Autoregulator-dependent Control of Extracellular Polysaccharide Production in Phytopathogenic Bacteria

Extracellular polysaccharides (EPSs) likely provide phytopathogenic bacteria a selective advantage both inside and outside plants. Despite the relatively scant knowledge about EPS biosynthesis in phytopathogenic bacteria, it clearly is a well controlled, complex, energy-intensive process. Unexpectedly, three phytopathogenic bacteria have been found to autoregulate EPS production in response to extracellular signal compounds (pheromones) that they produce. Like many bacteria, Pantoea stewartii subsp. stewartii produces a N-acyl-homoserine lactone (AHL) autoinducer. However, unlike most AHL-dependent autoinduction systems, that in P. stewartii subsp. stewartii somehow represses EPS production in the absence of autoinducer. Instead of an AHL-dependent system (which it also has), Ralstonia solanacearum uses a novel autoregulator identified as 3-hydroxypalmitic acid methyl ester to regulate EPS biosynthesis. A lack of this autoregulator in R. solanacearum results in repression of EPS biosynthesis by a complex two-component sensor/response regulator signal cascade. Xanthomonas campestris pv. campestris has two partially overlapping autoregulatory systems. The autoregulators are incompletely characterized, but one diffusible signal factor (DSF) is thought to be a fatty acid derivative and the other diffusible factor (DF) may be a butyrolactone. The autoregulation pathways in X. campestris pv. campestris are essentially unknown, but EPS production is controlled by both the DSF and DF systems, whereas production of extracellular enzymes and pigment production are regulated independently. In a confined micro-environment, population density and intercellular concentrations of an autoregulator will increase in parallel, so autoregulation is one way that bacteria can coordinate gene expression to synthesize EPS only at high cell density. However, because there is often limited evidence that it is actually cell density that is being detected, researchers should not assume a priori that autoregulation must function for quorum sensing. Some possible reasons for why phytopathogenic bacteria would benefit from delaying EPS production are discussed.