-solubilizing bacterial strain by both microbes and plants and serves as essential for plant
identified as Priestia sp. enhance 19% higher biomass and development (Chaudhary et al. 2022). The synthesis of plant
75% higher Se concentration in rice (Oryza sativa L.) than hormones is an essential feature of the beneficial micro-
untreated control (Lin et al. 2022). Another study have been organisms that also produce gibberellins, cytokinin, and
reported that, application of Se-solubilizing bacterial strain indole-3-acetic acid (IAA). Plant cells used auxins to aid in
namely Exiguobacterium sp. enhance growth and biofor- their differentiation and division. Plants that have cytokinin
tification in mustard plant (Marfetán et al. 2023). A study do not experience early leaf senescence. Another hormone
documented that Se-solubilizing bacterial strain identified that plants produce when under stress is called abscisic acid.
as Bacillus mycoides improves growth and physiological Gibberellins play a role in flowering, fruiting, shoot elonga-
parameters as compare to uninoculate control. The study tion, and seed germination. Under stressful environmental
document that, application of microbial consortium com- situations, microorganisms typically produce these hor-
prising of ­N2-fixation, P, K, and Se-solubilizing strains and mones to protect plants by adjusting the quantity of phyto-
single inoculum of these enhance growth of wheat plant hormones in the host plants. Several microorganisms that are
(Devi et al. 2024b). found in the rhizosphere, endosphere and phyllosphere are
well identified to produce hormones for the purpose of root
uptake or to regulate hormone balance in plants to promote
Siderophores producing biofertilizers growth and help to cope up with several stresses. Das et al.
(2023) observed that PGP bacterial strains such as Bacil-
A micronutrient termed iron (Fe) is essential for several lus subtilis, Paraburkholderia caribenis and Pseudomonas
processes in plants, including photosynthesis, respira- palmensis have great capacity to produce IAA, which helped
tion, chlorophyll, and other enzymatic activities (Rout to increase the nutrient uptake, root biomass, shoot weight
and Sahoo 2015). In an aerobic environment, the most in lentil plant. A study concluded that, IAA producing bac-
common form of iron that is inaccessible in nature is ­Fe+3 terium identified as Ochrobactrum haematophilum promote
that is most likely an insoluble oxyhydroxide or hydroxide apple growth (Jiang et al. 2023). A study reported that, cyto-
complex. Thus, microbes are the producers of siderophores kinin-producing bacterium identified as Pseudomonas fluo-
that constitute low molecular weight iron-binding protein rescens improve tomato (Solanum lycopersicum) growth by
molecules (Ellermann and Arthur 2017). Siderophores are enhancing resistant against drought stress (Mekureyaw et al.
soluble in water molecules that can be found both within 2022). Another study reported that, cytokinin producing
and outside of cells. ­Fe+3 are reduced to ­Fe+2 inside the bacterium namely Azospirillum brasilense increase yield,
cytoplasmic membrane by the siderophores within the antioxidant activity, and physiological parameters of wheat
bacteria, and ­Fe+2 are then transferred within the cyto- (Zaheer et al. 2022). A study concluded that, gibberellin
plasm by gating mechanisms (Aloo et al. 2022). The bac- producing Rhodobacter sphaeroides enhance photosynthetic
teria provide the host plant with this type of iron that is rate, chlorophyll content, stomatal conductance, and transpi-
readily available for growth. By allowing microorganisms ration rate (Kang et al. 2022).
to release the chelating chemical, siderophores enable
plants to assimilate iron. A study documented that, sidero-
phores producing bacterium identified as Bacillus aerophi-
lus enhances growth and copper accretion in Helianthus Delivery methods of biofertilzers
annuus L. (Kumar et al. 2021a). Another study reported
that, application of siderophore producing bacterial strain Vast arrays of crops are grown under various agro-climatic
namely Pseudomonas fluorescens promote the growth of conditions, and each crop has distinct planting and growing
cucumber (Al-Karablieh et al. 2022). In an investigation, habits (Grigorieva et al. 2023). Multiple strategies for apply-
siderophores producing bacterial strain isolated and identi- ing biofertilizer have been devised, taking into account the
fied as Pseudomonas fluorescence and Trichoderma spp. planting conditions and the parts that can be fertilized (root,
enhance growth of chickpea by lowering cadmium uptake seed, and shoot).
 Biologia

Foliar based applications dolomite; or calcium carbonate (Bashan et al. 2014). This
mechanism shields microorganisms from unfavorable envi-
In order to manage foliar diseases or promote plant growth, ronmental factors and the damaging effects of chemical fer-
liquid formulations are typically applied to the foliar portion tilizers. A study documented that, seed of Aegilops kotschyi
of the plant. The crop canopy's microclimate is the primary and wheat were coated by seed treatment method. The result
determinant of its effectiveness (Preininger et al. 2018). Its of the study revealed that application of biofertilizer in both
effectiveness is primarily reliant on the crop canopy's micro- of the crops enhance growth and physiological parameters
climate. The effectiveness and survivability of antagonist (Negi et al. 2022b).
phylloplane are significantly influenced by several param-
eters (Hernández-Fernández et al. 2021). ­N2-fixing microbes Soil based applications
fix atmospheric nitrogen close to its site of assimilation; and
phylloplane's food source is leaf leachates; biofertilizers Soil inoculation should be performed when a significant
use this food to eradicate cuticle-degrading pathogens by number of microbial strains are being introduced to the
means of competitive inhibition. A case study reported that, soil directly (Mawarda et al. 2020). Granular forms of soil
Klebsiella planticola and Enterobacter spp. used as foliar aggregates, talc, perlite, peat, or other granular materials are
biofertilizer which observed that, applications of bioferti- typically utilized in this approach. Granules sized between
lizer lower doses of mineral NPK fertilizers increased the 0.5 and 1.5 mm are preferable. Soil inoculation typically
yield of maize and wheat (Latkovic et al. 2020). A study entails scattering granules beneath, above, and beside the
documented that, foliar application of C. vulgaris enhanced seed in the seedbed. When using liquid biofertilizers, seed-
P. vulgaris growth and crop production (Refaay et al. 2021). lings planted in furrows can be sprayed with the inoculants;
Another study reported that, foliar application of Azotobac- hydroponic systems are another option. In addition to shield-
ter enhance growth, nutritional value of lettuce (Razmjooei ing inoculants from the damaging effects of fertilizer and
et al. 2022). A study concluded that, Chlorella vulgaris can preventing damage to the seed coats, soil treatment allows
be applied as biofertilizer through foliar method in tomato for control over the inoculants application rate and location
replacing conventional chemical fertilizer (Suchithra et al. (O'Callaghan et al. 2022). When employing seeding machin-
2022). According to a study by Dhote et al. (2023), foliar ery, there is less chance of losing portion of the inoculant.
application of microbes used as biofertilizer enhances Compared to seed treatment, soil inoculation increases the
growth and yield of chickpea Cicer arietinum L. Another likelihood of seeds coming into touch with a higher concen-
study revealed that, cyanobacterial formulations used as tration of biofertilizer (O’Callaghan 2016). However, there
foliar biofertilizer enhance plant growth and fruit quality in are certain drawbacks to this approach, most of which are
tomato under controlled condition (Bhardwaj et al. 2024). technical in nature. It requires specialized equipment and
bigger amounts of biofertilizer, which necessitate additional
Seed priming applications space for storage and transportation. There is a concurrent
relationship between these elements and increased expen-
Seed treatment is still the most popular method of applying ditures (Bashan et al. 2014). Granule-based soil inocula-
biofertilizers due to its convenience of use and low product tion has primarily been employed in affluent nations with
requirement for inoculation. Seed coating, slurry, and dust- the availability of sophisticated fertilization equipment and
ing are the three methods that inoculant can be applied to supplies.
the seeds (Asif et al. 2018). Direct mixing of the inoculant
and dried seeds occurs during dusting. The least successful
approach is believed to be this one since it could cause the Challenges of biofertilizer
microbes to stick to the seeds only weakly. Wetted seeds or commercialization
water alone are combined with bioinoculant before seeds are
added to the slurry (Kumar et al. 2022). In order to ensure The most prevalent feature among various biofertilizers is
that every seed has the right amount of microbes coated on the unpredictable nature of their results. It is necessary to
it, non-toxic commercial goods, gum arabic, carboxy methyl increase the performance of biofertilizers consistently (Khan
cellulose, sucrose solutions, and vegetable oils are employed et al. 2023a). The method used to develop the bioinoculants
as adhesives. If the biofertilizer isn't adhesive-free, you has a major impact on performance. Although the tech-
should mix either 1% milk powder or a 25% molasses solu- nology used to produce biostimulators is inexpensive and
tion with the suspension (Brahmaprakash et al. 2017). The environmentally benign, there are a number of drawbacks
final approach is to blend the seeds with the inoculant slurry that restrict its use and acceptability (Kumawat et al. 2021).
and then coat them with a finely ground inorganic inert These constraints can be of a technical, organizational struc-
material, such as talc; lime; clay; rock phosphate; charcoal; ture, budgetary, environmental, quality control, or human
Biologia

resources-related nature (Aloo et al. 2021). The application absence of superior carrier material can have a significant
of the appropriate amount of microbe based fertilizers as impact on the efficacy of biofertilizers (Malusà et al. 2016).
well as production and marketing technologies is impacted Reduced shelf life of biofertilizers is mostly caused by the
by the numerous restrictions and limitations. absence of an appropriate carrier material that would allow
potentially advantageous microorganisms to proliferate (Rai
Improvement of bacterial strains for mass et al. 2023). Two possible ways to lessen these disadvantages
production are the implementation of an integrated sterilizing facility
and the use of suitable sterilized carriers (Mącik et al. 2020).
A major limitation is the use of inappropriate, ineffective Another is identifying, through recommendations, the most
bacterial strains for the development of biofertilizers, which popular carrier materials across different nations and provid-
can result in an insufficient population density of viable ing producers with easy access to them.
microorganisms (Aloo et al. 2021). One of the major dis-
advantages is the lack of native microbial strains because Quality of bioinoculants
bioinoculants are soil and host-specific. Furthermore, the
chosen candidate microbial strains need to be able to sur- Understanding fundamental physiological and biochemical
vive in both liquid and solid carriers and exhibit competitive processes is crucial for producing bio-inoculants, as it pre-
efficiency against other strains in a variety of environmental serves the quality of the inoculants and, consequently, their
conditions (Sahu and Brahmaprakash 2016). Farmers also therapeutic effects (Khan et al. 2023a). Poor germination
find it unacceptable to apply biofertilizer when the envi- may result from the biofertilizer solution removing the seed
ronment is not at its ideal temperature or in problematic coat from the seed. The commercialization of biofertilizers
soil conditions (acidic, saline, or alkaline). These factors may be severely hampered by improper microbial inoculant
combined make bioenhancers, biostimulators, and biofer- formulation (Malusà et al. 2016). Nonetheless, the need for
tilizers less effective in these situations. Biofertilizers also premium inputs drives creative advancements in the effec-
cause mutations during bulk multiplication, which increases tiveness of biofertilizers. The following considerations need
production costs and quality control (Nath Bhowmik and to be made in order to develop high-quality bioinoculants:
Das 2018). A comprehensive investigation of this matter employing bioengineering techniques for efficient micro-
is required in order to eradicate such undesired alterations bial strain improvement; exchanging microbial inoculants
during the manufacturing of biofertilizer for particular crops. between countries with similar agro-climatic conditions and
investigating their potential beneficial effect of strains for an
Technical personnel and quality of production units individual crop; monitoring the efficacy of biological inocu-
lants throughout storage in order to avoid natural damage
Technical issues with the production of biofertilizers might (Suyal et al. 2016).
be caused by staff members who are inadequate, inexpe-
rienced, and lack technical qualifications (Kumawat et al. The shelf life of bioinoculants
2021). One significant barrier is that consumers are not
receiving the necessary training regarding the application Due to its brief shelf life, typically six months, it requires
of biofertilizers, which results in a lack of information and appropriate storage space. Because they are unable to keep
expertise about the use of bioinoculants (Odoh et al. 2020). the microbiological product for an extended period of time
An additional major barrier is that extension workers and under ideal conditions, this discourages business owners
customers are unaware of the inoculation procedure due to from creating more than what they could sell right away to
poor private management of the utilization process and a customers who buy more than they need right away (Negi
lack of time to apply biofertilizers during sowing (Odoh et al. 2024a; Suyal et al. 2016). The challenges facing the
et al. 2020). Most marketing staff members are ignorant of biofertilizer industries development are typically related to
appropriate application techniques. Insufficient technical poor demand brought by farmers ignorance of the mecha-
personnel with the necessary qualifications in the biofer- nisms underlying plant growth enhancement initiatives
tilizer production units could result in improper handling (Kumawat et al. 2021). Production costs are often still a
in mass manufacturing, transit, and storage establishments problem in these situations, as are low delivery methods
(Raimi et al. 2021). and limited demand that may be related to certain handling
and storage conditions. The shelf life of the inoculants, han-
Quality of carrier material dling, storage conditions, carrier material quality, and the
existence of contaminants all have an impact on how quickly
The use of different carrier materials by different produc- biofertilizers respond (Sahu and Brahmaprakash 2016). Con-
ers without considering the quality of the resources or the sequently, it is crucial to extend the shelf life of regionally
 Biologia

produced bioenhancers under suitable storage settings in method of applying bioinoculants, and production strate-
order to guarantee the viability of the microbial product for gies and expertise pertaining to enhanced biological ferti-
an extended duration (Kaur et al. 2022). lizer production techniques (Gautam et al. 2021). Some of
the actions that should be taken include providing technical
Facilities for production and storage of inoculants training to producers on the production of biofertilizers and
packet their quality control, providing manufacturers with techni-
cal information and projects, providing for consumer and
One of the biggest barriers to development is the absence agricultural extension personnel to receive training in order
of facilities that are appropriate for manufacturing (Sahu to popularize the technology, and establishing more effective
and Brahmaprakash 2016). Another issue is caused by the and extensive knowledge distribution (Kumawat et al. 2021).
insufficient availability of inputs at the right time. One key
factor threatening the quality of bio-enhancers is the lack of
facilities for storing packets of bioinoculants, that must be Importance of biofertilizers
kept cool and away from heat sources and direct sunlight
(Kumawat et al. 2021). Insufficient cold storage facilities Role of biofertilizers for agricultural sustainability
can subject bioinoculants to excessive heat, which would be
detrimental to the potent microbes that were inoculated. The The unregulated use of synthetic fertilizers in intensive agri-
consumer community faces uncertainty and risk due to the culture has a negative impact on the quality and health of
insufficient marketing facilities and lack of regular aware- the soil. Thus, a sustainable strategy is needed to prevent
ness regarding the usage of biofertilizers (Kumawat et al. the soil from deteriorating further, and biofertilizers pro-
2021). A great move toward enhancing the infrastructure vide an effective and practical solution for this (Kumar et al.
of the production unit is to hire soil microbiologists and a 2018). Thus, the two main challenges of today are feeding
qualified technician to oversee the manufacture of bioferti- the world's population and reducing chemical consumption.
lizer, control quality, and build suitable storage facilities in Reducing the use of harmful agrochemicals while enhancing
production centers (García de Salamone et al. 2019). fertility and health of the soil is a key component of sustain-
able agriculture (Kawalekar 2013). To enhance soil health
Equipment, laboratory, production, and storage and structure, the ideal ratio of inorganic and organic inputs
space must be maintained. The primary way to raise soil organic
carbon for the maintenance of soil quality and future agrar-
The lack of necessary equipment, power supplies, and suf- ian efficiency is by using natural fertilizers (Giri et al. 2019).
ficient inputs increases labor costs, as mass production is Currently, excessive use of chemical fertilizers has exac-
labor-intensive and slow in this context (Raimi et al. 2021). erbated soil and water contamination, decreasing land fertil-
It is crucial to have enough space for production, refrigerated ity (Negi et al. 2022b). The destruction of beneficial insects
storage, and laboratories. More acreage is needed to expand and bacteria predisposes the crops to illness. Biofertilizers
the biofertilizer producing plant. Other significant problems are a crucial part of organic farming since they support
include the lack of a subsidy structure and the appropriate soil sustainability and long-term fertility. They contribute
cost of bioinoculant trading (Mahanty et al. 2017). Nonethe- to raising soil productivity per unit area, enhancing farmers'
less, the growing markets for bioinoculants and consumer financial circumstances (Kaur and Purewal 2019). Because
education and knowledge on the use of biofertilizers have of its delayed release pattern, applying biofertilizers offers
made it easier and more motivating for entrepreneurs to enter long-term benefits. Thus, these biological inoculants are
the biofertilizer industry. a financially and environmentally responsible way to
encourage sustainability and productivity. Plant nutrition
Educational and training by extension specialist is enhanced by biofertilizers via a number of direct and
for biofertilizers application indirect methods (Sharma et al. 2024c). It directly aids in
the availability of nutrients through processes like nitrogen
The main problem is typically the lack of training and inad- fixation or solubilization, or it enhances root growth through
equate knowledge of field workers about the application of the synthesis of hormones like cytokinin and auxins (Uribe
biofertilizer, especially with regard to sustainable agriculture et al. 2010). Through the synthesis of different chemical
systems (Mahanty et al. 2017). Aside from the limitations metabolites or the induction of systemic resistance, it indi-
of individual resources and quality control, other signifi- rectly enhances the plant's ability to withstand disease. By
cant issues that must be addressed are ignorance of different breaking down soil debris and consequently increasing
agrochemicals, inoculum concentration, the proper time and nutrient content through mineralization or decomposition,
Biologia

it also aids in improving the structure of soil (Singh et al. Role of biofertilizers in the alleviation
2016). Overall, it's a practical strategy that boosts the output of environmental stresses
of small and marginal farmers because it provides a renew-
able and affordable source of nutrients to support sustain- A multitude of biotic and abiotic elements influence the
able agriculture in place of synthetic fertilizers (Sahoo et al. nation's crop yield. There have been several recommenda-
2012) (Table 1). tions made for treating stress in crops, and one of the most
significant of these has been the use of microbes as bioferti-
Role of biofertilizers in climate‑smart agriculture lizers (Chaudhary et al. 2022). Plant growth is enhanced by
metabolites such as exopolysaccharides, which enhance soil
Crops are experiencing significant biotic and abiotic stress structure and reduce plant water stress (Naseem et al. 2018).
due to changing climatic conditions. Beneficial microbes As an alternative to diverse synthetic pesticides, biofertiliz-
known as "biofertilizers" can help build and impart toler- ers can also function as effective biocontrol agents against
ance against a variety of abiotic stressors by adapting to a variety of phytopathogens (Singh et al. 2019). In order to
changing environmental conditions (Mohanty and Swain increase its resistance to pathogen attack, they are known
2018). In contrast to synthetic substances that might even- to produce a wide range of compounds, including hydro-
tually cause a shift in the climate, they offer a sustainable gen cyanide, ammonia, siderophores, and antibiotics. By
method of addressing climate change (Ajala et al. 2022). enhancing soil health and activating various stress-tolerant
Microorganisms that generate biofertilizers have an active mechanisms, such as antioxidant enzymes, biofertilizers
role in soil formation, soil structure maintenance, fertility support plant growth. They are particularly beneficial under
and overall ecosystem resilience (Bera et al. 2022). They various stress conditions (Kumar et al. 2021b). Thus, hostile
aid in maintaining the health of the soil and discourage the soils can be rendered habitable with the application of these
use of artificial chemicals, which eventually reduce the pro- organic compounds, improving plant growth at a reasonable
ductivity of the ecosystem. Biofertilizers assist plants with cost and in a way that is environmentally sustainable (Arora
absorbing nutrients, managing their growth, and modifying et al. 2018).
their physiological and biochemical characteristics, all of
which help them become more resistant to disease (Singh
et al. 2024). The global commercialization and practical
Since methane is produced in submerged environments, applications of biofertilizers
it is a crucial component of rice farming. It is a green-
house gas that raises the earth's temperature by retaining The commercialization of biofertilizer began in 1895 when
heat in the atmosphere and contributing to global warming Nobbe and Hiltner marketed the endophytic rhizobia product
(Hussain et al. 2020). Methane is the only source of carbon under the "Nitragin" brand. Rhizobium was initially com-
and energy for methanotrophs, often known as methane- mercialized in India by N. V. Joshi, who was responsible
oxidizing bacteria (MOB). They can serve as potential for promoting plant growth in leguminous crops (García-
sinks for capturing methane and are found in aerobic soils. Fraile et al. 2015). The Ministry of Agriculture launched
Although they might account for 15% of the world's total the National Project on Development (NAD) of bioinocu-
methane emissions, they are quite sensitive to change in lants (NPDB) during the ninth Five-Year Plan. This initia-
the neighboring environment (Guerrero-Cruz et al. 2021). tive aimed to popularize and promote the technology used
Additionally, because of their sluggish development rate in biofertilizer production. Additionally, a center for the
and adhesion to soil particles, they are challenging to iso- development of biofertilizers was established, along with
late. A novel bacterium called Methylacidophilum can six regional centers, and a national standard protocol for
thrive in harsh environments with high acidity and high quality control was developed for various biofertilizers
heat and can use methane gas as fuel (Schmitz et al. 2021). (Ghosh 2004). The Indian government and state organiza-
Moreover, several microbes have the capacity to absorb tions implemented various initiatives to increase bioinocu-
atmospheric carbon dioxide and convert it into calcium lant production, offering financial assistance and subsidies
carbonate. Nitrous oxide is another greenhouse gas that at different levels.
can be transformed into nitrogen gas by denitrifying bac- A new ruling on the regulation of biofertilizer production
teria (Zhan et al. 2021). By assisting in the mitigation of and marketing quality criteria concerning various types of
greenhouse gas emissions, these bacteria help to reduce the potentially advantageous microorganisms was issued by the
emissions of greenhouse gases per unit of food production. Ministry of Agriculture (Barquero et al. 2019). Seven criteria
Consequently, all of these microbial activities play a signifi- for quality control should be met by the bioproducts: maxi-
cant role in mitigating the effects of environment alteration mum viable population density, suitable moisture percentage
(Zhang et al. 2016). by weight of carrier-based products, levels of contamination,
 Biologia

Table 1  Plant growth promoting attributes of plant associated microbes as biofertilizer

S.N Microbiomes Host Plant growth promoting attributes References

1. Achromobacter xylosoxidans Rice Indole acetic acid production (1.20–60.13 μg/ Jhuma et al. (2021)
mL), phosphorus solubilization (0.02–1.81 μg/
mL), nitrogen fixation (70.24–198.70 μg/mL)
2. Acinetobacter guillouiae Wheat Nitogen fixation, solubilization of phosphorus Rana et al. (2020)
and potassium, production of sideophores,
­NH3, HCN and IAA
3. Acinetobacter schindleri Maize Nitrogen fixation Abadi et al. (2021)
4. Bacillus albus Wheat Phosphorus solubilization, IAA, siderophores Azaroual et al. (2020)
and 1- aminocyclopropane-1-carboxylate
deaminase production
5. Bacillus amyloliquefaciens Soil Salt tolerance up to 7.5%, production of IAA and Sharma et al. (2021)
­NH3
6. Bacillus aryabhattai Wheat IAA production, solubilization of inorganic Pang et al. (2022)
phosphorous, organic phosphorus and potas-
sium
7. Bacillus australimaris Rice Auxins production, phosphorus solubilization, Hernández et al. (2023)
potassium solubilization, and production of
siderophores
8. Bacillus cereus Wheat Nitrogen fixation, phosphorus, and zinc solubi- Zahra et al. (2023)
lization
9. Bacillus pumilus Artemisia Vulgaris Efficient drought tolerant, abscisic acid produc- Shaffique et al. (2024)
tion (2.18 ± 0.05 ng/mL), phosphorus solubi-
lization
10. Bordetella muralis Corn Nitrogen fixation and production of siderophores, Sukweenadhi et al. (2022)
IAA and ACC deaminase
11. Brachybacterium ginsengisoli Maize Nitrogen fixation Abadi et al. (2021)
12. Burkholderia cenocepacia Rice Production of IAA and N ­ H3, solubilization of Sherpa et al. (2021)
phosphorus and potassium, nitrogen fixation
13. Burkholderia cepacia Sweet corn The phosphorus-solubilization (125.33 ± 8.78 to Pande et al. (2020)
305.49 ± 10 μg/mL)
14. Cellulosimicrobium cellulans Corn Nitrogen fixation and production of siderophores, Sukweenadhi et al. (2022)
IAA and ACC deaminase
15. Chryseobacterium aquifrigidense Maize Nitrogen fixation Abadi et al. (2021)
16. Clavibacter michiganensis Maize Nitrogen fixation Abadi et al. (2021)
17. Comamonas testosteroni Soil High resistance and strong accumulation capac- Shi et al. (2021)
ity to cadmium
18. Curtobacterium oceanosedimentum Wheat Capability to tolerate high salt concentration Yadav et al. (2024)
(≥ 3.0 M NaCl)
19. Curtobacterium pusillum Wheat Capability to tolerate high salt concentration Yadav et al. (2024)
(≥ 3.0 M NaCl)
20. Enterobacter cloacae Rice Indole acetic acid production (1.20–60.13 μg/ Jhuma et al. (2021)
mL), phosphorus solubilization (0.02–1.81 μg/
mL) and nitrogen fixation (70.24–198.70 μg/
mL)
21. Enterobacter hormaechei Ziziphus lotus Production of IAA (550 mg/L) Fahsi et al. (2021)
22. Escherichia coli Wheat Nitrogen fixation, solubilization of P and Zn Zahra et al. (2023)
23. Ewingella americana Maize Nitrogen fixation, solubilization of P and K, pro- Rana et al. (2021)
duction of sideophores, ­NH3, HCN and IAA
24. Exiguobacterium indicum Wheat Capability to tolerate high salt concentration Yadav et al. (2024)
(≥ 3.0 M NaCl)
25. Exiguobacterium undae Wheat Capability to tolerate high salt concentration Yadav et al. (2024)
(≥ 3.0 M NaCl)
26. Klebsiella pneumonia Soil Nitrogen fixation and capability to tolerate high Khumairah et al. (2022)
salt concentration
Biologia

Table 1  (continued)
S.N Microbiomes Host Plant growth promoting attributes References

27. Kosakonia oryzendophytica Rice Production of IAA and N ­ H3, and solubilization Sherpa et al. (2021)
of P and K and nitrogen fixation
28. Ochrobactrum haematophilum Rice Nitrogenase activity (0.127–4.39 μmol Mir et al. (2022)
­C2H4/mg protein/h), IAA (12.24 ± 2.86 to
250.3 ± 1.15 μg/mL), phosphorus, zinc solubili-
zation and production of siderophores, HCN
and ACC deaminase
29. Ochrobactrum intermedium Soil Highest minimum inhibitory concentration of Renu et al. (2022)
150 mg/L of cadmium
30. Paenibacillus faecalis Soil Production of IAA, HCN, siderophores, and Waheed et al. (2024)
phosphorus solubilization
31. Paenibacillus hunanensis Rice Auxins production, phosphorus, and potassium Hernández et al. (2023)
solubilization, and production of siderophores
32. Paenibacillus polymyxa Rice Production of IAA and N ­ H3 and solubilization Sherpa et al. (2021)
of phosphorus and potassium and nitrogen
fixation
33. Pantoea agglomerans Soil Production of ACC, IAA, ­NH3, and phosphorus Bhise and Dandge (2019)
solubilization
34. Pantoea anthophila Rice Auxins production Hernández et al. (2023)
35. Pantoea deleyi Rice Auxins production Hernández et al. (2023)
36. Pantoea dispersa Chickpea Phosphorus solubilization and HCN production Tariq et al. (2023)
37. Paraburkholderia caribenis Lentil Production of IAA, siderophores and HCN Das et al. (2023)
38. Priestia aryabhattai Wheat Production of IAA, ACC deaminase, sidero- Shahid et al. (2022)
phores, and N ­ H3 and solubilization of phospho-
rus, and potassium
39. Priestia megaterium Soil Phosphorus solubilization, production of IAA Rehan et al. (2023)
and siderophores
40. Pseudochrobactrum asaccharolyticum Soil Production of IAA, HCN, siderophores, and Waheed et al. (2024)
phosphorus solubilization
41. Pseudomonas aeruginosa Rice Nitrogenase activity (0.127–4.39 μmol Mir et al. (2022)
­C2H4/mg protein/h), IAA (12.24 ± 2.86 to
250.3 ± 1.15 μg/mL), phosphorus, and zinc
solubilization
42. Pseudomonas fluorescens Soil Solubilization of inorganic phosphorus, nitrogen Ait Bessai et al. (2023)
fixation, production of plant defence com-
pounds
43. Pseudomonas frederiksbergensis Ziziphus lotus Nitrogen fixation and solubilization of potassium Fahsi et al. (2021)
44. Pseudomonas kribbensis Rice Production of IAA and N ­ H3 and solubilization Sherpa et al. (2021)
of phosphorus and potassium and Nitrogen
fixation
45. Pseudomonas palmensis Lentil Production of IAA, siderophores and HCN Das et al. (2023)
46. Pseudomonas plecoglossicida Soil Phosphorus solubilization, and production of Rehan et al. (2023)
IAA and siderophores
47. Pseudomonas stutzeri Soil Nitrogen fixation and capability to tolerate high Khumairah et al. (2022)
salt concentration
48. Rossellomorea aquimaris Soil Phosphorus solubilization, and production of Rehan et al. (2023)
IAA and siderophores
49. Serratia nematodiphila Corn Nitrogen fixation and production of siderophores, Sukweenadhi et al. (2022)
IAA and ACC deaminase
50. Stenotrophomonas pavanii Rice IAA production (1.20–60.13 μg/ mL), phos- Jhuma et al. (2021)
phorus solubilization (0.02–1.81 μg/mL) and
nitrogen fixation (70.24–198.70 μg/mL)
51. Streptomyces cinereoruber Soil Phosphorus solubilization, and production of Rehan et al. (2023)
IAA and siderophores
 Biologia

physical form, pH, carrier material particle size, and biofer- BSL-1 non-pathogenic microorganisms, wild-type microbes
tilizer efficiency (Maan and Garcha 2021). A minimum pop- (fungi and bacteria) derived primarily from soil and plants
ulation count of 5 × ­107 CFU/g and 1 × ­108 CFU/mL for the rather than from humans and animals are mainly used in the
solid and liquid carriers, respectively, must be maintained development of biofertilizers (Keswani et al. 2019). How-
in bacterial bioproducts. In goods that contain arbuscular ever, current studies often neglect the pathogenicity of these
mycorrhizal fungi (AMF), each gram of bio-product must strains, focusing primarily on their positive traits (Sacristán
have a minimum of 100 live propagules (Berninger et al. and García-Arenal 2008). Scientists may ignore potential
2018). For a bio-product designed to dissolve phosphate, a pathogenicity when evaluating bacteria believed to be non-
zone of solubilization of at least 5 mm on NBRIP's medium pathogenic, particularly those sourced from natural environ-
must be achieved. Additionally, the fertilizer should be able ments like soil and water (Khan et al. 2023b). Consequently,
to fix at least 10 mg N/g of sucrose utilized. Products with it has been discovered that a few of the microbes used in
AMF biostimulators should offer 80 infection sites in roots the biofertilizers belong to the BSL-2 family and exhibit
per gram of inoculums (Kumawat et al. 2021). opportunistic pathogenic behavior, there by posing a threat
The government's initiatives to support sustainable agri- to human health and the environment (Kumari et al. 2022).
culture have a significant hold on the biofertilizer enterprises Additionally, research into more effective isolation and
in Asia (Atieno et al. 2020). Approximately US$ 1.5 bil- selection methods has led to the discovery of new species
lion has been invested in the development of biofertilizer and genera that could potentially be used as biofertilizers.
products. The nation's sustainable agricultural practices are However, the absence of standard strains for pathogenic
currently improving, with about 1,000,000 hectares under comparability and the presence of potentially pathogenic
organic farming approaches (Gamage et al. 2023). Many organisms in hospital settings raise concerns about the secu-
governmental organizations, both public and private, are rity and appropriateness of these novel isolates for commer-
engaged in the manufacture of biofertilizer globally, includ- cial distribution (Brodmann et al. 2017). When used, these
ing in India. On average, approximately 45,000 tons of biofertilizers often multiply and become the predominant
biofertilizers are consumed annually in the nation, while bacteria. As a result, they interact with non-target plants,
less than half of that amount is produced (Kumawat et al. altering species composition and dominance, and in certain
2021). Biofertilizers are primarily produced in agroindus- situations, decreasing the biodiversity of neighboring plants
tries, state agriculture departments, national biofertilizer (Kardol et al. 2007). Moreover, these infectious diseases can
development centers, and both central and state agricultural significantly alter the local microbial community by pro-
institutions, as well as in the private sector (Mazid and Khan ducing growth regulatory compounds or antibiotics, which
2015) (Table 2). can alter the distribution and abundance of both beneficial
and pathogenic microorganisms in the ecosystem. Thus, this
might lead to changes in plant diversification and disruptions
Biosafety issue and risk assessment in nutrient cycles (Gneiding et al. 2008). Consequently, since
of microbial biofertilizers some of these isolates are linked to fatal illnesses, the use
of actinobacteria in biotechnology or agriculture requires
Biosafety refers to practices designed to reduce biological strict safety measures. Additionally, Arthrobacter oxydans
hazards to humans and the environment (Kimman et al. is frequently identified in clinical specimens, suggesting that
2008). Extensive research exists that provides compre- it may act as a pathogenic opportunistic organism (Mages
hensive guidance on handling microorganisms at different et al. 2008). It was later revealed that arbuscular mycorrhizal
biosafety levels (Emmert and Biosafety ATCoL 2013). The (AM) fungi may potentially be damaging to their host plants
term biosafety level (BSL) refers to the minimal safety pre- due to competition, which could disrupt plant growth and the
cautions that are advised when working with various risk dynamics of the ecosystem. Originally, it was believed that
groupings of microorganisms (Ta et al. 2019). Microor- AM fungi were mutualistic symbionts beneficial to plants
ganisms at BSL-1 are non-pathogenic and pose little harm (Fiorilli et al. 2024).
to individuals and the community (Keswani et al. 2019). It is strongly advised to conduct thorough categorization
While BSL-3 pathogens have a high individual risk but a and validation of PGP microbial strains under controlled
low community risk, BSL-2 microorganisms have a mod- settings before field application (Anand et al. 2022a). This
erate individual risk and a low communal risk. However, is crucial to prevent potential infections in the environment,
BSL-4 microbes pose a serious threat to individuals and even when the PGP microbes appear to be harmless (Shukla
communities since they are extremely pathogenic (Keswani et al. 2015). Adopting modern multidisciplinary techniques
et al. 2019). Beneficial microorganisms are employed in is essential for thoroughly examining the identification,
agriculture as biofertilizers in the context of organic farm- hazard appraisal, and ecological importance of each strain,
ing. Given current regulatory constraints and the selection of instead of relying only on the 16S rRNA gene sequence or
Biologia

Table 2  Commercial biofertilizer products

S.N Country Products Microbes Manufacturer Crop References

1. Argentina Liquid PSA P. aurantiaca Laboratorios BioAgro Wheat Celador-Lera et al.

S.A (2018)
2. Argentina Zadspirillum Azospirillum brasilense Semillera Guasch SRL Maize Celador-Lera et al.
(2018)
3. Argentina Rhizo Liq Bradyrhizobium sp., Rhizobacter Chickpea, Soybean, Adeleke et al. (2019)
Mesorhizobium Common bean, green
ciceri, Rhizobium spp. gram, Groundnut
4. Australia Myco-Tea Azotobacter chroococ- Nutri-Tech solution Tea Adeleke et al. (2019)
cum, B. polymyxa
5. Canada Rhizocell GC Nodulator B. amyloliquefaciens, B Lallen and plant care Beans, maize, carrot, Kenneth et al. (2019)
japonicum BASF rice, cotton
Inc
6. Brazil Bioativo PGPR consortia Embrafros Ltda Beans, maize, Kenneth et al. (2019)
sugarcane, rice, cereals
7. China CBF Bacillus mucilaginosus, China Biofertilizer AG Various cereals Celador-Lera et al.
B. subtilis (2018)
8. Germany FZB 24 fl, BactofilA 10 B. amyloliquefaciens, B. AbiTEP GmbH Vegetables, cereals Kenneth et al. (2019)
megaterium, P
fluorescens
9. India Ajay Azospirillum Azospirillum Ajay Biotech Cereals Celador-Lera et al.
(2018)
10. India Greenmax AgroTech Azotobacter, P. fluore- Biomax Various cereals Adeleke et al. (2019)
Life scens
Biomix, Biodinc, G
max
11. Spain InomixR B. polymyxa, B. subtilis Lab (Labiotech) Cereals Adeleke et al. (2019)
12. United Ammnite A 100 Azotobacter, Bacillus, Cleveland biotech Cucumber, tomato, Adeleke et al. (2019)
Kingdom Rhizobium, pepper
Pseudomonas
13. United Legume Fix Rhizobium sp., B. Legume Technology Common bean, Adeleke et al. (2019)
Kingdom japonicum Soybean
14. United Chickpea Nodulator Mesorhizobium cicero Becker Underwood Chickpea Adeleke et al. (2019)
States
15. United Cowpea Inoculant Rhizobium Becker Underwood Cowpea Adeleke et al. (2019)
States
16. United Quickroots B. amyloliquefaciens Monsanto Wheat, common bean Celador-Lera et al.
States (2018)
17. United Complete Plus Bacillus strains Plant Health Care Various crops Mustafa et al. (2019)
States

other conventional identification approaches (Khan et al. shown to be harmful or to have a negative impact on the
2023b). It is widely accepted that whole genome sequencing environment (Keswani et al. 2019). By implementing the
is a practical and economical method of acquiring thorough environmental and human safety index and adhering to the
phylogenomic data regarding isolates, including taxonomic precautionary principle, we can minimize potential dangers
relationships (Quainoo et al. 2017). Furthermore, determin- associated with the use of beneficial microbial inoculants
ing the safety of new bacterial isolates can be aided by the and ensure environmental and human.
molecular discovery and characterization of genes linked to
virulence (Culligan et al. 2014). The identification of appro-
priate microbes as next-generation bacterial inoculants will Conclusions
be aided by the standardization of methods and knowledge
sharing (Anand et al. 2022a). The environmental and human In agriculture and environments, biofertilizers are a prom-
safety index (EHSI) can be used to classify PGP isolates ising technique since they are an additional, sustainable,
and contrast them with identified microbes that have been and environmentally friendly supply of plant nutrients. It's
 Biologia

critical to recognize the benefits of biofertilizers and apply References

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