Swarming ofPseudomonas aeruginosaIs a Complex Adaptation Leading to Increased Production of Virulence Factors and Antibiotic Resistance

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Abstract
In addition to exhibiting swimming and twitching motility,Pseudomonas aeruginosais able to swarm on semisolid (viscous) surfaces. Recent studies have indicated that swarming is a more complex type of motility influenced by a large number of different genes. To investigate the adaptation process involved in swarming motility, gene expression profiles were analyzed by performing microarrays on bacteria from the leading edge of a swarm zone compared to bacteria growing in identical medium under swimming conditions. Major shifts in gene expression patterns were observed under swarming conditions, including, among others, the overexpression of a large number of virulence-related genes such as those encoding the type III secretion system and its effectors, those encoding extracellular proteases, and those associated with iron transport. In addition, swarming cells exhibited adaptive antibiotic resistance against polymyxin B, gentamicin, and ciprofloxacin compared to what was seen for their planktonic (swimming) counterparts. By analyzing a large subset of up-regulated genes, we were able to show that two virulence genes,lasBandpvdQ, were required for swarming motility. These results clearly favored the conclusion that swarming ofP. aeruginosais a complex adaptation process in response to a viscous environment resulting in a substantial change in virulence gene expression and antibiotic resistance.