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Journal of Bacteriology, September 2004, p. 6003-6014, Vol. 186, No. 18
0021-9193/04/$08.00+0     DOI: 10.1128/JB.186.18.6003-6014.2004
Copyright © 2004, American Society for Microbiology. All Rights Reserved.

DegU-P Represses Expression of the Motility fla-che Operon in Bacillus subtilis

Giuseppe Amati,{dagger} Paola Bisicchia,{dagger},{ddagger} and Alessandro Galizzi*

Dipartimento di Genetica e Microbiologia and Centro di Eccellenza in Biologia Applicata, Università degli Studi di Pavia, Pavia, Italy

Received 12 March 2004/ Accepted 17 June 2004

Bacillus subtilis implements several adaptive strategies to cope with nutrient limitation experienced at the end of exponential growth. The DegS-DegU two-component system is part of the network involved in the regulation of postexponential responses, such as competence development, the production of exoenzymes, and motility. The degU32(Hy) mutation extends the half-life of the phosphorylated form of DegU (DegU-P); this in turn increases the production of alkaline protease, levan-sucrase, and other exoenzymes and inhibits motility and the production of flagella. The expression of the flagellum-specific sigma factor SigD, of the flagellin gene hag, and of the fla-che operon is strongly reduced in a degU32(Hy) genetic background. To investigate the mechanism of action of DegU-P on motility, we isolated mutants of degU32(Hy) that completely suppressed the motility deficiency. The mutations were genetically mapped and characterized by PCR and sequencing. Most of the mutations were found to delete a transcriptional termination signal upstream of the main flagellar operon, fla-che, thus allowing transcriptional readthrough from the cod operon. Two additional mutations improved the {sigma}A-dependent promoter sequence of the fla-che operon. Using an electrophoretic mobility shift assay, we have demonstrated that purified DegU binds specifically to the PA promoter region of the fla-che operon. The data suggest that DegU represses transcription of the fla-che operon, and they indicate a central role of the operon in regulating the synthesis and assembly of flagella.

* Corresponding author. Mailing address: Dipartimento di Genetica e Microbiologia, Via Abbiategrasso 207, 27100 Pavia, Italy. Phone: 39-0382-505548. Fax: 39-0382-528496. E-mail: galizzi{at}ipvgen.unipv.it.

{dagger} G.A. and P.B. contributed equally to this work.

{ddagger} Present address: Smurfit Institute, Department of Genetics, Trinity College, Dublin, Ireland.

Journal of Bacteriology, September 2004, p. 6003-6014, Vol. 186, No. 18
0021-9193/04/$08.00+0     DOI: 10.1128/JB.186.18.6003-6014.2004
Copyright © 2004, American Society for Microbiology. All Rights Reserved.



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