ACTIVATION OF BACILLUS-SUBTILIS TRANSCRIPTION FACTOR SIGMA(B) BY A REGULATORY REGULATORY PATHWAY RESPONSIVE TO STATIONARY-PHASE SIGNALS

Alternative transcription factor-sigma(B) of Bacillus subtilis controls a stationary-phase regulon induced under growth conditions that do not favor sporulation. Little is known about the metabolic signals and protein factors regulating the activity of sigma(B). The operon containing the sigma(B) structural gene has the gene order orfV-orfW-sigB-rsbX, and operon expression is autoregulated positively by sigma(B) and negatively by the rsbX product (rsbX = regulator of sigma B). To establish the roles of the orfV and orfW products, orfV and orfW null and missense mutations were constructed and tested for their effects on expression of the sigma(B)-dependent genes ctc and csbA. These mutations were tested in two contexts: in the first, the sigB operon was under control of its wild-type, sigma(B)-dependent promoter, and in the second, the sigB operon promoter was replaced by the inducible P(spac) promoter. The principal findings are that (i) the orfV (now called rsbV) product is a positive regulator of sigma(B)-dependent gene expression; (ii) the orfW (now called rsbW) product is a negative regulator of such expression; (iii) sigma(B) is inactive during logarithmic growth unless the rsbW product is absent; (iv) the rsbX, rsbV, and rsbW products have a hierarchical order of action; and (v) both the rsbV and rsbW products appear to regulate sigma(B) activity posttranslationally. There are likely to be at least two routes by which information can enter the system to regulate sigma(B): via the rsbX product, and via the rsbV and rsbW products.