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

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14.5 Physiological Characterization of Rhizobacteriaj265<br />

presence of toxic metal concentrations may be more specifically related to climate,<br />

geographic location and soil management or contamination [65]. In such environments,<br />

most microorganisms have adapted through their physiological abilities to<br />

resist the stress conditions. For example, Bacillus, as one of the oligotrophic groups<br />

of soil bacteria, have shown relatively high growth rates in culture even at low<br />

concentrations of energy-yielding substrates (e.g. carbon and nitrogen) and have<br />

the ability to survive in nutrient-deficient soils. Arthrobacter are also capable of<br />

surviving in carbon- and nitrogen-deficient soils <strong>by</strong> following energy-efficient metabolic<br />

pathways [91]. Therefore, Arthrobacter are referred to as oligotrophs [92]. The<br />

oligotrophs can reproduce in the presence of minimum concentrations of organic<br />

matter, making them suitable candidates for plant root colonization in arid saline<br />

soils [93].<br />

Nutrients, in general, are limited in soil and bacteria, in response, reproduce<br />

slowly and may even remain dormant for long periods. Some of these microorganisms<br />

display considerable resistance to starvation [91]. A strong relationship between<br />

cellular adaptation of Arthrobacter globiformis to the conditions of chemostat growth<br />

and survival of starved suspensions for up to 56 days was reported [94]. A greater<br />

starvation resistance of Arthrobacter than Pseudomonas and Bacillus in a carbon-free<br />

medium was also reported in a number of studies [91,94,95].<br />

Endogenous metabolism is another important pathway for microbial survival in<br />

nature, and Arthrobacter may possibly use this mechanism to reduce metabolic rates<br />

to enhance their survival under starvation [96]. Arthrobacters were also considered to<br />

be the prominent members of the autochthonous microflora of soil that are able to<br />

maintain population levels over long periods in nutrient-limiting environments [97].<br />

By exhibiting a wide range of metabolic activities, they are able to utilize various lowmolecular-weight<br />

organic compounds and some more complex compounds as<br />

carbon and energy sources. In culture, they display a wide nutritional diversity and<br />

can even utilize aromatic compounds to survive under nutrient deficiencies. In our<br />

previous studies, some of the salt-tolerant bacterial strains such as A. globiformis<br />

ArG1 and A. tumescens ArT16 isolated from nutrient-poor saline and arid soils were<br />

able to grow in nitrogen-free medium and were considered as oligonitrophiles with<br />

the ability to fix organic nitrogen (Figure 14.2).<br />

A wide variety of free-living heterotrophic bacteria are known to utilize ammonium<br />

nitrogen as the preferred nitrogen source. Under conditions of limited ammonium<br />

nitrogen, many such bacteria are able to increase their affinity for ammonium<br />

nitrogen through depression of the assimilatory system in which the task of ammonium-nitrogen<br />

assimilation is performed <strong>by</strong> glutamine synthetase [98]. Among the<br />

bacteria capable of nitrogen assimilation, glutamine synthetase is generally the catalyst<br />

for assimilating ammonium nitrogen into organic compounds resulting from fixation<br />

of atmospheric nitrogen [99]. The troposphere serves as a vast reservoir of nitrogen<br />

(78%) that can be utilized when the preferred form of nitrogen in soil becomes limited,<br />

so that oligonitrophiles, once adapted, can easily incorporate nitrogen through biological<br />

nitrogen fixation. Adding carbon-enriched materials in excess could impose<br />

nitrogen scarcity without removing the nitrogen source. Lactate, malate and succinate<br />

are all labile carbon sources for microorganisms to enhance the nitrogenase activity.

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