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BACTERIAL ROOT ZONE COMMUNITIES, BENEFICIAL ALLELOPATHIES AND PLANT DISEASE CONTROL<br />

2.4. Microbe-microbe Disruption of Quorum Sensing Mechanisms<br />

It should now be apparent that quorum sensing plays a significant role in the biology<br />

and regulation of both plant-microbe and microbe-microbe interactions. And while<br />

pathogenic bacteria depend significantly on quorum sensing regulation to coordinate<br />

the saprophytic and parasitic phases of their life cycles, plants and their adherent root<br />

zone microbial communities have evolved mechanisms by which to disrupt this strategy.<br />

For example, Variovorax paradoxus, a relatively common soil organism, is able<br />

to utilize (degrade) acyl-HSL signaling molecules as an energy source (Leadbetter<br />

and Greenberg, 2000) with the effect that those classes of bacteria relying on acyl-<br />

HSLs for cell-to-cell signaling molecules will be kept ‘unaware’ of their own presence<br />

and population density. Pierson et al. (1998) demonstrated that approximately 8% of<br />

rhizobacteria recovered at random from the surfaces of wheat roots could specifically<br />

stimulate quorum sensing gene regulated expression in adjacent P. aureofaciens<br />

bacteria. Thus different bacteria appear able to exchange quorum sensing signals,<br />

with the possibility of forming functional mixed communities (Bauer and Robinson,<br />

2002).<br />

Several instances have been reported of soil bacteria in possession of enzymes<br />

designed to degrade or inactivate acyl-HSLs (Bauer and Robinson, 2002; Dong et al.,<br />

2001, 2002; Leadbetter, 2001; Whitehead et al., 2001). A case in point is the lactonase<br />

enzyme, AiiA, from Bacillus cereus, which, it is believed, opens the lactone ring in<br />

acyl-HSLs, thereby reducing signal strength in the order of a 1000 fold (Dong et al.,<br />

2000). In circumstances where B. cereus and E. carotovora co-exist as commensals<br />

in field soils, AiiA is able to inactivate the acyl-HSL autoinducer in E. carotovora<br />

rendering the pathogen avirulent.<br />

Clearly, understanding the mechanism of signal synthesis, and being able to<br />

identify signal synthesis inhibitors, has implications for developing quorum sensingtargeted<br />

antivirulence molecules, or engineering beneficial communities which utilize<br />

acyl-HSL signals as an energy source and so inhibit pathogenic trait expression. In<br />

the former case, AHL signal-inactivating molecules (the so-called ‘quorum quashing’<br />

moieties) - namely AHL-lactonases and AHL-acylases - have already been identified<br />

in a Ralstonia sp. isolated from a mixed-species biofilm (Lin et al., 2003).<br />

However, while selection pressures upon component bacterial populations in a<br />

community will include biological pressures created by microbially mediated<br />

moderation of habitats, external abiotic pressures created by environmental<br />

perturbations - such as climate or crop management practices - also exist. As a result,<br />

even though “... microorganisms are potentially everywhere, [the] environment<br />

selects...” (Alexander, 1997).<br />

3. MICROBIAL ANTAGONISM AND DISEASE CONTROL<br />

3.1. Microbially Induced <strong>Biologica</strong>l Control in Soils<br />

Since every living soil sample will yield organisms with antagonistic activity to some<br />

other organism, or group of organisms, it has almost become axiomatic that<br />

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