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Page 2 Plant-Bacteria Interactions Edited by Iqbal Ahmad, John ...

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100j 5 Diversity and Potential of Nonsymbiotic Diazotrophic <strong>Bacteria</strong> in Promoting <strong>Plant</strong> Growth<br />

the microbial cells begin to release small signaling molecule mediated sensing<br />

response pathways. This effect has been defined as quorum sensing [206]. These<br />

microbially derived signal molecules are placed into two main categories: (i) amino<br />

acids and short peptide pheromones commonly utilized <strong>by</strong> Gram-positive bacteria<br />

[207,208] and (ii) fatty acid derivatives such as acyl homoserine lactone (AHL)<br />

utilized <strong>by</strong> Gram-negative bacteria. Cellular processes regulated <strong>by</strong> QS in bacteria<br />

are diverse and includes genetic competence development. Quorum-sensing signals<br />

and identical two-component regulatory systems are used <strong>by</strong> plant-interacting<br />

bacteria (mutualistic or pathogenic associations) to coordinate, in a cell density<br />

dependent manner or in response to changing environmental conditions, the<br />

expression of important factors for host colonization and infection. The success<br />

of invasion and survival within the host also requires that rhizobia and pathogens<br />

suppress and/or overcome plant defense responses triggered after microbial recognition,<br />

a process in which surface polysaccharides, antioxidant systems, ethylene<br />

biosynthesis inhibitors and virulence genes are involved [209]. The role of<br />

AHL and AHL analogues was also reported in Rhizobium–legume symbiosis and<br />

Pseudomonas fluorescens [210,211]. Similarly, QS systems are widespread mechanisms<br />

of gene regulation in both pathogenic and plant associative bacteria, thus<br />

requiring in-depth investigation in modulating various PGP traits and plant–<br />

bacteria interactions.<br />

5.7<br />

Critical Gaps in PGPR Research and Future Directions<br />

The inoculated PGPR may release various secondary metabolites as plant growth<br />

promoting substances. The bioproduction of these substances in contact with roots<br />

is most likely subject to direct uptake <strong>by</strong> plant roots before being catabolized <strong>by</strong> soil<br />

microbes or being immobilized in soil. It has been demonstrated that these microbially<br />

derived plant growth promoting substances can promote plant growth and<br />

development. Therefore, there is a need to provide evidence and their role. This can<br />

be explored <strong>by</strong> monitoring the synthesis of PGP substances in the rhizosphere <strong>by</strong><br />

developing analytical techniques for the separation and detection of PGP substances<br />

such as plant growth regulator in the soil and screening of microbes for the production<br />

of PGP substances in the absence or presence of a precursor.<br />

In vitro activities exhibited <strong>by</strong> various PGPR may not give the expected results<br />

under field conditions. The failure of PGPR to produce the desired effects after seed/<br />

seedling inoculation is frequently associated with their inability to colonize plant<br />

roots. The process of root colonization is complex. Several traits associated with the<br />

survivability, tolerance, competence with indigenous rhizospheric microorganisms,<br />

expression of root colonizing traits and so on are important [49]. In many agroclimatic<br />

situations such as harsh climates, population pressure, land constraints and<br />

decline of traditional soil management practices, reduced soil fertility often exists.<br />

Therefore, considering the varied agroclimatic conditions, continuous research is<br />

needed to develop region-specific bioinoculants with rhizospheric competence and

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