Role of the FliA-FlgM regulatory system on the transcriptional control of the flagellar regulon and flagellar formation in Salmonella typhimurium

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Abstract
In the flagellar regulon of Salmonella typhimurium, the flagellar operons are divided into three classes, 1, 2, and 3, with respect to transcriptional hierarchy. The class 2 operons are controlled positively by the class 1 genes, flhD and flhC. The class 3 operons are controlled positively by fliA and negatively by flgM. It has been shown that FliA is a sigma factor specific for class 3, whereas FlgM is an anti-sigma factor which binds FliA to prevent its association with RNA polymerase core enzyme. Therefore, the FliA-FlgM regulatory system has been believed to control specifically the class 3 operons. In the present study, we showed that the flgM mutation enhanced the expression of class 2 by more than fivefold. When a fliA mutation was present simultaneously, this enhancement was not observed. These results indicate that the FliA-FlgM regulatory system is involved not only in the expression of class 3 but also in that of class 2. However, though neither flhD nor flhC mutants could express the class 2 operons, the fliA mutants permitted the basal-level expression of those operons. Therefore, FlhD and FlhC are indispensable for the expression of class 2, whereas FliA is required only for its enhancement in the FlgM-depletion condition. Furthermore, we showed that the flgM mutation resulted in a two- to threefold increase in flagellar number. On the basis of these results, we propose that the relative concentration of FliA and FlgM may play an important role in the determination of flagellar numbers produced by a single cell.