Stanley NR, Findlay K, Berks BC, Palmer T:Escherichia coli strains blocked in Tat-dependent protein export exhibit pleiotropic defects in the cell envelope. J Bacteriol 2001, 183:139–144.CrossRefPubMed 34. Chanal A, Santini CL, Wu L-F: Specific inhibition of the translocation of a subset of Escherichia coli TAT substrates by the

TorA signal peptide. J Mol Biol 2003, 327:563–570.CrossRefPubMed Veliparib 35. Ali A, Johnson JA, Franco AA, Metzger DJ, Connell TD, Morris JG Jr, Sozhamannan S: Mutations in the extracellular protein secretion pathway genes ( eps ) interfere with rugose polysaccharide production in and motility of Vibrio cholerae. Infect Immun 2000, 68:1967–1974.CrossRefPubMed 36. Connell TD, Metzger DJ, Wang M, Jobling MG, Holmes RK: Initial studies

of the structural signal for extracellular transport of cholera toxin and other proteins recognized by Vibrio cholerae. Infect Immun 1995, 63:4091–4098.PubMed 37. Sandkvist M: Type II secretion and pathogenesis. Infect Immun 2001, 69:3523–3535.CrossRefPubMed 38. Zhu J, Mekalanos JJ: Quorum sensing-dependent biofilms enhance colonization in Vibrio cholerae. Dev Cell 2003, 5:647–656.CrossRefPubMed FRAX597 purchase 39. Ize B, Porcelli I, Lucchini S, Hinton JC, Berks BC, Palmer T: Novel Phenotypes of Escherichia coli tat mutants revealed by global gene expression and phenotypic analysis. J Biol Chem 2004, 279:47543–47554.CrossRefPubMed 40. Ghose AC: Adherence and colonization properties of Vibrio cholerae and diarrhoeagenic Escherichia coli. J Med Res Indian 1996, 104:38–51. 41. Heithoff DM, Mahan MJ:Vibrio cholerae biofilms: Stuck between a rock and a hard place. J Bacteriol 2004, 186:4835–4837.CrossRefPubMed 42. Sperandio V, Giron JA, Silveira WD, Kaper JB: The OmpU outer membrane protein, a potential adherence factor of Vibrio cholerae. Infect Immun 1995, 63:4433–4438.PubMed 43. Taylor RK: Bacterial adhesion to mucosal surfaces. J Chemother 1991, 3:190–195.PubMed 44. Posey JE, Tyrosine-protein kinase BLK Shinnick TM, Quinn FD: Characterization of the twin-arginine translocase secretion system of Mycobacterium smegmatis. J Bacteriol 2006, 188:1332–1340.CrossRefPubMed

45. Lipp EK, Huq A, Colwell RR: Effects of global climate on infectious disease: the cholera model. Clin Microbiol Rev 2002, 15:757–770.CrossRefPubMed Authors’ contributions LZ and ZZ performed most of the experiments in this study. LZ confirmed the function of tatABC in V. cholerae. ZZ constructed some new deletion mutants, repeated and complemented the data of the experiments, and prepared the draft. HJ provided plasmids, performed TMAO experiments, and conceived the experiments. JZ performed reverse transcription-PCR and confocal microscopy. YX performed the complementation assay of the E. coli tat gene mutants with the tat genes of V. cholerae. MY taught molecular techniques, performed cell culture, and provided critical discussion about the methodology. SG and JX participated in the design and coordination of the study.