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High-level transcription ofeps , an operon encoding biosynthesis of an exopolysaccharide virulence factor of the phytopathogenRalstonia (Pseudomonas )solanacearum , requires the products of at least seven regulatory genes (phcA ,phcB ,xpsR ,vsrA-vsrD , andvsrB-vsrC ), which are organized in three converging signal transduction cascades. BecausexpsR and thevsrB-vsrC two-component system are the most downstream cascade components required for activation ofeps , we explored how these components control transcription from theeps promoter (Peps ). Deletion and PCR mutagenesis identified an upstream region of Peps (nucleotides −82 to −62) that is critical for transcription activation by VsrB-VsrC and XpsR and also is required for negative control of Peps by the putativeeps regulator EpsR. Using PCR mutagenesis we generated thevsrC1 allele that encodes a response regulator that constitutively activates Peps in the absence of its cognate sensor, VsrB. However, activation of Peps byvsrC1 still requiredxpsR . Unexpectedly, the amino acid substitution conferring the constitutive phenotype on VsrC1 is 12 residues from its C terminus, outside the known functional domains of response regulators. Finally, a modified DNase I footprinting method was used to demonstrate specific binding of both VsrC1 and VsrC to the −72 to −62 upstream region of Peps .

Multicomponent Transcriptional Regulation at the Complex Promoter of the Exopolysaccharide I Biosynthetic Operon of Ralstonia solanacearum