Regulation of gene expression during swarmer cell differentiation in Proteus mirabilis

The gram-negative bacterium Proteus mirabilis can exist in either of two cell types, a vegetative cell characterized as a short rod and a highly elongated and hyperflagellated swarmer cell. This differentiation is triggered by growth on solid surfaces and multiple inputs are sensed by the cell to initiate the differentiation process. These include the inhibition of flagellar rotation, the accumulation of extracellular putrescine and O-antigen interactions with a surface. A key event in the differentiation process is the upregulation of FlhD(2)C(2), which activates the flagellar regulon and additional genes required for differentiation. There are a number of genes that influence FlhD(2)C(2) expression and the function of these genes, if known, will be discussed in this review. Additional genes that have been shown to regulate gene expression during swarming will also be reviewed. Although P. mirabilis represents an excellent system to study microbial differentiation, it is largely understudied relative to other systems. Therefore, this review will also discuss some of the unanswered questions that are central to understanding this process in P. mirabilis.