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STM2209  and  STM2208  are contiguous loci annotated as putative protein-coding genes in the chromosome of  Salmonella enterica . Lack of homologs in related Enterobacteria and low G+C content suggest that  S. enterica  may have acquired  STM2209-STM2208  by horizontal transfer.  STM2209  and  STM2208  are co-transcribed from a promoter upstream  STM2209 , and their products are inner (cytoplasmic) membrane proteins. Analysis with the bacterial adenylate cyclase two-hybrid system suggests that STM2209 and STM2208 may interact. Expression of  STM2209-STM2208  is subjected to phase variation in wild type  Salmonella enterica  serovar Typhimurium. Switching frequencies in LB medium are 6.1×10 −5  (OFF→ON) and 3.7×10 −2  (ON→OFF) per cell and generation. Lack of DNA adenine methylation locks  STM2209-STM2208  in the ON state, and lack of the LysR-type factor OxyR locks  STM2209-STM2208  in the OFF state. OxyR-dependent activation of  STM2209-STM2208  expression is independent of the oxidation state of OxyR.  Salmonella  cultures locked in the ON state show alteration of O-antigen length in the lipopolysaccharide, reduced absorption of bacteriophage P22, impaired resistance to serum, and reduced proliferation in macrophages. Phenotypic heterogeneity generated by  STM2209-STM2208  phase variation may thus provide defense against phages. In turn, formation of a subpopulation unable to proliferate in macrophages may restrain  Salmonella  spread in animal organs, potentially contributing to successful infection.

STM2209-STM2208 (opvAB): A Phase Variation Locus of Salmonella enterica Involved in Control of O-Antigen Chain Length