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gmhA  encodes a phosphoheptose isomerase that catalyzes the biosynthesis of heptose, a conserved component of lipopolysaccharide (LPS). GmhA plays an important role in  Yersinia pestis  biofilm blockage in the flea gut.  waaA ,  waaE , and  coaD  constitute a three-gene operon  waaAE - coaD  in  Y. pestis .  waaA  encodes a transferase that is responsible for binding lipid-A to the core oligosaccharide of LPS. WaaA is a key determinant in  Y. pestis  biofilm formation, and the  waaA  expression is positively regulated by the two-component regulatory system PhoP/PhoQ. WaaE is involved in LPS modification and is necessary for  Y. pestis  biofilm production. In this study, the biofilm-related phenotypic assays indicate that the global regulator CRP stimulates  Y. pestis  biofilm formation  in vitro  and on nematodes, while it has no regulatory effect on the biosynthesis of the biofilm-signaling molecular 3′,5′-cyclic diguanosine monophosphate. Further gene regulation experiments disclose that CRP does not regulate the  hms  genes at the transcriptional level but directly promotes the  gmhA  transcription and indirectly activates the  waaAE-coaD  transcription through directly acting on  phoPQ -YPO1632. Thus, it is speculated that CRP-mediated carbon catabolite regulation of  Y. pestis  biofilm formation depends on the CRP-dependent carbon source metabolic pathways of the biosynthesis, modification, and transportation of biofilm exopolysaccharide.

CRP Is an Activator of Yersinia pestis Biofilm Formation that Operates via a Mechanism Involving gmhA and waaAE-coaD