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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.

Role of {sigma}D in Regulating Genes and Signals during Myxococcus xanthus Development

Poorna Viswanathan,1 Mitchell Singer,2 and Lee Kroos1*

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 {sigma}D, an alternative {sigma} 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 {sigma}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. {sigma}D RNA polymerase is dispensable for transcription of this subset, but a distinct regulatory class, which includes genes essential for sporulation, requires {sigma}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 {sigma}D synthesized during growth occupies a position near the top of a regulatory hierarchy governing M. xanthus development, analogous to {sigma} factors that control biofilm formation of other bacteria.


* Corresponding author. Mailing address: Department of Biochemistry and Molecular Biology, Michigan State University, East Lansing, MI 48824. Phone: (517) 355-9726. Fax: (517) 353-9334. E-mail: kroos{at}msu.edu.


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.




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