Current Research in Environmental Sustainability 3 (2021) 100063

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Current Research in Environmental Sustainability

journal homepage: www.sciencedirect.com/journal/current-research-in-environmental-sustainability

Biofertilizers: A Nexus between soil fertility and crop productivity under

abiotic stress
Aliyu Ahmad Mahmud a, Sudhir K. Upadhyay b, Abhishek K. Srivastava c, Ali Asger Bhojiya a, *
a
Faculty of Agriculture and Veterinary Sciences, Mewar University, Chittaurgarh 312901, Rajasthan, India
b
Department of Environmental Science, V.B.S. Purvanchal University, Jaunpur 222003, UP, India
c
Department of Biotechnology, M.H.P.G. College, Jaunpur 222001, UP, India

A R T I C L E I N F O A B S T R A C T

Keywords: High food demand for the world's teeming population necessitates the intensification of crop production in
Biofertilizers modern agriculture, which requires the extensive use of synthetic fertilizers for higher crop yield. The excessive
Chemical fertilizers use of chemical fertilizers, despite the high nutrients contents and ability to grow crops faster, discovered to be
Crop yield
dangerous to the health and environment besides polluting the groundwater and atmosphere in the future. The
Eco-friendly
Sustainable food security
alternative to these, biofertilizers arose today due to their attributes towards eco-friendly, cost-effective, and easy
to apply in the agricultural field. Biofertilizers are a batch of diverse microorganisms, which can induce plant
growth-promotion activities along with soil health, even under abiotic stress conditions. Biofertilizers maybe
plant growth-promoting rhizobacteria, arbuscular mycorrhizal fungi, and as well as the consortia of other
beneficial microbes. Biofertilizers can sustain plant growth performance, even in a challenging environment. The
performance of perfect-candidate-biofertilizer in the agricultural field depends on crop type, properties of in­
oculants, technical background, and environmental condition. Biofertilizers can, directly or indirectly, help in
attaining food security compared to the harmful effect of chemical fertilizers. A direct mechanism of Bio­
fertilizers refers to phyto-stimulation and nutrient mobility, while an indirect mechanism poses bio-control ac­
tivity. Direct mechanisms involve phytohormone production and phosphate, potassium, zinc, etc. solubilization.
While, indirect-mechanism is HCN production, siderophore production, antibiotic production, etc. The present
review elucidates the diversity of microbial inoculants (biofertilizers), their impacts on agricultural production
through rising soil fertility, and overall crop yield. In line with related literature worked out by different
researchers.

1. Introduction
The sustainability of the agricultural sector is a clef for feeding the
emerging population and economic exports of a country; therefore, the
growth and survival of a nation indirectly depend on its agriculture.
Over the decades, various innovations had elevated by scientists to make
the agricultural sector more efficient (Ajmal et al., 2018). The major
constraint of crop production in the developing world is characterized
by the inaccessibility of the essential plant nutrients due to lack of suf­
ficient quantity and type of fertilizer (Itelima et al., 2018). Fertilizer is
defined in many kinds of literature as any material often applied to the
soils that provide one or more essential nutrients for plant growth and
development (Vanlauwe and Giller, 2006). The use of chemical pesti­
cides and chemical fertilizers virtue of their potential for quick nutrients
release and growing the plants more rapidly and efficiently served their
purpose up to a point (Sneha et al., 2018). However, the continuous use
of chemical fertilizers results in the deterioration of soil quality and
gradual loss of soil fertility, which might further lead to the accumula­
tion of heavy metals in plant tissue, affecting the nutritional contents of
the yield and edibility (Farnia and Hasanpoor, 2015).
Biofertilizer is the microbial inoculants that contain the culture of
dormant or live cells of the effective strains of N-fixing, P-solubilizing/
mobilizing, K-solubilizing (Itelima et al., 2018; de Vives-Peris et al.,
2020; Fasusi et al., 2021). Microorganisms at their cellular level which is
often applied to seeds, soils, or compost material to accelerate the mi­
crobial activities by such organisms through their multiplication and
enhance the nutrient's availability, which can be easily accessible by the
plants (Boraste et al., 2009). The biological activity of microbial

* Corresponding author.
E-mail addresses: aliasger@mewaruniversity.co.in, aliasger786in@yahoo.com (A.A. Bhojiya).

https://doi.org/10.1016/j.crsust.2021.100063
Received 21 November 2020; Received in revised form 28 June 2021; Accepted 30 June 2021
Available online 9 July 2021
2666-0490/© 2021 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license
(http://creativecommons.org/licenses/by-nc-nd/4.0/).
A.A. Mahmud et al.
inoculants helps in mobilizing the availability of nutrients and recovery
of nutrients, thereby improving the soil quality in general (Upadhyay
et al., 2012a; Upadhyay and Singh, 2014; Yadav and Sarkar, 2019).
Biofertilizers have the potential to impede the nitrification process for a
long period while improving the fertility status of the soil (Sun et al.,
2020; Fasusi et al., 2021). They are among the vital constituents of In­
tegrated nutrient management (INM) strategies for meeting both the
soil's productivity and sustainability and at the same time keeping the
environment safe, being pollution-free, economic and source of renew­
able nutrients to the plants to augment synthetic fertilizers in the sus­
tainable production system (Yadav and Sarkar, 2019). Panda (2011)
documented that the impact of bio-fertilizers for yield improvement
ranges between 35% to 65%. However, to potentially exploit the ben­
efits of biofertilizers based on the consistency of their performance.
These require frequent researches in different dimensions due to the
complex biological interaction among the host, other rhizosphere
microflora and fauna as well as the environment (Berg, 2009; Panda,
2011). The use of bio-fertilizers, therefore, supplies and enhance the
productivity per area in a comparatively short time, consume smaller
amounts of energy, reduces contamination of soil and water, increases
soil fertility, and encourages antagonism and biological control of
phytopathogenic organisms (Sneha et al., 2018). This review paper
elucidates the recent updates of potential-biofertilizers in crop produc­
tion due to an increasingly diverse biological activity for soil enrich­
ment, and yields increased per unit area, including abiotic stress
conditions.
2. Biofertilizer and food security
The introduction of ‘Green Revolution’ technologies significantly
Fig. 1. Schematic representation; Influence of biofertilizers on plant growth-performance and soil health. [VAM = Vesicular–arbuscular mycorrhiza, ISP=Increase
soil porosity, ISA = Increase soil aggregation].
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Current Research in Environmental Sustainability 3 (2021) 100063
brings a positive impact on agricultural production initially. However,
excessive use of chemical fertilizers destroyed not only the texture and
other physicochemical properties of the soil (Ye et al., 2020) but are also
harmful to insects, worms, and other microorganisms in soils, which
ultimately reduce both the quantity and quality of production (Khandare
et al., 2020; Jaipunya, 2020). Owing to the current rapid human pop­
ulation there was a huge demand for foods on the one hand, and on the
other hand, farmers aspire for higher profit (Popp et al., 2013). As such,
excessive use of synthetic chemical fertilizers and exploitation of un­
limited underground water were adopted by the farmers (Stefanakis
et al., 2017). A dramatic increase in production and crop yield associ­
ated with these inputs motivated the farmers towards their excessive use
beyond the plants' consumption (Liu et al., 2015). As a consequence, this
resulted in secondary effects on both the plant itself, soil, and environ­
ments like increased atmospheric temperature and unpredictable cli­
matic change (Raza et al., 2019). Besides, the soil organic matter can be
reduced due to soil acidification caused by the excessive use of chemical
fertilizers, which is a threat to plants' growth and development (Naher
et al., 2016). Nevertheless, the issue of food security is related to
balanced fertilization of agricultural soils that sustainably increased
productivity while providing sufficient, safe, and nutritious food
(Stewart and Roberts, 2012). All things being equal, using both organic
and inorganic sources of fertilizers followed by practicing the 4R fer­
tilizer management strategies i.e. (i) right fertilizer source, (ii) right rate
of fertilizer use, (iii) right time and (iv) right target place (Kumar et al.,
2018) will help ensure appropriate nutrients utilization and maximized
productivity (Naher et al., 2016; Gao et al., 2020). Hence, food demand
for the vigorous emerging population can attain sustainably.
A.A. Mahmud et al. Current Research in Environmental Sustainability 3 (2021) 100063

3. Biofertilizers; classification and mechanism of action Table 1

Types of biofertilizers, and its role in crop growth-performance.
Several inoculations and their interaction with crop plants have been Type of Contributions to plant Beneficial Crops References
exploiting for the production of biofertilizers Fig. 1 classified bio­ biofertilizer growth
fertilizers into different groups based on their nature and mode of action Rhizobium Fixes 40-200 kg N/ha. Leguminous crops Panda, 2011;
(Itelima et al., 2018). The microbial inoculants used in crop production Meets up to 80–90% such as; Cowpea, Mazid and
such as nitrogen fixers, phosphorus solubilizers/mobilizers, potassium of N crop Gram, Pea, G/nut, Khan, 2014
mobilizers, PGPR, etc. (Fig.1.) incorporated on a variety of soils and crop requirement. Soybean, Berseem,
Maintains soil Lucerna etc.
types, hence; the knowledge of the variety and abundance of microor­ fertility.
ganisms in biofertilizers is a key to potentially exploits their impact in Increases the yield by
agriculture, including under abiotic stress (Sahu and Brahmaprakash, 10–30%.
2016; Fasusi et al., 2021). The application of biofertilizers is quite sig­ leaves residual N for
the benefit of
nificant for the nutrients enhancements and overall increase in plant
subsequent crops.
production in the sight of INM with the added benefit of growing crops Azotobacter Fixes about 15-20 kg Mustard, Das, 2019
on the least fertile soil (Panda, 2011). N/ha/yr. Sunflower, Grapes,
Based on the functional properties, biofertilizers can classify into Supplies 20-40 mg N/ Sugarcane,
various types such as Nitrogen Fixing Biofertilizer, Phosphate Bio­ ha. Banana, Water
Promotion of growth melon, Papaya,
fertilizers, Biofertilizers for Micro-nutrients, and Plant Growth Promot­ substances such as Coconut,
ing Rhizobacteria, etc. Nitrogen-Fixing biofertilizers raise the nitrogen Indole Acetic Acid, Plantation and
level of soil by fixing atmospheric nitrogen and making it available to Giberellic acid etc. Forest crops
the plants. Examples: Azotobacter, Nostoc, Rhizobium, Azospirillum (Ite­ Results in 10–15%
increase in crop yield.
lima et al., 2018). Phosphate Biofertilizers include phosphorous solu­
Biological control of
bilizing biofertilizers (PSB) and phosphorus mobilizing biofertilizers plant diseases and
(PMB). PSB solubilizes the insoluble phosphate from organic and inor­ destroy plant
ganic phosphate sources. Examples include species of Bacillus, Pseudo­ pathogens.
monas, Penicillium, Aspergillus, etc. (Etesami et al., 2017). PMB transfer Maintains the fertility
status of the soil.
the phosphorus from the soil to the root cortex. Examples: Arbuscular
Azospirillum Fixes N up to 10-20 kg Maize, Rice, Mazid and
Mycorrhiza (AM fungi). Biofertilizers for micro-nutrients include silicate per hectare. Wheat, Finger Khan, 2014;
and zinc solubilizers bacteria. These bacteria degrade silicates and Enhances the intake of Millet, Sorghum, Kumar et al.,
aluminum silicates in soil. Example: Bacillus sp. Plant Growth Promoting both minerals and Bajra, Oil seeds 2017
water.
Rhizobacteria (PGPR) are the bacteria present in the rhizospheric region
Increases roots
(Upadhyay et al., 2019). They promote plant growth by acting as Bio­ developments.
protectants, Biostimulants, and by improving nutrient availability. Improves the
Table 1 highlights the various types of biofertilizers, their contributions vegetative growth and
to plant growth, and the crops that benefitted from the application of yield of crops.
Azolla Fixes up to 900 kg N/ Rice Sahu et al.,
these biofertilizers. Mechanism of action refers to the biological and
ha. 2012
chemical process by which microorganisms contained in biofertilizers Use as green manure
exert their effects applied to the plant's rhizosphere (Vejan et al., 2016). due to their large
The rhizosphere is a thin layer of soil surrounding the roots character­ biomass.
Blue-green algae Fixes almost 20-30 kg Rice, Banana Mishra et al.,
ized by high levels of biochemical activities and comprised of plant,
N/ha in lowland rice 2013; Kumar
bacteria, fungi, and soil constituents (Nihorimbere et al., 2011). More­ field. et al., 2017
over to these zones were found to contain microbial cells of more than Promote the
1010 per gram of root (Egamberdieva et al., 2008), and over 30,000 production of growth
different bacterial species (Mendes et al., 2011). Interestingly, these substances.
Phosphate- Solubilizes 50–60% of Wheat, Potato, Alori et al.,
interactions between plant-root and microbial populations have char­
Solubilizing the fixed P in the soil Rice, Millets, Oils 2017; Mazid
acterized as symbiosis. As the former access the unavailable nutrients Microorganism and 10–20% increase seeds, Pulses and and Khan,
elements into available form through decomposition, the later benefit (PSM) in yield of almost all Vegetables etc. 2014
from root exudates such as carbohydrate, proteins, sugars, vitamins, crops.
Convert unavailable
mucilage, amino acids, and organic acids (Bonfante and Genre, 2010; de
inorganic P present in
Vives-Peris et al., 2020), which modify biochemical properties of the soluble source to the
rhizosphere through the application of these root exudates by acting as a form available to the
messenger between the microbes and the plants (Pii et al., 2015; de plants by the release
Vives-Peris et al., 2020). of organic acids (such
as glutamic acid,
Biofertilizers mediates plant growth performance by direct and in­
oxalic acid, citric
direct mechanisms (Table 2). Direct mechanisms influence the plant acids, tartaric acid,
growth activity directly. It includes nitrogen fixation, phytohormones succinic acid and
production, phosphate solubilization, etc. The indirect mechanism fumaric acid).
doesn't influence plant growth directly, but it protects the plant from the Promote the
production of
deleterious effects of plant pathogens. The production of HCN, anti­ phytohormone.
biotic, siderophore, lytic enzymes (such as chitinases, cellulases, 1,3-glu­ Potassium- Functions in Its applied in Etesami
canases, proteases, and lipases), etc., are the indirect-mechanism of Solubilizing mobilizing almost all crops as et al., 2017
biofertilizers that lyse the cell walls of many pathogenic fungi. Bacteria (KSB) elementary K in to well as in
available form for combination with
crop uptake. other biofertilizers
Enhanced
(continued on next page)

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A.A. Mahmud et al. Current Research in Environmental Sustainability 3 (2021) 100063

Table 1 (continued ) Table 2

Type of Contributions to plant Beneficial Crops References
Mechanism and action of different types of biofertilizer.
biofertilizer growth Biofertilizers Example Mechanisms and References
action
solubilization of fixed
micro nutrients such Nitrogen fixing Rhizobium sp., These groups of Itelima et al.,
as zinc. bacteria Azospirillum sp., organisms worked by 2018
PGPR Increases crop yield Used in all crops Vejan et al., Bacillus sp. and Blue fixing atmospheric N
with the range from 2016 green algae and convert them to
7% to 33%. plant usable forms in
Promotes plant the soil and root
growth through nodules of
increased leguminous plants for
phytohormones and plants uptake.
water& minerals However, These
uptake. biofertilizers are crop
Suppress the growth specific.
of harmful bacteria on Phosphate Aspergillus sp. Worked by Etesami et al.,
the root surface. solubilizing Bacillus sp., and solubilizing insoluble 2017
Micro-nutrients These are strains of Used in all crops in Kumawat biofertilizer Pseudomonas sp. form of phosphate in
Biofertilizer microorganisms helps combination with et al., 2019 (PSB) to soluble form for
in the transformation other type of plant use as most soil
of unavailable biofertilizers P are in insoluble
micronutrients form, this task is
presents in organic accomplish by these
and minerals matter to organisms through
available for plant organic acids
uptake, they includes; secretion which lower
Sulfur solubilizers, the soil pH and
Silicon solubilizers, enhance the
Zinc mobilizers etc. dissolution of
unavailable bound
forms of P thereby
3.1. Direct mechanism making them
available to plants.
Phosphate Mycorrhiza These microorganism Ahemad, 2015
3.1.1. Biological nitrogen fixation
mobilizing works by mobilizing
Biological N2 fixation (BNF) converts atmospheric N2 to plant- biofertilizrers the insoluble P in the
utilizable forms by nitrogen-fixing microbes employing a complicated (PMB) soil by scavenging
enzyme system known as nitrogenase to convert nitrogen to ammonia. phosphates from soil
BNF is a cost-effective and ecologically friendly alternative to chemical layers to which they
are applied. PMB are
fertilizers. Nitrogen-fixing organisms are broadly classified as (a) sym­ broad spectrum bio-
biotic N2 fixing bacteria, which form a symbiosis with leguminous plants fertilizers
(e.g. rhizobia) and non-leguminous trees (e.g. Frankia), and (b) non- Plant growth Pseudomonas sp., Hormones and Souza et al.,
symbiotic (free-living, associative, and endophytes) nitrogen-fixing promoting Agrobacterium, antimetabolites are 2015; Vejan
biofertilizer Achromobacter, produced by these et al., 2016
forms, which include cyanobacteria (Anabaena, Nostoc), Azospirillum,
(PGPB) Xanthomonasetc. group of
Azotobacter, Gluconoacetobacter diazotrophicus, and Azocarus, etc. microorganism
(Bhattacharyya and Jha, 2012; Aasfar et al., 2021; Jain et al., 2021, a). through their action
Upadhyay et al. (2009) amplified nitrogen fixation nif H gene in the which help to
PGPR isolate Arthobacter (SU18). do Carmo Dias et al. (2021) assessed promote the roots
growth, increases the
the genetics and regulation of nitrogen-fixing (nif) genes in Paenibacillus rate of organic matter
brasilensis PB24. BNF regulatory genes, glnK and amtB (encoding GlnK decomposition,
signal transduction protein and AmtB transmembrane protein, respec­ enhance the
tively), and glnR and glnA genes (encoding the transcription factor GlnR mineralization
process and
and glutamine synthetase) were found in the PB24 genome.
availability of plant
nutrients with
3.1.2. Plant hormones production ultimate increase in
The production of plant hormones is the key feature of microbes crop yield. PGPB are
which is used as a biofertilizer. Beneficial microbes produce plant hor­ also crop specific
Potassium Bacillus sp. and These groups of Etesami and
mones such as IAA (indole-3-acetic acid), gibberellins (GA), etc. Solubilizing Aspergillus niger microorganism's Maheshwari,
(Upadhyay et al., 2009, 2012a; Upadhyay and Singh, 2014). IAA acts as Biofertilizer produces organic 2018
a plant growth regulator and is synthesized by using L-tryptophan (Yu (PSB) acids in their course
et al., 2017), IAA mainly regulates plant cell division, cell differentia­ of action which aids
the decomposition of
tion, increases root length, etc. (Santner et al., 2009). GA triggers root
silicates compound
shoot elongation, seed germination, flowering and fruiting in the plant. and cause the
The movement and concentration of GA are essential for the different detachment of metal
developmental stages of plants and useful for crop. Out of 130 known ions thereby making
GAs, few found in plants, fungi, and bacteria, which are bioactive, and them available for
plant uptake. (PSB)
most bioactive GAs reported such as GA1, GA3, GA4, and GA7 (Binen­ are broad spectrum
baum et al., 2018). bio-fertilizers.
Bacillus sp.
(continued on next page)

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A.A. Mahmud et al. Current Research in Environmental Sustainability 3 (2021) 100063

Table 2 (continued ) et al., 2015), and Azospirillum can release organic acid and solubilize
Biofertilizers Example Mechanisms and References zinc in the rhizospheric region.
action

Potassium These groups of Etesami et al.,

3.1.4. Siderophore production
Mobilizing microbes work by 2017 Siderophores are low-molecular-weight iron-binding protein mole­
Biofertilizer mobilizing the cules that aid in the chelation of ferric iron (Fe3+) in the environment
(PMB) unavailable forms of (Saha et al., 2016). Siderophores produced by microorganisms has been
potassium (in the
widely recognized as bio-remediation, bio-controls, bio-sensors, and as a
form of silicates) in
the soil. Similarly, chelation agent. It also takes part in weathering of soil minerals (Ahmed
Some species and Holmström, 2014; Saha et al., 2016). Microbial siderophores deliver
commonly known as Fe to plants when Fe is scarce, allowing them to grow faster. When there
phosphate is a lack of iron, these molecules work as solubilizers for iron from
solubilizers has been
found to play the role
minerals or organic substances. In general, siderophores form 1:1
of (PMB), these complexes with Fe3+, which are subsequently taken up by the cell
include; Bacillus spp. membrane of bacteria, where the Fe3+ is reduced to Fe2+ and released
and Aspergillus spp. into the cell. Iron (Fe) is one of the major minerals found on the earth's
Zinc Solubilizer Serratia sp. These groups of Kour et al.,
surface, however, it is inaccessible to plants in the soil. To deal with the
Biofertilizer microbes primarily 2019
(ZSB) use the acidification fact that iron is typically found in nature in the form of Fe3+, which is
mechanism for very insoluble, PGPR produces siderophores (Upadhyay et al., 2009).
solubilization of Zn. Several bacteria were reported to play the role of siderophore produc­
These microbes tion in plant growth. For example, bacterial siderophores isolated from
produce organic acids
mainly 2-ketoglu­
roots of A. thaliana L., F. rubra, and Agrostis catapillaris L., discovered
conic acid and effective in reducing metal toxicity. Besides phytoremediation and
gluconic acid in soil growth improvement of the plant's growth under contaminated soil
which solubilize Zn. (Grobelak and Hiller, 2017). Inoculation of siderophore-producing
Sulfur oxidizing Thiobacillus sp. Work primarily to Zhi-Hui et al.,
bacterium (Bacillus subtilis CAS15) showed antagonized effects on the
biofertilizer oxidized sulfur in its 2010
(SOB) elemental form to growth of fungal pathogen Fusarium wilt, which also promotes the
sulfates for plant growth of pepper (Yu et al., 2011). Also, the growth and yield attributing
uptake characters of maize were increased due to increased iron transport to
plants as a result of siderophore production by Pseudomonas aeruginosa
(RSP5 and RSP8) strains (Sah et al., 2017).
3.1.3. Nutrient solubilizers
Phosphorus (P) is readily available in soils in both organic and
3.2. Indirect mechanism
inorganic forms (Khan et al., 2009). The bulk of soil phosphorus is
contained in insoluble forms, but plants only absorb soluble forms such
3.2.1. HCN and Ammonia production
as monobasic (H2PO-4) and dibasic (HPO-2 4 ) ions (Bhattacharyya and Jha,
HCN and ammonia production are considered to be indirect plant
2012). Phosphate-solubilizing microorganisms (PSM) have phosphate-
growth promoters. HCN is a volatile substance that has antifungal
solubilizing activity that can offer accessible fertilizers (McKenzie and
properties. Ammonia production can assist the host plant meet its ni­
Roberts, 1990). PSBs are potential biofertilizers among the different
trogen requirements while also reducing pathogen invasion. HCN is
PSMs, due to rapid adaptation in the rhizosphere and organic acid
frequently employed as a biocontrol agent in agriculture systems
production. The production of organic acids is the primary mechanism
because of its high toxicity against phytopathogens. However, HCN is
of inorganic phosphate solubilization (Bhattacharyya and Jha, 2012).
also utilized in the chelation of metal ions and is indirectly implicated in
Song et al. (2021) demonstrated the beneficial effects of PSB on plant
making phosphate available (Rijavec and Lapanje, 2016). Few PGPR
and soil health and observed that PSB produces low molecular weight
produce and synthesize HCN, and acts as biological fertilizers or
organic acids such as oxalic acid, fumaric acid, formic acid, α-ketoglu­
biocontrol agents to boost crop yields (Rijavec and Lapanje, 2016).
taric acid, lactic acid, maleic acid, propanedioic acid, acetic acid, and
Heydari et al. (2008) identified Pseudomonas fluorescence, a cyanogenic
acrylic acid. Bacterial genera like Pseudomonas, Burkholderia, Azoto­
strain which act as a possible biocontrol isolate, increased stem, root
bacter, Enterobacter, Bacillus, Rhizobium, Beijerinckia, Serratia, Micro­
length, germination rate in the rye, wild barley, and wheat. Ammonia
bacterium, Flavobacterium, and Erwinia are reported as the most
production also plays an important role in plant growth by the accu­
significant phosphate-solubilizing bacteria (Upadhyay et al., 2009;
mulation of nitrogen and helps in promoting root and shoot growth and
Etesami et al., 2017; Alori et al., 2017; Bhojiya et al., 2021).
biomass production (Dutta and Thakur, 2017). Rodrigues et al. (2016)
Potassium solubilizing bacteria (KSB) have been shown to solubilize
observed the production of ammonia in 45% of the endophytic bacterial
K-bearing minerals and convert insoluble K to soluble forms that plants
samples from leaf, stem, root and rhizosphere of the commercial sug­
may absorb. Acidothiobacillus ferrooxidans, Paenibacillus spp., Bacillus
arcane variety-RB 867515.
licheniformis, and Burkholderia cenocepacia, Klebsiella variicola, Entero­
bacter cloacae, Bacillus cereus and Bacillus circulans releases organic acids
3.2.2. Chitinase production
which can solubilize K minerals (e.g., biotite, feldspar, illite, muscovite,
The production of chitinase by microbes has been considered as one
orthoclase, and mica) (Zhang and Kong, 2014; Ahmad et al., 2021).
of the strategies for controlling plant pathogens. Chitinase induces the
Plants may take up zinc as a divalent cation, but only a small per­
breakdown of the cell wall, which affects the structure's integrity and
centage of total zinc is soluble in soil (Kabata-Pendias and Pendias,
hence inhibits pathogen development. Chitinase attacks chitin (1,4-N-
2001). The remaining zinc is found as insoluble compounds and min­
acetyl-glucosamine), which is an essential component of the fungal cell
erals (Kour et al., 2019). Zinc deficiency arises as a result of zinc
wall (Sadfi et al., 2001). Chitinolytic activities found in B. thuringiensis,
shortage in the soil, and it is one of the most common micronutrient
B. licheniformis, B. circulans, B. cereus, and B. subtilis (Blake et al., 2021;
deficiencies. Pseudomonas aeruginosa HMR1, a PGPR which showed zinc
Kwon et al., 2021), and gram negative bacteria such as P. fluorescens,
solubilizing activity (Bhojiya et al., 2021). Rhizobium strains (Naz et al.,
Enterobacter agglomerans, Serratia marcescens, and Pseudomonas aerugi­
2016), Bacillus aryabhattai (Ramesh et al., 2014), Bacillus sp. (Hussain
nosa (Pramanik et al., 2021; Mendez-Santiago et al., 2021).

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A.A. Mahmud et al.
3.2.3. Antibiotic production
PGPM strains serve as an inhibitory factor for a variety of pathogenic
microbes by producing metabolites that might lead to pathogen growth
suspension at the minute, the level has opened up new possibilities.
Today, microorganisms' antagonistic properties are being investigated
as a possible replacement for chemical pesticides, which have had
devastating consequences for our ecosystem (Vurukonda et al., 2018).
Bacteria suppresses phytopathogens in a variety of ways, including
competing for available nutrients and space, producing bacteriocins,
lytic enzymes, and antibiotics (Lugtenberg and Kamilova, 2009). The
production of one or more antibiotics is the mechanism most commonly
associated with the ability of plant growth-promoting bacteria to act as
antagonistic agents against phytopathogens (Glick et al., 2007). For
example, antibiotics such as 2,4-Diacetyl phloroglucinol, Phenazine-1-
carboxylic acid, Phenazine-1- carboxamide, Pyoluteorin, Pyrrolnitrin,
Oomycin A, Viscosinamide, Butyrolactones, Kanosamine, Zwittermycin-
A produced by Pseudomonas fluorescens and P. aeruginosa. Bacillus subtilis
and B. amyloliquefaciens produced antibiotic such as Subtilin, Subtilosin
A, Tasa, Sublancin Bacilysin, Chlorotetain, Mycobacillin, Rhizocticins,
Bacillaene.
3.2.4. Biocontrolling activity
In an attempt to substitute current approaches of extensive fungi­
cides usage, often associated with plant pathogen-resistance develop­
ment in a long run, there is increasing concern on the use of microbial
antagonists to control disease infection triggered by soil-borne and air-
borne pathogenic bacterial and fungal microbes. The indirect way by
which rhizobacteria promote plant development is through serving as
biocontrol agents (Glick, 2012). Biocontrol microorganisms emerged in
the agricultural field as a safe alternative to chemical pesticides (Vur­
ukonda et al., 2018). The main mechanisms of biocontrol action in PGPR
include competition for nutrients, niche exclusion, induced systemic
resistance, and the production of antibiotic compounds or enzymes
capable of fungal cell lysis (Lugtenberg and Kamilova, 2009). Different
microorganism serves the role of bio-control agents, the most common
biocontrol agents currently used belong to the group of plant growth-
promoting rhizobacteria (PGPR) (Pii et al., 2015; Paterson et al.,
2017). A limited study revealed that root-associated microbes, through
their positive interaction with the plant root, might influence plant
physiological changes under pathogen attack, thereby increasing the
resistivity of the plants, particularly aboveground parts by phytopha­
gous insect infection (Pangesti et al., 2013), through the production of
compounds such as ethylene, salicylic, or jasmonic acids, plant defense
is then guaranteed. Alternatively, volatile organic compounds (VOCs)
that have insecticidal properties might be produced by beneficial mi­
crobes, such as root-colonizing pseudomonads, which directly act
against plant-feeding insects (Kupferschmied et al., 2013). In several
literatures, the biocontrol activity against many fungal diseases due to
antagonistic relationships between beneficial microbes and pathogens
have been explained (Baker, 1991; Trivedi et al., 2017). For example,
the genus Streptomyces has been extensively studied as a biocontrol
agent of soil-borne fungal pathogens through the production of various
antifungal metabolites because of its intense antagonistic activity (El-
Tarabily and Sivasithamparam, 2006; Himaman et al., 2016; Vurukonda
et al., 2018; Dukare et al., 2019). The genus Pseudomonas was also
successful in preventing fusarium pathogen infection by seeds inocula­
tion of different crop species (rice, Bermuda grass and annual bluegrass
seeds) through the production of growth inhibition compounds (Verma
et al., 2018). Moreover, yeast is a pathogen that often colonized the
wounded fruits shortly especially under the favorable condition in a
nutrient-rich environment, the use of bio-control agents have been
successfully reported to confer pathogen invasion by colonizing yeast in
wound protection (Spadaro and Droby, 2016; Ghorbanpour et al.,
2018).
6
Current Research in Environmental Sustainability 3 (2021) 100063
3.2.5. Induced systematic resistance (ISR)
ISR is a phenomenon triggered by a certain group of PGPMs such as
PGPRs when the defense mechanisms of the plant are stimulated and
primed to repel pathogenic infections (Borriss, 2011). Under varying
degrees of plant disease by plant pathogens such as bacterial, fungal,
viral, nematode, and oomycete, ISR is known to reduce disease severity
on diverse plant species (Walters et al., 2013). Plant resistance to
harmful bacteria, fungi, and viruses can be induced through the inter­
action of certain rhizobacteria with plant roots. Furthermore, jasmonate
and ethylene signaling within the plant are involved in ISR, and these
hormones promote the host plant's defense responses against several
plant pathogens (Glick, 2012). Lipopolysaccharides, flagella, side­
rophores, cyclic lipopeptides, 2,4-diacetylphloroglucinol, homoserine
lactones, and volatiles like acetoin and 2,3-butanediol are just a few of
the bacterial components that induce ISR (Lugtenberg and Kamilova,
2009). Recent studies showed that microorganisms like Bacillus atro­
phaeus, Bacillus subtilis, Trichoderma harzianum, Trichoderma asperellum
and Paenibacillus dendritiformis trigger ISR in Tomato and Chilli Pepper
against Meloidogyne incognita, Botrytis cinerea and Colletotrichum trun­
catum (Ayaz et al., 2021; Yadav et al., 2021; Zhou et al., 2021). Many
scientists have studied the systemic disease protection in both green­
house and field experiments on Cercospora leaf spot and Erwinia tra­
cheiphila disease respectively (Zehnder et al., 2001; Bargabus et al.,
2004; Raymaekers et al., 2020), which involves the use of both Gram-
negative and Gram-positive strain of PGPR (Compant et al., 2005). For
example, the use of biocontrol agents Trichoderma spp. through myco­
parasitism phenomenon and production of antibiotics observed to sup­
pressed the severity of the disease, particularly disease caused by soil-
borne plant pathogens (Akhter et al., 2015). Nevertheless, the myco­
parasitic ability of T. virens mutants deficient did not affect the biological
activity of these strains. Instead, a very strong correlation between the
terpenoid phytoalexin defense and control of Rhizoctonia solani exists in
cotton seedlings which are triggered by this strain (Howell et al., 2000).
Further, a greenhouse experiment conducted by Pourya et al. (2020)
reported that wheat seeds treated with biofertilizers (Biofarm and pro­
bio 96) could be environmentally safe to manage insect pest Sitobiona­
venae as a result of induced host-plants resistance. The genus Penicillium
spp. have also been extensively tested for their ability to impact ISR in
plants and were discovered very much effective against plant pathogenic
bacterial, fungal, and viral disease attack (Elsharkawy et al., 2012;
Hossain et al., 2014; Hossain and Sultana, 2015).
4. Biofertilizers for the alleviation of abiotic stress
Abiotic stress has been adversely affecting the growth and produc­
tivity of crops, especially those raised under field conditions (Gill and
Tuteja, 2010; Upadhyay and Singh, 2014; Upadhyay et al., 2019). This
situation is becoming aggravated with changing climate and improper
agricultural practices such as excessive use of chemical fertilizers and
pesticides beyond recommended crops and soil's threshold level (Liu
et al., 2017). Consequently, this results in a significant reduction in
agricultural productivity and the degraded ecosystem (Begum et al.,
2019). Thus, there is an urgent need for the development of new man­
agement techniques that are eco-friendly to both soil and its surrounding
environment. Several forms of biofertilizers like plant growth-
promoting rhizobacteria (PGPR), arbuscular mycorrhizal fungi (AMF),
mycorrhizal helping bacteria (MHB), as well as consortia of other
beneficial microbes, has been successfully applied to sustained plants
growth and developments in a challenging environment (Mazid and
Khan, 2014; Odoh et al., 2020; Paul et al., 2020; Basu et al., 2021; Fasusi
et al., 2021). Biofertilizers induce nutrient uptake in the plant by raising
nutrient mobility in the rhizospheric region (Gou et al., 2020). Bio­
fertilizers releases organic acids, enzymes and signaling compounds, etc.
(de Vives-Peris et al., 2020), which participate in the different nutrient
cycling and maintain beneficial microbial population in the root sur­
roundings (Table 2). Raza et al. (2019) reported that different abiotic
A.A. Mahmud et al.
stress, like climate-related (e.g. Drought, Flooding, Heat, etc.) and soil-
related (Salt, Fertility, Heavy metals, etc.) has been scientifically re­
ported to interfering with normal plant growth.
4.1. Biofertilizers under drought stress
Drought is abiotic stress associated with a shortage of water to meet
plants' requirements. It is said to occur when the rates of water uptake by
the plants are far below the rate of water loss from the plant's surface
through transpiration (Changhai et al., 2010). Consequently, normal
plant growth and development become hampered due to the reduction
of water content in their relative cells (Utsumi et al., 2019). Several
earlier reports indicate the challenge of drought stress and its related
consequences on the breeding of crops and yield (De Oliveira et al.,
2017). Subsequently, exploration of the microbial potentials in the soils
and their interactions offers a significant role in decreasing the rising
jeopardy of drought conditions (Igiehon and Babalola, 2018). Farooq
et al. (2009) documented that, synthesis of osmolytes enzymes increased
as an adaptation mechanism by the plants when exposed to this condi­
tion of water deficits. Also, the production of phytohormones or syn­
thesis of molecules such as IAA, gibberellins, cytokinins, and abscisic
acid is enhancing by PGPR (Upadhyay et al., 2009, 2011, 2016; Kenneth
et al., 2019). Hence, an increase in root length, growth, total number,
surface area, as well as the formation of root hairs and lateral roots for
increased absorption of water are among the changes plants experienced
(Raza et al., 2019).
4.2. Biofertilizers under heavy metals toxicity
Since the beginning of the industrial revolution in the year 18th
century, anthropogenic pollution of soils play a brutal impact on the
public health sector, the global economy, and environmental conser­
vation (White et al., 1997). Most importantly, the release of inorganic
pollutants into the environment (such as heavy metals or radioactive
isotopes) becomes a wide area of concern in the agricultural sector
owing to their deleterious effects on food crop production (Odoh et al.,
2020). Hydrocarbon products and their derivatives/congeners often
accumulate and concentrate with the aids of biological agents in the
food chain through the soil ecosystem (Nwankwegu et al., 2018). Some
of the heavy metals found in the soils such as lead, copper, nickel, zinc,
mercury, cadmium, chromium, arsenic, cobalt, etc., due to their non-
degradability, bio-accumulate in the soil environment over a long
period (Popp et al., 2013; Chibuike and Obiora, 2014), as a result,
ruining food crop production with health risk to both animals and
human. In agriculture, a decrease in crop yield and further economic loss
are the detrimental effects of these heavy metal pollutants (Chibuike and
Obiora, 2014). Phyto-remediation is a natural mechanism developed by
a few plants to degrade, stabilize, volatile, or mitigate pollutants from
their surrounding environment in an attempt to clean up contaminated
soils (Ahemad, 2015). Nonetheless, the phytoremediation potentials of
most plant species are reducing because of the toxic effects of heavy
metals on the plants (Glick, 2003). White (2001) and Glick (2003) re­
ported that phytoremediation enhancing through the incorporation of
bacteria into the soil, commonly described as “microbe-assisted phy­
toremediation”. A few groups of microorganisms play a critical role in
bioremediation technology, includes PGPR, AMF, phosphorus solubi­
lizing bacteria, and MHB (Ahemad, 2015). Previous researchers isolated
heavy metal tolerant bacteria belonging to the genus Pseudomonas sp.
and Serratia sp. with various plant growth-promoting activities such as
the production of plant growth-promoting hormone (IAA), ammonia,
HCN, siderophores, and phosphate solubilization (Bhojiya and Joshi,
2016; Jain et al., 2020; Bhojiya et al., 2021). Secretion of proteins, an­
tibiotics, organic acids, and other chemical molecules are some of the
strategies developed by these plants to curb the menace of pollutants in
their environment (Wei et al., 2017). White et al. (1997) and Ahmad
et al. (2018) reported that the coexistence of microorganisms in the soil's
7
Current Research in Environmental Sustainability 3 (2021) 100063
rhizosphere zone like bacteria, fungi, etc., can serve as active metal
sequestering in metal-stressed soils as well as growth-promoting bio-
inoculants for plants.
4.3. Biofertilizers under heat and cold stress
Extremes temperature acts as a stress condition for plant growth and
development (Akter and Islam, 2017). It implies a rise or fall in a tem­
perature at a specific time, more than the critical edge (Hasanuzzaman
et al., 2013). Heat stress is associated with several adverse effects to
plant growth ranging from loss of microbial diversity, nutrients loss, and
soil fertility to changes concerning plants' physiological, morphological,
biochemical, and molecular composition (Chodak et al., 2015). This
abiotic stress in the tropical regions where the temperature is relatively
higher has impacted agricultural developments in most communities
with consequent weather change and variation in the farming calendar
(Odoh et al., 2020). Besides, during high-temperature stress, cellular
changes like reactive oxygen species may occur, which is a result of the
change in plant metabolism, build the worst scenario is the reduction in
crop productivity (Hasanuzzaman et al., 2013). Reduction in plant water
use efficiency, cell turgidity, seedling growth, and seed germination are
all attributed due to heat stress (Akter and Islam, 2017). Similarly, this
stress condition plays a role in the enhancement of leaf senescence,
cellular function disturbances, photosynthetic enzyme deactivation, and
damages to chloroplast through the provision of oxidative stress con­
dition (Sharma et al., 2012).
On the other hand, the effect of low temperature (cold and frost)
lessened the free movement of biomolecules (kinetic energy) cause
decreased enzymatic reactions and cell membrane flexibility. These
resulted in photosynthesis evasion, cell division, imbalance of water
transport, and variation in crop growth and development in the arctic
(cold) environment (Odoh et al., 2020). Consequently, Lamaoui et al.
(2018) documented that the least impact of these abiotic stress on the
smooth growth and development of the plants as reported by many re­
searchers, and at all states is the tendency to lower crop yields by over
50%. Plants over the year, evolved a coping mechanism to avoid this
stress condition, have developed and induced accumulation of
numerous hydrophilic and osmolytes proteins like dehydrin. Chakra­
borty et al. (2018) and Lamaoui et al. (2018), based on their research
compiled that, several beneficial bacteria like thermophiles and cry­
ophiles proved to have the ability to induce stability in plants during
extreme temperature. However, this mechanism depends on several
factors such as; type of stress, species of microbes, plant's genotype, and
most importantly, the relationship between plant and microbes in the
soil. Abd El-Daim et al. (2014) investigated the role of rhizospheric
bacteria in the management of heat stress in Wheat (Triticum aestivum).
They treated the seeds of wheat (cultivars Olivin and Sids1) with Bacillus
amyloliquefaciens UCMB5113 or Azospirillum brasilense NO40 and found
that treated seeds showed tolerance to heat as compared to control.
Khan et al. (2020) isolated and studied the role of thermotolerant
B. cereus SA1 for heat tolerance in the soybean plant. They suggested
that B. cereus SA1 could be used for the mitigation of damage in soybean
plants due to heat stress. Mukhtar et al. (2020b) determined the po­
tential of thermotolerant PGPR, Bacillus cereus in mitigating the heat
stress effects in tomato plants. They concluded that this PGPR bacterial
strain proved as a potential candidate for improving tomato crop growth
under heat-stressed conditions. Zubair et al. (2019) designed the study
to alleviate cold stress on the wheat plant with the help of psychrophilic
PGPR bacteria belonging to the Bacillus genera. They found that cold-
tolerant Bacillus strains help in inducing stress response by regulating
abscisic acid, lipid peroxidation, and proline accumulation pathways.
4.4. Biofertilizers under salinity stress
The saline soil is increasingly getting global attention in the arid and
semi-arid regions (Mongi et al., 2010). High salts concentration in
A.A. Mahmud et al.
agricultural soil affects plant growth by impairing their physiological
mechanisms due to the toxicity of cells by excess Na+ and Cl− ions (Guo
et al., 2019). Disruption of photosynthetic efficiency, stomatal conduc­
tance, membrane organization, gas exchange, water statue as well as
disorganization of enzyme structure (Huang et al., 2015), and other
macromolecules among other are the noxious effects of these stress
condition. Besides, other metabolic activities, plants' cell division,
growth, development, and productivity may be directly or indirectly
affected (Jouyban, 2012). As a natural adaptation against salt stress
conditions, several strategies adopted by plants like its association with
soil fungi described as AMF (Odoh et al., 2020) and salinity tolerant
PGPR (Upadhyay et al., 2012b). Generally, a high level of salts (e.g NaCl,
Na2SO4, Na2CO3, or magnesium-based salts) in the rhizosphere of arid
and semi-arid soils affecting physiological processes in plants classified
into three basic types of stress, namely; oxidative, osmotic, and ionic
(Saxena et al., 2017). Oxidative stress inhibits cell growth and meta­
bolism in plants by stimulating the release of oxygen (Gill and Tuteja,
2010). While osmotic stress induces natural drought conditions, thereby
reducing water use efficiency because of altered water potential. Ionic
stresses, on the other hand, interrupt the ionic balance in the whole plant
system (Saxena et al., 2017). The PGPR has been exploited efficiently in
several kinds of researches to enhanced plant growth and development
under high saline conditions (Upadhyay et al., 2009, 2011, 2019). It is
accomplished either directly by activities of microbes or indirectly due
to the established relationship between the plants and the microbes
(Upadhyay et al., 2016). According to Ansari et al. (2013), the existence
of endophytic fungi in the plant roots like AMF and Piriformospora indica
efficiently induce host defense as a response to salt stress, thus,
ameliorating ions toxicity. Guo et al. (2019) postulated that the accu­
mulation of osmolytes and other low molecular weight compounds like
amino acids increased because of the presence of PGPR, which elicits
both the physiological and developmental functions of the plants. Sta­
bilizing the plant growth under unfavorable environmental conditions
through the inflection of osmotic pressure in both the cytoplasm and cell
membrane is possible in the presence of the enzyme and osmolytes
(Cherel and Gaillard, 2019). Mukhtar et al. (2020a) evaluated the plant
growth-promoting abilities of bacterial strains and their effect on maize
growth under salinity stress. All the bacterial strains (Bacillus safensis
HL1HP11 and Bacillus pumilus HL3RS14, Kocuria rosea HL1RP8,
Enterobacter aerogenes AT1HP4, and Aeromonas veronii AT1RP10 used as
inoculants in the study positively affected the maize growth under saline
stress. El Semary et al. (2020) applied microbial inoculums of cyano­
bacterial strain Cyanothece sp. and the rhizobacterium Enterobacter
cloacae, alone or in combination with one another to alleviate the
salinity stress in plants. They found that the application of these mi­
crobes as biofertilizers at the highest salinity level promotes the adverse
effect of salinity mitigation to a greater extent. In another study, Phour
and Sindhu (2020) applied Pseudomonas argentinensis and
P. azotoformans to mitigate the saline stress in Indian Mustard (Brassica
juncea L.). They concluded that this Pseudomonas sp. enhance the pro­
ductivity in mustard crop under salinity stress and could be used as a
biofertilizer.
5. Biofertilizers and soil productivity
Over half of the century, the world's attention focussed on the effort
to increased food crop production to cater to the challenges of the
increasing food demands (Strassburg et al., 2014). In recent years,
several means aim at enhanced plant growth have been developed
(Sneha et al., 2018). The PGPR referred to as “Bio-fertilizer” emerged to
be one of the organic fertilizers (Mishra et al., 2013; Basu et al., 2021).
These biofertilizers often applied to the soil or used as seed treatments
nourish the plant through the indirect supply of the essential nutrients to
the plant for growth and metabolism (Upadhyay et al., 2012a; Mishra
et al., 2013; Basu et al., 2021). The process of plant nourishment has
accomplished when all the essential plant nutrients are accessible to the
8
Current Research in Environmental Sustainability 3 (2021) 100063
plant through the biological activities of the microbes.
The perfect biofertilizers can assemble their population in the rhi­
zospheric region of the selective plant by quorum sensing molecules
(Reinhold-Hurek et al., 2015), these microbial assemblies benefit the
plant's growth for a long time through nutrient cycling and raising soil
fertility. Biofertilizers are not fertilizers themselves but potentially
enhance the release of naturally reserved nutrients in the soil (Schutz
et al., 2018). The type of fertilizer used for increased crop production
depends on the materials used to supply plant nutrients as fertilizers. In
general, any fertilizer material used should contain an adequate and
balanced quantity of nutrients for optimum plant growth. Since the
amount of nutrients released is small compared to the total applied each
year from the organic source, the remaining parts are actively released
by the process of chemical and biological activity. Thus, biofertilizers
are employed to enhance the conversion of unavailable nutrients present
in the soil to available form for plant absorption (Verma et al., 2017), as
shown in Fig.1.
6. Methods of biofertilizers application
Different techniques have use for biofertilizer application takes into
consideration such as type of crops, properties of inoculants, constraints
related to farmers, technical background, and environmental conditions
(Mahmood et al., 2016). Each technique has advantages and disadvan­
tages. Besides, Muraleedharan et al. (2010) documented certain pre­
cautions that need to observe before application, which includes; storing
the biofertilizer products at optimum temperature, neither below 0 ◦ C
nor above 35 ◦ C, avoidance of keeping used solution overnight, and
direct exposure to sunlight. The ultimate aim is to enhance the avail­
ability of plant nutrients, often incorporating the microorganisms near
the surface of the root zone. The different techniques based on dry and
liquid formulations of biofertilizers, depicted in Table 3.
7. Biofertilizers role on crop production
The soil fertility and crop productivity synonymously, though the
two terms significantly differ as the former described the inherent ability
of the soil to produce essential plant nutrients in sufficient quantity
(Panda, 2011; Basu et al., 2021). However, both the two terms depend
on the available essentials plant nutrients such as N, P, K, Ca, Mg, S, Cu,
Cl, Si, etc., which produced from the natural process of organic matter
decomposition besides the addition through chemical fertilizers. Mi­
crobial inoculants purposely enhance the decomposition process for the
release of those essential nutrients and induce overall crop productivity
(Pii et al., 2015). At present, the world population is estimated at 8
billion, targeted production of 321 million tons to feed the teeming
population (Conway, 2012). To meet up with the current and future
demands, the solitary use of chemical fertilizers is no longer a lasting
solution due to their economic and negative impacts on soil health. As
such, the use of biofertilizers assumes special attention for their eco­
nomic, eco-friendly, and sustainability in nature, besides improving the
soil's physical, chemical, biological, and in turn, crop yield per unit area
(Kumar et al., 2017). Additionally, biofertilizers are promising in diverse
extreme conditions such as heat, salts, drought, pH, and other distur­
bances from the environment such as crude protein pollutants as well as
heavy metals (Odoh et al., 2020). The altering of the microbial popu­
lation around the plant's rhizosphere, thereby distressing the biological
activities of the soils' ecosystem (Vejan et al., 2016). Effects of bio­
fertilizers on various crops yield at different locations, depicted in
Table 4.
Furthermore, biofertilizer application enhances the function and
structure of soil microorganisms (Fig.1.) in addition to its influence on
the physicochemical properties of the soil (Berg, 2009). The impacts of
PGPR vary significantly upon introduction into the soil's rhizosphere
among the indigenous populations. Some microbial populations may
remain unchanged, and others may be altered, either in a positive way
A.A. Mahmud et al.
Table 3
Method of biofertilizers application based on dry and liquid formulation.
Method of Dry formulation
application
Seed 200 g of biofertilizer is suspended in 300–400 mL of water and
treatment mixed gently with 10 kg of seeds using an adhesive like gum
acacia, jiggery solution, etc. The seeds are then spread on a clean
sheet/cloth under shade to dry and used immediately for sowing.
Seedlings This method is used for transplanted crops. For rice crop, a bed is
made in the field and filled with water. Recommended
biofertilizers are mixed in this water and the roots of seedlings are
dipped for 8–10 h and transplanted
Soil 4 kg each of the recommended biofertilizers is mixed in 200 kg of
application compost and kept overnight. Its then mixed with the soil the
following day before seed sowing/planting.
Standing crop A mixture of biofertilizer incorporated in to the soil after pruning
operation then followed by irrigation. The quantity depends on
the size of the plant; the overall aim is to ensure enough
inoculation around the root zone to enhanced biological activity.
Foliar Application of prepared-biofertilizers at two stages; firstly, the
application mixture sprinkled when the crop had two-three formed leaves.
And the second stage, at nine-ten leaves-formed. However, the
second stage is particular for maize plants, and for wheat, the
second stage applies after tillering of the plant.
by enhancing their growth, or negatively through inhibiting their
growth (Mendes et al., 2011). The bacterial population in the chlor­
othalonil contaminated rhizosphere of Vicia faba was increased when
treated with Stenotrophomonas acidaminiphila BJ1 probiotic strain
(Zhang et al., 2017). Nonetheless, many researchers characterized the
impacts of microbial inoculation on soil microorganisms (Etesami et al.,
2017). Upadhyay et al. (2012a) observed the positive effects on growth
and antioxidant status of Wheat under saline conditions when inocu­
lated with two PGPR strains, Bacillus subtilis SU47, and Arthrobacter sp.
SU18. Gangwar et al. (2013) observed the increase in root length, shoot
length, root dry weight, and shoot dry weight of Mung Bean (Vigna
radiata L.) in response to dual inoculation of plant growth-promoting
Pseudomonas putida and Trichoderma viride.
8. Application trends of biofertilizers vs. chemical fertilizers
Biofertilizers are made up of selected living microbial cells that
provide nutrients to plants via their root system. The microbes in these
fertilizers make use of various mechanisms to provide the plants with
nutrients. They can fix nitrogen, solubilize phosphates, mobilize phos­
phates, etc. Further, the excessive usage of chemical fertilizer in agri­
cultural fields ended up causing the degradation of the soil and polluting
the environment. Due to these beneficial qualities of biofertilizers, the
global market size of biofertilizers is estimated at $1.49 billion in 2019
and is expecting to hit $3.28 billion by 2027 (Biofertilizers market
research report, 2020). It is anticipated that the Asia-Pacific region will
have 34% of the total demand, while Europe and Latin America will also
move their consumption to these biofertilizers due to the regulation of
chemical fertilizers (Ajmal et al., 2018). Globally, various researchers
designed studies that show the reduction in the usage of chemical fer­
tilizers and encourage the application of biofertilizers (Table 5).
9. Limitations to the use of biofertilizers in crop production
In general terms, the use of biofertilizers increases the sustainability
of agricultural production with few side effects to the environment
particularly, the perceived smell of the fresh inoculants. Similarly, there
are certain limitations associated with the use of biofertilizers in agri­
cultural sectors.
Different sort of constraints has been documented in the emerging
field of biofertilizers from the points of production to the last points of
practical field application by farmers, these limitations are; technical
Current Research in Environmental Sustainability 3 (2021) 100063
Liquid formulation References
3-5 mL and 250 mL of liquid culture is recommended per kg and Malusa et al., 2012;
one acre respectively, with minimum colony forming unit (Cfu = Sabalpara and
1 × 108). However, this depend on the size of seeds used, larger Mahatma, 2016
seeds generally require high amount compared to small seed crops
1% jaggery solution is prepared firstly by mixing 200 g jaggery in Sabalpara and
20 L of water, liquid biofertilizer is then mixed with the jaggery Mahatma, 2016; Kumar
solution were the seedlings is dipped for 30 min before planting et al., 2017
1 L/ha of liquid biofertilizers is recommended, 200 kg/ha of FYM Malusa et al., 2012
or pulverised soil can be applied near the root zone prior to
irrigation or incorporated and applied as fertigation were drip
system of irrigation is employed
Generally, 50 mL of biofertilizer is recommended per plant. This is Malusa et al., 2016;
also varied with the growth stage of the plant Kumar et al., 2017
300m3 of diluted liquid bacterial inoculum is enough for per m2. Habibzadeh et al.
The mixture is applied using a plastic haversack sprinkler (2012); Latkovic et al.
(2020)
problems such as; lack of good quality carrier materials, inadequate
qualified technical personnel in the production units and low level of
awareness and adoption to the farmers due to different inoculation
method and long-time effects on the crop compared to chemical fertil­
izers (Bhardwaj et al., 2014; Odoh et al., 2020). One of the major-
limitations of biofertilizer is that it is difficult to store for long time i.
e. they have a short shelf life. Shelf life can be increases by thermo-
tolerant, drought-tolerant, genetically modified strains, and liquid bio­
fertilizer formulation (Brar et al., 2012). The lack of region-specific
indigenous microbial strains is another major limitation. Biofertilizers
are not only specific to particular crops but are also soil specific. Mi­
crobial strains of one agroclimatic region cannot perform well to their
potential as biofertilizers in the other agroclimatic region. The probable
reason could be a lack of competitive ability and survivability over the
local microbes. Infrastructures constraints such as; inaccessibility of
essential application equipment like sprayers, shortage of power supply,
and poor road network for the conveyance of biofertilizers to the field
(Ajmal et al., 2018; Odoh et al., 2020) and marketing constraints such
as; problems due to 3R's of fertilizer management i.e. right inoculants at
the right place and at the right time and inadequate retail outlets of the
producers) (Backer et al., 2018). However, all these limitations affect the
production, marketing, and usage of biofertilizers in agricultural crop
production (Kumar et al., 2017; Malusa et al., 2016).
10. Conclusion and remarkable perspective
Biofertilizers have been used to boost up crop production by aug­
menting the plant's available nutrients through the organic matter
decomposition process. Two main reasons-necessitate the use of bio­
fertilizers in today's crop production. The first is to increase the use of
biofertilizers, which results in the corresponding increase in crop yield,
and the second is the long-term use of synthetic fertilizers degrades the
soil besides other threats to our health and environment. The efficacy of
biofertilizers can enhance by sound knowledge and long-time practical
experience on a diverse soil type. The incorporation of biofertilizers in
crop production is still not well understood by our farmers despite its
low cost and environmental safety. Multiple mechanisms exist for the
action of biofertilizers to improve plant growth. In nature, few micro­
organisms are present which possess all the PGP mechanisms against the
adverse environment, and thus genetic modification is the best option to
improve the microbial strain. Several types of research need to be con­
ducted on different farming communities to steepen with the
9
A.A. Mahmud et al. Current Research in Environmental Sustainability 3 (2021) 100063

Table 4 Table 5
Biofertilizers performance at selected locations. Impact of biofertilizers in reduction the consumption of chemical fertilizers.
Location Biofertilizer Crop grown Effect References Country Biofertilizers Rate of Crop Reference
reduction in
Xinjiang P. putida Rs-198 Vitis The weight, Lu et al.,
the use of
Province, liquid vinifera L. length, and 2020
chemical
China biofertilizer width increases
fertilizer (%)
(Rs198LBF) 17.2%, 6.2%,
and 4.4% India Azotobacter and PSB 25 Wheat Khandare
respectively, in (Triticum et al., 2020
the addition of aestivum
60 mL of L.)
Rs198LBF (Rs- Azospirillum, 25 Brinjal Padhiary
198) Azotobacter, PSB and Azad B-3 and Dubey,
Yinchuan, Trichoderma Tomato Yield increase Ye et al., (Vesicular Arbuscular 2020
Ningxia harzianum T7 from82.85 to 2020 Mycorrhiza)
Province, 86.78 kg/plot Iran Azotobacter vinlandi 50 Basil Dehsheikh
China and PSB included (Ocimum et al., 2020
Sulaymania, Azoto barwar1 Cucumis Fruit Yield per Saeed Pseudomonas putida sp.)
Iraq sativus green house/kg et al., 2015 and Pantoea
increase to agglomerans
4340.6 as Egypt Azotobacter 50 Maize Gao et al.,
control chroococcum SARS 101 (Zea mays 2020
(4298.3). and Bacillus circulans L.)
Lamjung Azotobacter Gautam 63.11% yield Mahato & ARC-SWERI- 2
Nepal variety of increases over Kafle, Thailand 50 Rice Jaipunya,
wheat the control 2018 Biofertilizers (San-Pah- 2020
Egypt Azotobacter (Zea mays Cob Weight (g), Gao et al., Tawng 1
chrocoocum, L.) Cob Length 2020 sticky)
arbuscular (cm), Row Malaysia Consortium of 50 Rice Naher et al.,
mycorrhizal Number/ nitrogen fixing 2016
fungi (AMF), Cob100 and bacteria (Bacillus sp.
Bacillus Grain Weight Sb13, Burkholderia sp.
circulans (g) were Sb16) and phospahte
367.2%, solubiling bacteria
39.71%, (Bacillus sp. PSB9)
38.83% and Zimbabwe Bacillus 25–50 Maize Mtaita
48.76% amyloliquefaciens, (Zea Mays. et al., 2019
respectively, Paenibacillus L)
higher as polymyxa, Rhodobacter
control capsulatus,
(chemical Lactobacillus
fertilizers only) acidophillus,
Egypt Azotobacter Wheat The grains and El-Sawah Saccharomyces
chroococcum (Triticum straw yield et al., 2018 cerevisiae, Trichoderma
MF135558 (N2- aestivum L.) increased by harzianum, Aspergillus
fixing bacteria), 19.01% as oryzae,
Klebsiella compared to China Biofertilizer 50 Wheat Cisse et al.,
oxytoca control 2019
MF135559 (P-
solubilizing
bacteria) and Declaration of competing interest
Rhizobium
pusense
MF135560 (K-
The authors declare no conflict of interest.
releasing
bacteria) Acknowledgments
Indonesia Biofertilizer Sugarcane The treatment Djajadi
containing N (BL) increased the et al., 2020
Authors are grateful to Mewar University, Chittaurgarh, Rajasthan,
fixing and P number of stalk
solubilizing by 12.06% over and V.B.S. Purvanchal University, Jaunpur, U.P. for providing all the
bacteria the control necessary facilities to carry out this work.
Sudan biofertilizer (Zea mays The grain yield Obid et al.,
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