Chapter 13

Microbial Biofortification: A Green

Technology Through Plant Growth
Promoting Microorganisms

Amir Khan, Jyoti Singh, Viabhav Kumar Upadhayay, Ajay Veer Singh,

and Shachi Shah

Abstract  The hidden hunger or malnutrition is considered to be the most dignified

global challenge to human kind. Malnutrition afflicts approximately more than one
billion of world’s population in both developed and developing countries.
Malnutrition includes diet related chronic diseases as well as overt nutrient deficien-
cies which leads to morbidity, reduced physical and mental growth. However, strat-
egies to enhance supplementation of mineral elements and food fortification have
not always been successful. Plant growth promoting microorganisms are known to
fortify micro- and macro-nutrient contents in staple food crops through various
mechanisms such as siderophore production, zinc solubilization, nitrogen fixation,
phosphate solubilization, etc. Inoculation of potential microorganisms along with
mineral fertilizers can increase the uptake of mineral elements, yield and growth.
Therefore, biofortification of staple food crops by the implications of plant growth
promoting microorganisms has an ability to attain mineral elements, is advocated as
novel strategy not only to increase concentration of micronutrient in edible food
crops but also to improve yields on less fertile soils.

Keywords  Microbial biofortification · Plant growth promoting microorganisms ·

Malnutrition · Zinc · Micronutrient

A. Khan · J. Singh · V. K. Upadhayay · A. V. Singh (*)

Department of Microbiology, College of Basic Sciences and Humanities, GBPUA&T,
Pantnagar, Uttarakhand, India
S. Shah
School of Interdisciplinary and Transdisciplinary Studies, Indira Gandhi National Open
University, New Delhi, India

© Springer Nature Singapore Pte Ltd. 2019 255

S. Shah et al. (eds.), Sustainable Green Technologies for Environmental
Management, https://doi.org/10.1007/978-981-13-2772-8_13
256 A. Khan et al.

13.1  Introduction

For sustainable agriculture, the use of microbial based biofertilizers has prestigious
role in enhancing level of crop productivity and in food safety. Microorganisms as
invisible soil engineers maintain soil health, construct a hub for different biogeo-
chemical cycles (Gadd 2010) and many soil microorganisms such as bacteria, acti-
nomycetes, cyanobacteria and mycorrhiza present an eco-friendly approach for
improved uptake of nutrients and enhanced plant growth. Microorganisms particu-
larly plant growth promoting rhizobacteria (PGPR) are dwelled in rhizospheric
region and efficiently colonize the roots of plants and confer tolerance in plants
against several abiotic and biotic stresses (Prasad et al. 2015). Plant growth promot-
ing microorganisms make nutrients available to plants by numerous mechanisms
such as atmospheric nitrogen fixation, solubilizing the nutrients fixed in the soil
matrix, and production of phytohormones. Such microorganisms make sure for fur-
ther enhancement of micronutrients in plants, as they play a key role in organic mate-
rial mineralization and as well as transforming inorganic nutrients. Microorganisms
can also influence nutrient availability through presenting different characteristics
such as chelation, solubilization and oxidation or reduction (Khan 2005) and also
conferred resistance from pathogens causing diseases to the host plant by the secre-
tions of antibiotics (Bonfante and Genre 2015). The aim of modern agriculture sys-
tem, besides augmented crop yield, is also to produce nutritious safe food crops with
improved level of micronutrients in the edible portion of crop plants. Human popula-
tion is mainly dependent on crop based foods for the basic diet, and having foods
with poor level of essential micronutrients creates serious health issues in humans.
Deficiency of micronutrients (zinc, iron, selenium, copper, manganese and vitamins)
in both humans and plants is narrated as ‘hidden hunger’ (Sharma et al. 2016), and
bestows threat of malnutrition among world population. Therefore, implementation
of biofortification strategy is an important mode for providing the preeminent solu-
tion for producing food crops with elevated level of necessary micronutrients.
‘Biofortification’ and ‘standard fortification’ are two different terms, where bioforti-
fication is related with consigning the nutrients aggregation inside plant cells whereas
latter involves use of additives with the foods. Biofortification process deals with
several approaches for enhancing bioavailability of key nutrients in crops.

13.2  Micronutrient Associated Malnutrition

Micronutrient deficiency occurs in humans, where populations of developing coun-

tries intake diet in the form of staple foods characterized by reduced bioavailability
of essential micronutrients. Prevalence of micronutrient deficiency increase the risk
of extensive disease burden in low and middle income countries (Black 2014),
where populations of impecunious people cannot afford costlier nutrient rich foods
and other nutrient supplements and suffers from wide varieties of micronutrients
13  Microbial Biofortification: A Green Technology Through Plant Growth Promoting… 257

malnutrition associated ailments. According to the United Nations System Standing

Committee on Nutrition (UNSSCN) (2004) micronutrients starvation are associated
with more than 50% of all child mortality and also present the foremost risk factor
for maternal mortality. Zinc (Zn), iron (Fe), and selenium (Se) are considered as
important micronutrients and these are required in appropriate amount through rou-
tine diet for maintaining several life processes. Deficiency of one or more micronu-
trients creates negative impact on human health express in wide arrays of diseases
(Fig.  13.1). The micronutrient zinc is most essential for all organisms including
humans, and also has important structural roles in several proteins. Zinc deficiency
is most prevalent micronutrient dearth and is associated with numerous human
health related issues such as impairments of physical growth, greater risk of various
infections, retarded growth, deferred wound healing, diarrhea, skeletal abnormali-
ties and increased risk of abortion (Salgueiro et al. 2000).
Deficiency of iron causes chlorosis in plants and results in reduced crop yield, and
eventually affects human health through food-chain, specifically to people whose diets
generally rely on plant resources. Iron deficiency engenders nutritional anemia and
also associated with impaired immune functions in children and as well impaired neu-
rocognitive development (Murray-Kolb 2013). Selenium is another instance of essen-
tial micronutrient possessing role in wide range of metabolic pathways such as
antioxidant defense and thyroid hormone metabolism. Selenium (Se) deficiency may
linked with numerous ailments including heart diseases, reduced male fertility, hypo-
thyroidism, weakened immune system and high risk of infections, cancer, oxidative
stress-related conditions and epilepsy (Hatfield et al. 2014). Deficiency of vitamins
(Vitamin B6, Vitamin B12, Vitamin C, Vitamin E and folic acid), zinc and iron also

Retarded growth
Zinc Delayed wound healing
Micronutrients malnutrition

Skeletal abnormalities
Impairments in physical
growth and Increased risk of
infection

Iron Anemia
fatigue and
impaired immune functions

Hypothyroidism and Reduced

Selenium male fertility
Weakened immune system and
cretinism
Mental slowing
Enhanced susceptibility to infections

Fig. 13.1  Schematic representation of health effects of micronutrient (Zn, Fe and Se) malnutrition
in humans
258 A. Khan et al.

results in DNA damage through using similar strategies as radiation and various chem-
icals, and therefore considered a major factor to cause cancer and other disabilities. To
circumvent problems associated with micronutrients malnutrition, investigation of
those strategies are required which can improve the nutrient assimilation in plants.

13.3  Approaches for Biofortification

13.3.1  Biofortification Through Genetic Modification

Biofortification of vital food crops through genetic amendment and various biotechno-
logical techniques is a sustainable solution for alleviating the micronutrient malnutri-
tion. The techniques of genetic modification are being optimized for the development
and production of healthy foods in addition to step up in the levels and activity of
biologically active components in food crop system. Techniques of genetic modifica-
tion have typically been targeted at escalating yields of staple food crops in developing
countries. Though, the food crops with improved nutritional quality have gathered less
consideration. An excellent example of a genetically modified biofortified crop is
golden rice. Ordinary rice is not able to synthesise beta-carotene; however, due to
genetic modification, golden rice can produce rice with beta-carotene in it. Moreover,
stearidonic acid assimilation in soybean crop is also reported through genetic transfor-
mation (Singh et al. 2017).

13.3.2  Transgenic Approaches

Biotechnological techniques accredit the screening and selection of flourishing gen-

otypes, isolation as well as cloning of favorable traits and formation of transgenic
crops for sustainable agriculture system. Transgenic approaches can be used to
increase the micronutrient content of staple food crops such as legumes and cereals,
which can be achieved by insertion of specific genetic trait with the ability to pro-
duce the desired nutrients that are typically deficient in recipients. It involves the
characterization, insertion or deletion of specific gene to improve the desired trait
like nutritional quality from donor organisms. This may be achieved by the intro-
duction of genes that code for trace element binding proteins, over expression of
storage proteins already present or the expression of other proteins that are respon-
sible for micronutrient uptake in plants. Furthermore, metabolic pathways from any
microorganism and other organisms can also be applied into crops to utilize alterna-
tive pathways for metabolic engineering. Thus, these technologies provide a power-
ful tool that is unconstrained by the gene pool of the host. In addition, the transgenic
approaches can be targeted to the edible portions of commercial crops (Hirschi
2009). As shown in Table 13.1, several crops have been genetically modified with
traits of macronutrient and micronutrient that may provide reimbursement to
­consumers (Newell 2008).
13  Microbial Biofortification: A Green Technology Through Plant Growth Promoting… 259

Table 13.1  Genetically modified crops with description of macronutrient and micronutrient
assimilation
Characteristics Crop (details of characteristics)
Protein and amino acids
Quality of protein Maize (amino acid composition; protein↑)
and level
Potato (amino acid composition; protein↑)
Rice (amino acid composition; protein↑)
Soybean (amino acid balance)
Sweet potato (protein↑)
Essential amino acid
Maize (Lys↑, Met↑, Trp↑)
Potato (Met ↑)
Sorghum (Lys↑)
Soybean (Lys↑, Trp↑, Cys↑, Met↑)
Oils and fatty acids
Canola (lauric acid↑; + ω-3 fatty acids; 8:0 and 10:0 fatty acids↑; lauric
and myristic acids↑; oleic acid↑; γ-Linolenic acid)
Cotton (oleic acid↑, oleic + stearic acids↑)
Grass, legumes (↓trans-fatty acids)
Linseed (+ ω-3 and ω-6 fatty acids)
Maize (oil↑)
Oil palm (oleic acid↑ or stearic acids↑, oleic acid↑,+palmitic acid↓)
Rice (α-linolenic acid↑)
Soybean (oleic acid↑, α-linolenic acid↑, stearidonic acid↑, Arachidonic
acid↑)
Carbohydrates
Fructans Maize (fructan↑)
Potato (fructan↑)
Starch Rice (amylase↑)
Wheat (amylose↑)
Micronutrients and functional metabolites (Vitamins and carotenoids)
Canola (vitamin E↑)
Maize (vitamin E↑, vitamin C↑, provitamin A)
Mustard (+β-carotene)
Soyabean (Vitamin E)
Potato (β-carotene and lutein↑)
Rice (+β-carotene, Vitamin B9↑)
Wheat (provitamin A↑)
Strawberry (vitamin C↑)
Tomato (folate↑, phytoene and β-carotene↑, lycopene↑)
(continued)
260 A. Khan et al.

Table 13.1 (continued)
Characteristics Crop (details of characteristics)
Mineral availabilities
Alfalfa (phytase↑)
Carrot (calcium↑)
Lettuce (iron↑)
Rice (iron↑, zinc↑)
Maize (phytase↑, ferritin↑)
Soybean (phytase↑)
Wheat (phytase↑, iron↑, zinc↑)
Alfalfa (phytase↑)
Carrot (calcium↑)
Lettuce (iron↑)
Barley (zinc↑, phytase↑)
Modified from Singh et al. (2017)

13.3.3  Agronomic Biofortification

Biofortification through agronomical approach can be achieved through the impli-

cation of nutrient-rich fertilizers to foliage or soil to increase the micronutrient con-
centration in edible crop parts and thus increase the intake of essential micronutrients
by consumers. Interaction between micronutrient and macronutrient can influence
the efficiency of agronomic biofortification. Good quantity of macro elements (N, P
and K) in crop has a positive effect on development of root architecture and trans-
portation of nutrients from vegetative tissues to the seeds. Consequently, there is
increased concentration of micronutrients in edible parts of the food crop (Prasad
et al. 2014). However, when food crops are grown where mineral elements become
straight away unavailable in the soil system, targeted application of soluble chemi-
cal fertilizers to foliar parts and roots are practised.

13.3.4  Chemical Fertilizer

Effectiveness of chemical fertilizer on crop performance is influenced by type of

fertilizer. The interaction between chemical fertilizer and different forms of nutri-
ents can have positive, negative or neutral effect on food crop in yield and nutrient
bioavailability. The implication of fertilizer with soil is often the most efficient man-
ner. However, soils often contain huge amounts of iron, but only little amount of
iron is phytoavailable. The implication of inorganic Fe fertilizers to such soils is
more often futile as it rapidly becomes unavailable to plant root system through
adsorption, oxidation reactions and precipitation. For this reason, Fe chelators are
often used as soil Fe fertilizers (Rengel et al. 1999). Zinc is commonly applied to
13  Microbial Biofortification: A Green Technology Through Plant Growth Promoting… 261

crops as ZnSO4 or as synthetic chelators (Shuman 1998). The application of Zn

fertilizers to the soil system is effective for increasing Zn concentrations in cereal
grains, growing mostly in soils and foliar applications of either ZnSO4 or Zn chela-
tors can increase Zn concentrations in plant via ample Zn mobility in the phloem.
Similarly, applications of Zn fertilizers in soil and foliar can increase Zn concentra-
tions in leaf, tuber and fruit (Shuman 1998; Rengel et al. 1999).

13.3.5  B
 iofortification Through Plant Growth Promoting
Microorganisms (PGPM)

Biofortification of crops through implications of PGPMs can be considered as a

promising accompanying measure, which along with transgenic varieties, can lead
to augmented micronutrient concentrations in food crop system, besides improving
yield and soil fertility. Plant growth promoting microorganism’s have been reported
to biofortify the micronutrient contents in food crops besides improving the soil
fertility and crop yield (Rana et  al. 2012). In addition, plant growth promoting
microorganisms also facilitate the plant growth through N2 fixation, insoluble phos-
phorus solubilization, production of phytohormones, lowering of ethylene concen-
tration, antibiotics and antifungal metabolites synthesis and induced systemic
resistance. In this way, PGPM are also known to boost the soil fertility in return, the
plant acquiesce by supplying essential nutrients, growth regulators and enhancing
the ethylene mediated stress by 1-aminocyclopropane-1-carboxylate (ACC) deami-
nase production along with improved plant stress tolerance to drought, salinity,
metal and toxicity of pesticide (Singh and Prasad 2014; Singh and Singh 2017).
Moreover, the potentiality of PGPM in agriculture is progressively increased as it
provides an attractive approach to replace the exploitation of chemical fertilizers,
pesticides and other supplements. Subsequently, biofortification of crops through
application of PGPMs can be therefore considered as a potential supplementary
approach, which along with breeding varieties, can escort to augment the concentra-
tions of micronutrient in wheat crop, besides improving yield and soil fertility
(Singh et al. 2017).

13.4  M
 echanisms of Plant Growth Promoting
Microorganisms

A variety of plant growth promoting microorganisms (PGPM) has been reported to

enhance plant growth and productivity by means of various mechanisms. Illustration
of some of the mechanisms is as follows.
262 A. Khan et al.

13.4.1  Iron Chelation

Iron is a vital component for all forms of life including prokaryotes as well as
eukaryotes. It is the component of electron transport carrier, cofactor of various
enzymes and important part of various constituent such as hemoglobin. Due to the
aerobic environment conditions, iron is present in its oxidized form (Fe3+, insoluble
at neutral pH) instead of reduced form (Fe2+, soluble at neutral pH) which are taken
up by plants. To sequester the iron, many fungi, bacteria and some plants have an
unusual adaptation to produce low molecular weight compounds called as sidero-
phores, a group of low molecular weight compounds (<10 KD) those have immense
affinity towards Fe3+ ions. Siderophores are PGPM secreted compounds that are
ultimately taken up by plants therefore transporting molecule of iron to the plants
cells. Plant roots might be able of take up siderophore and use them as sources of
iron. Therefore, microbial siderophore can enhance plant growth by improving iron
uptake as well as by inhibiting the plant pathogen by means of competition ulti-
mately leads to the iron biofortification in plants and their grains (Srivastava et al.
2013). Many researchers has been reported the siderophore production in wide
range of bacterial species viz. Bacillus, Pseudomonas, Azotobacter, Arthrobacter,
Burkholderia, Enterobacter, Rhodospirrilum, Serratia, Azospirillum and Rhizobium
and fungal species viz. Aspergillus Penicillium Rhizopus, Syncephalastrum (Leong
and Neilands 1982; Das et al. 2007; Duran et al. 2016; Srivastava et al. 2013). There
are many types of siderophore such as hydroxamate, catacholate and carboxalate
that are secreted by microbes varies from species to species. Furthermore, mixed
type of siderophore has been secreted by many baceterial species (Wandersman and
Delepelaire 2004). Hence we can say that use of siderophore producing PGPM is
better approach over other conventional methods such as chemical fertilizers to
enhance iron content in plants and grains.

13.4.2  Zinc Solubilizer

Zinc is one of the essential nutrients required for growth and metabolic activities.
Zinc ions takes part in many physiological activities, it act as cofactor in various
enzymes; take part in defense; play role in cell division and growth in prokaryotes
as well as in eukaryotes. Zinc ions are highly reactive in nature and present in close
interaction with soil constituents therefore soluble zinc is very low in soil. Generally,
it is found in the form of oxides, phosphates and carbonates. Plant associated micro-
organisms adopt several mechanisms to solubilize zinc such as chelation (Whiting
et al. 2001), reduction in soil pH (Subramanian et al. 2009), or through improving
root growth and root absorptive area (Burkert and Robson 1994). Zinc chelation by
microbes and making them available for plants roots is a well known phenomenon.
Microbes produce chealating compounds, which forms complex upon binding with
zinc. In addition, they releases chealated zinc at the root surface and enhance the
13  Microbial Biofortification: A Green Technology Through Plant Growth Promoting… 263

zinc availability ultimately lead to the zinc biofotification in plants. Whiting et al.
(2001) reported production of metallophores as the possible strategies used by bac-
teria to chelate Zn. Reduction in soil pH also enhance availability of zinc. Decline
in pH has been reported, when Pseudomonas and Bacillus spp. solubilized zinc
complex compounds (ZnS, ZnO and ZnCO3) into zinc ions in a broth culture
(Saravanan et al. 2004). Zinc solubilization methods differ from one microorganism
to another to improve Zn availability in soil system and plant tissues. Many microbes
including many bacterial and fungal species (Pseudomonas, Microbacterium,
Enterobacter, Bacillus, Arbuscular mycorrhizae) have the incredible capability to
solubilize Zn from complex compounds (Whiting et al. 2001; Fasim et al. 2002;
Subramanian et al. 2009) and consequently take part in improvement of food quality
and nutrient status of plants and grains.

13.4.3  Biofertilizers

In the present era, use of chemical fertilizers to enhance plant growth, productivity
and to replenish soil nutrient status is very common. But several problems coincide
with the use of chemical fertilizers such as high cost, unavailability of large portion
of nutrients, toxic and non-degradable nature, leading to enhancement of environ-
mental pollution and making land unsuitable for cultivation. Therefore as an alter-
native strategy, application of biofertilizers can be used to enhance crop productivity
and biofortification of nutrients in grains. Biofertilizers which are the fertilizers
based on source of biological origin such as microbes (including bacteria and fungi)
are used instead of synthetic compounds. Use of biofertilizers is an ecofriendly
approach as they are of biological origin and keeps environment healthy due to its
low persistence. The objective of biofertilizers is to increase the soil organic con-
tents improve soil structure and reduce loss of nutrients such as nitrogen, iron, zinc,
calcium and phosphorus (Lal and Greenland 1979). Biofertilizers serves as source
of all nutrients due to their ability to solubilize complex form of nutrients into solu-
ble form (Singh et al. 2010, 2013, 2018). Plants utilize phosphorus in the form of
orthophosphate (Pi). Plant growth promoting microorganisms possess various
mechanisms to enhance phosphate solubilization (Fig.  13.2). Jones et  al. (1998)
reported 3.1–4.7 times more efficacy in mycorrhizal associated plants for phospho-
rus uptake than nonmycorrhizal plants. Commercially biofertilizers are developed
by coating of various bacteria (Azotobacter, Azospirillum, Rhizobium, Pseudomonas
and Bacillus) on seeds, a process called bacterization. Azotobacter chrococcum
secretes azotobacterin, Bacillus megaterium secretes phosphobacterin, which is
used for preparation of biofertilizers (Kumar and Bohra 2006). These bacteria may
or may not form symbiotic association but enhances lateral root hairs of plants to
increase mineral and water absorption, also increases nitrogen availability, secretes
plant growth stimulating substances such as vitamin, auxins, gibberellic acid, cyto-
kinins which leads to increase in photosynthesis capacity ultimately enhancing
nutrient status in plants. Rhizobial biofertilizers are able to fix 50–150 N/ha/annum.
264 A. Khan et al.

Fig. 13.2  Phosphate solubilization mechanisms of plant growth promoting microorganisms

It has been well known, that application of biofertilizers with plants, significantly
increases plant growth, high nutrient status, low level of pathogen attack (Gupta
et al. 2003; Yadav et al. 2016). Therefore, such free living and symbiotic microor-
ganisms are promoted to reduce the dependence on chemical fertilizers. Some bio-
fertilizers are listed in Table 13.2.

13.4.4  Biocontrol Agents

Under natural environmental conditions, plants are continuously exposed to various

pathogenic bacteria and fungi, which causes disease in plants leading to the reduc-
tion in crop production or death of the plants. Therefore, plant diseases need to be
controlled to maintain the quality and nutritional status of plants. Different
approaches may be used to prevent or control plant diseases. In present circum-
stances, many pesticides are used to prevent disease but the disadvantage is that the
pest may adapted towards pesticide and it is also cost intensive as well as act as
environmental pollutant due to its persistence. Therefore, there is a novel approach
i.e. application of biocontrol agent to suppress or demise pathogen growth.
Biocontrol agents are microorganisms, which controls the fungi, insect, pest and
any other pathogen (Beattie 2006). Generally bacteria, fungi, virus and protozoans
are used as biocontrol agents. Generally plant growth promoting microorganisms
produces various substances that protect plants against pathogens by direct interac-
tions i.e. antagonistic activity or indirectly by inducing host resistance (Induce sys-
temic resistance or systemic acquired resistance). Plant growth promoting
microorganisms that indirectly act on pathogens may have some mechanisms to
control plant pathogen such as the following (Table 13.3).
13  Microbial Biofortification: A Green Technology Through Plant Growth Promoting… 265

Table 13.2  Biofertilizers, their mode of action and crops benefitted

Organisms Action Crop References
Bacteria Rhizobium Symbiotic nitrogen All leguminous Bagali (2012)
leguminosarum fixation crops
Bacillus Phosphate Mustard Kang et al. (2014)
megaterium solubilization
Bacillus subtilisMicronutrient Cotton Yao et al. (2006)
solubilizer
Pseudomonas Micronutrient Bean Alemu (2013)
fluorescens solubilizer
Azotobacter sp. Free living nitrogen Leguminous Bagali (2012)
fixation crops
Azospirillum sp. Associated Symbiotic Leguminous Bagali (2012)
nitrogen fixation crops
Fungi Penicillium bilaiae Phosphate Coffee, Malhi et al. (2013)
solubilization Casurina
Glomex sp. Phosphate Coffee, Malhi et al. (2013)
solubilization Casurina
Cyanobacteria Nostoc Free living nitrogen Rice Vaishampayan
fixation et al. (2001)
Anabana Free living nitrogen Rice Vaishampayan
fixation et al. (2001)
Anabana- Azolla Symbiotic nitrogen Rice Vaishampayan
fixation et al. (2001)

Table 13.3  Major biocontrol agents, their target pathogen and mechanism of action
Biocontrol
agent Target pathogen Crop Action References
Bacillus R. solani, All Cotton Lytic enzymes Shaikh and Sayyed
thuringiensis phytopathogen (2015)
Pseudomonas Erwinia Potato, Siderophor, Shaikh and Sayyed
fluorescence carotovora,Puccinia wheat, Sugar Antibiotics (2015)
ultimum, Fusarium beat production
glycinia
Strepromyces S. sclerotiorum Potato, Lytic enzymes Shaikh and Sayyed
sp. Tomato (2015)
Trichoderma Botrytis cinerea, Bean, Lytic enzymes, Woo et al. (1999),
harzianum Meloidogyne javanica Tomato Competition Sahebani and
Hadavi (2008) and
Puyam (2016)
Trichoderma Sclerotium rolfsii Groundnut Lytic enzymes, Hirpara et al.
viride Competition (2017) and Puyam
(2016)
Pseudomonas Bipolaris maydis Maize Antibiotics Shaikh and Sayyed
cepacia production (2015)
266 A. Khan et al.

1. The PGPR may have ability to produce siderophore that chelates iron, which
makes iron unavailable for plant pathogens (Singh et al. 2017)
2. The PGPR may possess capacity to secrete some anti-pathogenic metabolites
such as antibiotics, cell wall degrading hydrolytic enzyme (Glucanases, chitin-
ases, proteases, lipase, pectinases) or hydrogen cyanide (HCN), which sup-
presses pathogen growth (Maksimov et al. 2011).
3. The PGPR may compete for nutrients and niche with pathogen (Kamilov et al.
2005).
4. The PGPR may stimulate Induced Systemic Resistance (ISR) or Systemic

Acquired Resistance (SAR) (Van loon et al. 1998).
A wide range of microorganisms are reported to act as biocontrol agents, as they
possess one or more than one mechanisms to suppress pathogen attack or growth.
Due to less persistence in environment, specificity for target pest, cost effectiveness
and ecofriendly nature, biocontrol agents are good alternative to pesticides.

13.5  Conclusion

Development of crops with elevated concentration of micronutrients is immensely

and urgently needed to combat the problem of micronutrient deficiency. Plant
growth promoting microorganisms (PGPMs) make interaction with plants and exert
plant growth promoting activities and enhance the capability of the plant for uptake
of micronutrients from surrounding soils. Zinc solubilization and siderophore
secreting microorganisms enhance the level of zinc and iron in the various edible
portions of crop plants and provide an alternative strategy to fortify micronutrients
and produce micronutrients rich foods. Application of such microorganisms inocu-
lants reduces the dependency on costly biofortification approaches i.e. agronomic
and genetic approaches. In future, formulation of microorganisms possessing mul-
tiple beneficial traits can be applied for biofortification strategies to tackle the prob-
lem of hidden hunger. Using plant growth promoting microorganisms for
biofortification as part of green technology approach can be a better strategy to
achieve environmental sustainability.

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