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Journal of Bacteriology, May 2006, p. 3246-3256, Vol. 188, No. 9
0021-9193/06/$08.00+0 doi:10.1128/JB.188.9.3246-3256.2006
Copyright © 2006, American Society for Microbiology. All Rights Reserved.
D in Regulating Genes and Signals during Myxococcus xanthus Development
Department of Biochemistry and Molecular Biology, Michigan State University, East Lansing, Michigan 48824,1 Section of Microbiology and Center for Genetics and Development, The University of California-Davis, Davis, California 956162
Received 14 January 2006/ Accepted 14 February 2006
Starvation-induced development of Myxococcus xanthus is an excellent model for biofilm formation because it involves cell-cell signaling to coordinate formation of multicellular mounds, gene expression, and cellular differentiation into spores. The role of
D, an alternative
factor important for viability in stationary phase and for stress responses, was investigated during development by measuring signal production, gene expression, and sporulation of a sigD null mutant alone and upon codevelopment with wild-type cells or signaling mutants. The sigD mutant responded to starvation by inducing (p)ppGpp synthesis normally but was impaired for production of A-signal, an early cell density signal, and for production of the morphogenetic C-signal. Induction of early developmental genes was greatly reduced, and expression of those that depend on A-signal was not restored by codevelopment with wild-type cells, indicating that
D is needed for cellular responses to A-signal. Despite these early developmental defects, the sigD mutant responded to C-signal supplied by codeveloping wild-type cells by inducing a subset of late developmental genes.
D RNA polymerase is dispensable for transcription of this subset, but a distinct regulatory class, which includes genes essential for sporulation, requires
D RNA polymerase or a gene under its control, cell autonomously. The level of sigD transcript in a relA mutant during growth is much lower than in wild-type cells, suggesting that (p)ppGpp positively regulates sigD transcription in growing cells. The sigD transcript level drops in wild-type cells after 20 min of starvation and remains low after 40 min but rises in a relA mutant after 40 min, suggesting that (p)ppGpp negatively regulates sigD transcription early in development. We conclude that
D synthesized during growth occupies a position near the top of a regulatory hierarchy governing M. xanthus development, analogous to
factors that control biofilm formation of other bacteria.
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