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

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The enriched soil microbial communities in the mycorrhizosphere are often<br />

organized in the form of biofilms and probably horizontal gene transfer (HGT)<br />

among cohabiting microbial species and between plant and microbe occurs [56].<br />

Many plant-associated Pseudomonas rhizobacteria produce signal molecules for<br />

quorum-sensing regulation, which were absent from soilborne strains [57]. This<br />

indicates that quorum-sensing systems exist and are required in the mycorrhizosphere.<br />

Microbial colonies on root surfaces consist of many populations or strains,<br />

and positive and negative interpopulation signaling on the plant root occur [58],<br />

which may play an important role in the efficiency of the use of biofertilizers.<br />

In addition to the above-described interactions between AMF and rhizobacteria,<br />

certain bacterialike organisms (BLOs) reside in the AM fungal cytoplasm, first<br />

described <strong>by</strong> Mosse [53]. Khan [59] illustrated AMF spores, collected from semiarid<br />

areas of Pakistan, containing 1–10 small spherical endospores without any<br />

subtending hyphae of their own. Ultrastructural observations clearly revealed their<br />

presence in many field-collected AM fungal isolates. Because of their unculturable<br />

nature, further investigation of BLOs was hampered but current advanced electron<br />

and confocal microscopic and molecular analysis techniques have allowed us to<br />

learn more about their endosymbiotic nature. Minerdi et al. [60] reported the presence<br />

of intracellular and endosymbiotic bacteria belonging to the genus Burkholderia<br />

(a nitrogen-fixing genus) in fungal hyphae of many species of Gigasporaceae. The<br />

authors used genetic approaches to investigate the presence of nitrogen-fixing genes<br />

and their expression in this endosymbiont and found nifHDK genes in the endosymbiont<br />

Burkholderia and their RNA messengers demonstrating that they possess a<br />

molecular basis for nitrogen fixation. This discovery, as stated <strong>by</strong> Minerdi et al. [60],<br />

indicates that a fungus that improves phosphorus uptake might also fix nitrogen<br />

through specialized endobacteria. Endosymbiont Burkholderia may have an impact<br />

on AMF–PGPR–MHB–plant associations and metabolism. This finding suggests a<br />

new application scenario worth pursuing.<br />

Recent methodological developments in molecular and microscopical techniques<br />

together with those in genomes, bioinformatics, remote-sensing, proteomics and so<br />

on will assist in understanding the complexity of interactions existing between<br />

diverse plants, microbes, climates and soil.<br />

13.2.3<br />

<strong>Plant</strong> Growth Promoting Rhizobacteria<br />

13.2 Rhizosphere and Microorganismsj249<br />

Rhizobacteria include mycorrhization helper bacteria (MHB) and PGPR, which<br />

assist AMF in colonizing the plant root [61,62], phosphorus solubilizers, free-living<br />

and symbiotic nitrogen fixers, antibiotic-producing rhizobacteria, plant pathogens,<br />

predators and parasites [63]. The most common bacteria in the mycorrhizosphere<br />

are Pseudomonas [64], while different bacterial species exist in the hyphosphere.<br />

Like AMF, rhizobacteria such as pseudomonads are also ubiquitous members of<br />

the soil microbial community and have received special attention as they also exert<br />

beneficial effects on plants <strong>by</strong> suppressing soilborne pathogens, synthesizing phytohormones<br />

and promoting plant growth [65–68]. Many fluorescent Pseudomonas

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