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

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Table 11.3 MajorgenesinvolvedinthebiosynthesisofrhamnolipidsinPseudomonasaeruginosa[121].<br />

Gene G + C %<br />

promoter<br />

researchers showed that rhlC is coordinately regulated with a rhlAB quorum-sensing<br />

system. Major genes involved in this system are listed in Table 11.3.<br />

The rhl system also regulates the stationary phase sigma factor encoded <strong>by</strong> rpoS,<br />

which is involved in the regulation of numerous genes important for survival under<br />

adverse conditions [116]. A second quorum-sensing system, located at a different<br />

region in the P. aeruginosa chromosome, also influences the expression of rhamnolipid<br />

biosynthesis. This second system is encoded <strong>by</strong> lasR (31% homology to rhlR)<br />

and lasI (28% homology to rhlI) wherein lasI encodes the autoinducer N-(3-oxododecanoyl)-L-homo-serine<br />

lactone [122]. The interaction between both systems was<br />

described as a hierarchical quorum-sensing cascade, with lasR and lasI as the master<br />

regulators [123].<br />

Olvera et al. [124] proved that the algC gene, involved in alginate production<br />

through its phosphomannomutase activity and in LPS synthesis through phosphoglucomutase<br />

activity, participates in rhamnolipid biosynthesis in P. aeruginosa. The<br />

phosphoglucomutase activity of AlgC is responsible for the production of glucose-1phosphate,<br />

the precursor of dTDP-glucose and ultimately of dTDP-l-rhamnose,<br />

whereas products of other alg genes are involved neither in rhamnolipid production<br />

nor LPS synthesis. Pearson et al. [117] have shown that both quorum-sensing<br />

systems can induce expression of rhamnolipids and other virulence genes but with<br />

different efficiencies due to specificity of the transcriptional activator with its cognate<br />

autoinducer.<br />

11.4.3<br />

Regulation<br />

Size<br />

(nt)<br />

Peptide<br />

length<br />

Gene<br />

product<br />

(kDa)<br />

pI Function<br />

11.4 Rhamnolipidsj223<br />

RhlA 65.8 54 887 296 32.5 7.4 Rhamnosyl- transferase 1<br />

RhlB 67.9 54 1280 427 47 8.4 Rhamnosyl- transferase 1<br />

RhlC 70.7 54 975 325 35.9 nd Rhamnosyl- transferase 2<br />

RhlG Nd 54 768 256 26.8 nd NADPH-dependent ketoacyl reductase<br />

RhlR 61.7 70 726 242 26.5 7.0 Transcriptional regulator<br />

RhlI 64.8 64.8 NA NA NA NA NA<br />

The synthesis of rhamnolipids in P. aeruginosa under nitrogen depletion, when the<br />

cells shift into stationary growth phase, has been reported [125,126]. Ochsner et al.<br />

[115] observed the expression of P. aeruginosa genes for rhamnolipid synthesis in<br />

P. fluorescens and P. putida only under nitrogen-limited conditions. Addition of<br />

nitrogen inhibits the production of rhamnolipids <strong>by</strong> resting cells in Pseudomonas<br />

sp. DSM 2874 [102]. Guerra-Santos et al. [127] reported that higher levels of

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