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    Emergence of Plant Growth Promoting Rhizobacteria for an

    Environmentally Sustainable AgricultureNeha Upadhyay, Radha Rani, Shivesh Sharma*

    Department of Biotechnology,

    MNNIT Allahabad, Allahabad -211004 U.P.Abstract

    Plant growth promoting rhizobacteria (PGPR) are valuable bacteria commercialized as biofertilizer and biopesticides. Any technology that deals with natural environment is always turning to an innovative

    way and PGPR isolates are categorized as an eco-friendly microbial tool. PGPR inoculants are emerged as a novel technique that reduces the excessive use of chemical fertilizers and pesticide. The industrial

    requirement for PGPR is progressively increased, as it propounds attractive way to maintain a sustainable agriculture system. The successful commercialization of PGPR strains depend on the association

    between the scientific organizations and industries. A number of plant growth promoting rhizobacterial isolates has been reported that enhance plant growth either directly or indirect ly by inhibiting the growth

    of plant pathogen. Production of phytohormones (IAA), solublizing phosphate and nitrogen fixation are some of the mechanism that directly involve in plant growth and promotion. Indirect mechanism for

    plant growth promotion involves production of antifungal substances, HCN production, production of s iderophores and cell wall degrading enzymes. The miscellaneous and favourable interactions between

    plant and PGPR inoculants lead towards a hopeful solutions for environmentally susta inable agr iculture. Recent progress renders the use of PGPR as biopesticides as well as biofertilizers. Its long term

    benefits for successful agriculture are presented here.

    IntroductionFertilizers are essential components of modern agriculture because they provide

    essential plant nutrients. However, overuse of fertilizers and pesticide can cause

    unanticipated environmental impacts such as accumulation of toxic chemical

    substances in the soil, depletion of organic carbon content, micro flora and fauna,

    soil fertility having deleterious effects on crop productivity. One potential way to

    decrease negative environmental impacts resulting from continued use of chemical

    fertilizers is inoculation with plant growth-promoting rhizobacteria (PGPR). These

    bacteria influencing root growth and morphology or by aiding other beneficial

    symbiotic relationships exert beneficial effects on plant growth and development

    and many different genera have been commercialized for use in agriculture. During

    the past couple of decades, the use of plant growth promoting rhizobacteria (PGPR)

    for sustainable agriculture has increased tremendously in various parts of the world.

    Significant increases in growth and yield of agronomically important crops inresponse to inoculation with PGPR have been repeatedly reported (1, 2, 5).

    Plant Growth Promoting Rhizobacteria (PGPR)

    PGPR (Biofertilizers & Biofungicides) are Plant

    Growth-Promoting Rhizobacteria defined as root-

    colonizing bacteria that exert beneficial traits on

    plant growth and development. Root colonization

    comprises the ability of PGPR to establish on or in

    the root or rhizosphere to multiply, survive and

    colonize along the growing root in the presence of

    the indigenous microflora. Genera of PGPR

    generally include Acinobacter, Agrobacter,

    Arthrobacter, Azospirillum, Bacillus,

    Bradyrhizobium, Frankia, Pseudomonas,

    Rhizobium, Serratia, Thiobacillus, and others. In

    addition to plant growth promotion, PGPR are also

    used for controlling several plant pathogens,

    enhancement of nutrient up-take and in

    rhizomediation.

    .

    Mechanism of actionPGPR has a significant impact on plant growth

    and development in both indirect and direct

    ways. the direct promotion of plant growth by

    PGPR generally entails providing the plant withcompound that is synthesized by the bacterium

    or facilitating the uptake of nutrients from the

    environment On the other hand, indirect

    promotion of plant growth occurs when bacteria

    prevent some of the deleterious effects of a

    phytopathogenic organism by one or more

    mechanisms. (Glick, 1995; Glick et al.1999).

    Phosphate solublizationPGPR stimulate plant growth directly through increase in nutrition

    acquisition, such as phosphate solubilization, or by rendering the

    inaccessible nutrients available to the plants. Phosphorous is taken

    up from soil solution as phosphate (Pi, H2PO4

    _). Although soils

    generally contain a large amount of total P but only a small

    proportion is available for uptake by the plants. Several phosphate

    solubilizing microorganisms (PSMs) are now recorded to convert

    the insoluble form of phosphorus to soluble form through

    acidification, secretion of organic acids or protons (7) and chelation

    and exchange reactions. Phosphate-solubilizing microorganisms

    (PSM) include a wide range of symbiotic and non symbiotic

    organisms, such as Pseudomonas, Bacillus, and Rhizobiumspecies;

    actinomycetes; and various fungi-like Aspergillus and Penicillium

    species (7).

    Siderophore ProductionIron is an essential nutrient of plants, but it is relatively insoluble in soil

    solutions. Siderophores are low molecular weight, extracellular compounds

    with a high affinity for ferric iron, that are secreted by microorganisms to take

    up iron from the environment and their mode of action in suppression of

    disease were thought to be solely based on competition for iron with the

    pathogen (Bakker et al., 1993; Duijff et al., 1997). Plant roots prefer to absorb

    iron as the more reduced ferrous (Fe2+) ion, but the ferric (Fe3+) ion is more

    common in well aerated soil although it is easily precipitated in iron-oxide

    forms (Salisbury and Ross, 1992). Plants commonly excrete a soluble organic

    compound (chelators and phytosiderophores) which binds Fe3+ and helps to

    maintain it in solution. Chelators deliverthe Fe3+ to the root surface where it

    is reduced to Fe2+ and immediately. Some rhizospheric bacteria also produce

    siderophores and there is evidence that a number of plant species can absorb

    bacterial Fe3+-siderophore complexes (Wang et al., 1993). Majority of

    research on microbial siderophores in the rhizosphere is associated with their

    biocontrol activities due to their competitive effects with plant pathogens.

    Production of PhytohormonesPGPR facilitates plant growth by the production of plant growth regulators or phytohormones like IAA, gibberllic

    acid and cytokinins (5). Tryptophan has been identified as main precursor molecule for biosynthesis of IAA in

    bacteria. IAA controls a diverse array of functions in plant growth and development and acts as a key component

    in shaping plant root architecture such as root vascular tissue differentiation, regulation of lateral root initiation,

    polar root hair positioning, and root gravitropism (4).

    Development of PGPR

    inoculant

    Challenges in Field Application of PGPRThe application of PGPR for control of fungal pathogens in greenhouse systems shows considerable promise (30), due in

    part to the consistent environmental conditions and high incidence o f fungal disease in greenhouses. Achieving consistent

    performance i n the field where there is heterogeneity of abiotic and biotic factors and competition with indigenous

    organisms is more difficult. Knowledge of these factors can aid in determination of optimal concentration, timing and

    placement of inoculant, and of soil and crop management strategies to enhance survival and proliferation of the i noculant.

    The concept of engineering or managing the rhizosphere to enhance PGPR function by manipulation of the host plant,

    substrates for PGPR, or through agronomic practices, is gaining increasing attention (8 ). Development of better

    formulations to ensure survival and activity in the field and compatibility with chemical and biological seed treatments isanother area of focus; approaches include optimization of growth conditions prior to formulation and development of

    improved carriers and application technology (8,3).

    ConclusionAs our understanding of the complex environment of

    the rhizosphere, of the mechanisms of action of PGPR,

    and of the practical aspects of inoculant formulation and

    delivery increases, we can expect to see new PGPR

    products becoming available. The success of these

    products will depend on our ability to manage the

    rhizosphere to enhance survival and competitiveness of

    these beneficial microorganisms (4). Rhizosphere

    management will require consideration of soil and crop

    cultural practices as well as inoculant formulation and

    delivery (3,8). The use of multi-strain inocula of PGPR

    with known functions is of interest as these

    formulations may increase consistency in the field (8).

    They offer the potential to address multiple modes of

    action, multiple pathogens, and temporal or spatial

    variability . PGPR offer an environmentally sustainable

    approach to increase crop production and health. The

    application of molecular tools is enhancing our ability

    to understand and manage the rhizosphere and will lead

    to new products with improved effectiveness.

    References1. Ahemad, M., Khan, M.S., 2009a. Effect of insecticide-tolerant and plant growth promoting Mesorhizobium on

    the performance of chickpea grown in insecticide stressed alluvial soils. J. Crop Sci. Biotechnol. 12, 213 222.

    2. Ahemad, M., Khan, M.S., 2011k. Pseudomonas aeruginosa strain PS1 enhances growth parameters of greengram[Vigna radiata (L.) Wilczek] in insecticide-stressed soils. J. Pest Sci. 84, 123131.

    3. Bashan, Y. 1 998. Inoculants of plant growth-prom oting bacteria for use in agriculture. Biotechnol. Adv . 1 6:7 2

    9-7 7 0.

    4. Bhattacharyya, P.N., Jha, D.K., 2012. Plant growth-promoting rhizobacteria (PGPR): emergence in agriculture

    World J Microbiol Biotechnol) 28:13271350 DOI 10.1007/s11274-011-0979-9

    5. Kloepper JW, Schroth MN (1981) Relationship of in vitro antibiosis of plant growth promoting rhizobacteria to

    plant growth and the displacement of root microflora. Phytopathology 71:102010246. Glick, B.R., C.L. Patten, G.Holguin and D.M. Penrose, 1999. Biochemical and genetic Mechanism used by plant

    growth Promoting Bacteria. Imperial College Press, London, UK.7. Richardson AE, Barea JM, McNeill AM, Prigent-Combaret C (2009) Acquisition of phosphorus and nitrogen in

    the rhizosphere and plant growth promotion by microorganisms. Plant Soil 321:305339.8. Nelson, L.M. Plant Growth Promoting Rhizobacteria (PGPR): Prospects for New Inoculants 2004 Plant

    Managem ent Network. Accepted for publication 14 January 2004. Published 1 March 2004.

    Selection and characterization of

    PGPR strainsPGPR strains have diverse applications in

    agriculture, horticulture and forestry. Specific

    PGPR strains are initially selected from several

    hundreds of root-colonizing bacteria isolated

    from excised roots of field grown plants. During

    the experiment, those PGPRs that consistently

    caused statistically significant increases in root

    or shoot development or both are selected for

    further testing in agricultural field.

    Recently, selections of efficacious PGPR

    strains have been made by mass screening

    technique (4). Primary screenings of new

    isolates are done based on physiological,

    nutritional and biochemical characteristics as in

    BergeysManual of Determinative Bacteriology.

    While DNA and RNA homology tests are also

    considered as most reliable tools for the

    characterization of potent PGPR strains (4).

    Table: Examples of plant growth promoting

    rhizobacteria tested for various crop types

    Nitrogen

    FixationNitrogen is one

    Nitrogen is one of

    the most common

    nutrients required

    for optimal plant

    growth and

    productivity. Approx

    60 % of the earths

    available nitrogen is

    fixed by the process

    of biological

    nitrogen fixation and

    represents an

    economically

    beneficial and

    environmentally

    sound alternative tochemical fertilizers

    (Ladha et al., 1997.

    The process of N2fixation is carried

    out by nitrogenase

    enzyme coded by nif

    genes (Kim and

    Rees, 1994).

    Fig.3 Mechanism of P solublization

    Fig.2 mechanism of Plant growth promotion by PGPR

    Fig.1 Schematic representation of PGPR

    Fig.4 Function of siderophores