Abstract
Vibrio parahaemolyticus possesses two alternate flagellar systems adapted for movement under different circumstances. A single polar flagellum propels the bacterium in liquid (swimming), while multiple lateral flagella move the bacterium over surfaces (swarming). Energy to rotate the polar flagellum is derived from the sodium membrane potential, whereas lateral flagella are powered by the proton motive force. Lateral flagella are arranged peritrichously, and the unsheathed filaments are polymerized from a single flagellin. The polar flagellum is synthesized constitutively, but lateral flagella are produced only under conditions in which the polar flagellum is not functional, e.g., on surfaces. This work initiates characterization of the sheathed, polar flagellum. Four genes encoding flagellins were cloned and found to map in two loci. These genes, as well as three genes encoding proteins resembling HAPs (hook-associated proteins), were sequenced. A potential consensus polar flagellar promoter was identified by using upstream sequences from seven polar genes. It resembled the enterobacterial sigma 28 consensus promoter. Three of the four flagellin genes were expressed in Escherichia coli, and expression was dependent on the product of the fliA gene encoding sigma 28. The fourth flagellin gene may be different regulated. It was not expressed in E. coli, and inspection of upstream sequence revealed a potential sigma 54 consensus promoter. Mutants with single and multiple defects in flagellin genes were constructed in order to determine assembly rules for filament polymerization. HAP mutants displayed new phenotypes, which were different from those of Salmonella typhimurium and most probably were the result of the filament being sheathed.
References
59
Referenced
91
-
Alam , M. , and D. Oesterhelt . 1987 . Purification, reconstitution and polymorphic transition of halobacterial flagella . J. Mol. Biol. 194 : 495 - 499 .
(
10.1016/0022-2836(87)90677-2
) / J. Mol. Biol. / Purification, reconstitution and polymorphic transition of halobacterial flagella by Alam M. (1987) -
Alm , R. A. , P. Guerry , and T. J. Trust . 1993 . The Campylobacter ~28 flaB flagellin promoter is subject to environmental regulation . J. Bacteriol. 175 : 4448 - 4455 .
(
10.1128/JB.175.14.4448-4455.1993
) / J. Bacteriol. / The Campylobacter ~28 flaB flagellin promoter is subject to environmental regulation by Alm R. A. (1993) -
Allen , R. , and P. Baumann . 1971 . Structure and arrangement of flagella in species of the genus Beneckea and Photobacterium fischeri . J. Bacteriol. 107 : 295 - 302 .
(
10.1128/jb.107.1.295-302.1971
) / J. Bacteriol. / Structure and arrangement of flagella in species of the genus Beneckea and Photobacterium fischeri by Allen R. (1971) -
Altschul , S. F. , W. Miller , E. W. Myers , and D. J. Lipman . 1990 . Basic local alignment search tool . J. Mol. Biol. 215 : 403 - 410 .
(
10.1016/S0022-2836(05)80360-2
) / J. Mol. Biol. / Basic local alignment search tool by Altschul S. F. (1990) -
Atsumi , T. , L. McCarter , and Y. Imae . 1992 . Polar and lateral flagellar motors of marine Vibrio are driven by different ion membrane forces . Nature (London) 355 : 182 - 184 .
(
10.1038/355182a0
) / Nature (London) / Polar and lateral flagellar motors of marine Vibrio are driven by different ion membrane forces by Atsumi T. (1992) -
Belas , R. , M. Simon , and M. Silverman . 1986 . Regulation of lateral flagella gene transcription in Vibrio parahaemolyticus . J. Bacteriol. 167 : 210 - 218 .
(
10.1128/JB.167.1.210-218.1986
) / J. Bacteriol. / Regulation of lateral flagella gene transcription in Vibrio parahaemolyticus by Belas R. (1986) -
Bergman , K. , E. Nulty , and L. Su. 1991 . Mutations in the two flagellin genes of Rhizobium meliloti . J. Bacteriol. 173 : 3716 - 3723 .
(
10.1128/JB.173.12.3716-3723.1991
) / J. Bacteriol. / Mutations in the two flagellin genes of Rhizobium meliloti by Bergman K. (1991) -
Broach , J. , C. Neumann , and S. Kustu . 1976 . Mutant strains (nit) of Salmonella typhimurium with a pleiotropic defect in nitrogen metabolism . J. Bacteriol. 128 : 86 - 98 .
(
10.1128/JB.128.1.86-98.1976
) / J. Bacteriol. / Mutant strains (nit) of Salmonella typhimurium with a pleiotropic defect in nitrogen metabolism by Broach J. (1976) -
Canale-Parola , E. 1978 . Motility and chemotaxis of spirochetes. Annu . Rev. Microbiol. 32 : 69 - 99 .
(
10.1146/annurev.mi.32.100178.000441
) / Rev. Microbiol. / Motility and chemotaxis of spirochetes. Annu by Canale-Parola E. (1978) -
Chow , W. - Y. , and D. Berg . 1988 . Tn5tac1, a derivative of transposon Tn5 that generates conditional mutations . Proc. Natl. Acad. Sci. USA 85 : 6468 - 6472 .
(
10.1073/pnas.85.17.6468
) / Proc. Natl. Acad. Sci. USA / Tn5tac1, a derivative of transposon Tn5 that generates conditional mutations by Chow W. (1988) -
Driks , A. , R. Bryan , L. Shapiro , and D. J. DeRosier . 1989 . The organization of the Caulobacter crescentus flagellar filament . J. Mol. Biol. 206 : 627 - 636 .
(
10.1016/0022-2836(89)90571-8
) / J. Mol. Biol. / The organization of the Caulobacter crescentus flagellar filament by Driks A. (1989) -
Friedman , A. , S. R. Long , S. E. Brown , W. J. Buikema , and F. Ausubel . 1982 . Construction of a broad host range cosmid cloning vector and its use in the genetic analysis of Rhizobium mutants . Gene 18 : 289 - 296 .
(
10.1016/0378-1119(82)90167-6
) / Gene / Construction of a broad host range cosmid cloning vector and its use in the genetic analysis of Rhizobium mutants by Friedman A. (1982) -
Geis , G. , H. Leying , S. Suerbaum , U. Mai , and W. Opferkuch . 1989 . Ultrastructure and chemical analysis of Campylobacter pylori flagella . J. Clin. Microbiol. 27 : 436 - 441 .
(
10.1128/JCM.27.3.436-441.1989
) / J. Clin. Microbiol. / Ultrastructure and chemical analysis of Campylobacter pylori flagella by Geis G. (1989) -
Gerl , L. , and M. Sumper . 1988 . Halobacterial flagellins are encoded by a multigene family . J. Biol. Chem. 263 : 13246 - 13251 .
(
10.1016/S0021-9258(18)37697-X
) / J. Biol. Chem. / Halobacterial flagellins are encoded by a multigene family by Gerl L. (1988) -
Gill , P. R. , and N. Agabian . 1983 . The nucleotide sequence of the Mr ~ 28,500 flagellin gene of Caulobacter crescentus . J. Biol. Chem. 258 : 7395 - 7401 .
(
10.1016/S0021-9258(18)32192-6
) / J. Biol. Chem. / The nucleotide sequence of the Mr ~ 28,500 flagellin gene of Caulobacter crescentus by Gill P. R. (1983) -
Guerry , P. , R. A. Alm , M. E. Power , S. M. Logan , and T. J. Trust . 1991 . Role of two flagellin genes in Campylobacter motility . J. Bacteriol. 173 : 4757 - 4764 .
(
10.1128/JB.173.15.4757-4764.1991
) / J. Bacteriol. / Role of two flagellin genes in Campylobacter motility by Guerry P. (1991) -
Gutierrez , C. , J. Barondess , C. Manoil , and J. Beckwith . 1987 . The use of transposon TnphoA to detect genes for cell envelope proteins subject to a common regulatory stimulus . J. Mol. Biol. 195 : 289 - 297 .
(
10.1016/0022-2836(87)90650-4
) / J. Mol. Biol. / The use of transposon TnphoA to detect genes for cell envelope proteins subject to a common regulatory stimulus by Gutierrez C. (1987) -
Helmann , J. D. 1991 . Alternative sigma factors and the regulation of flagellar gene expression . Mol. Microbiol. 5 : 2875 - 2882 .
(
10.1111/j.1365-2958.1991.tb01847.x
) / Mol. Microbiol. / Alternative sigma factors and the regulation of flagellar gene expression by Helmann J. D. (1991) -
Homma , M. , D. DeRosier , and R. M. Macnab . 1990 . Flagellar hook and hook-associated proteins of Salmonella typhimurium and their relationship to other axial components of the flagellum . J. Mol. Biol. 211 : 819 - 832 .
(
10.1016/S0022-2836(05)80266-9
) / J. Mol. Biol. / Flagellar hook and hook-associated proteins of Salmonella typhimurium and their relationship to other axial components of the flagellum by Homma M. (1990) -
Homma , M. , H. Fujita , S. Yamaguchi , and T. Iino . 1984 . Excretion of unassembled flagellin by Salmonella typhimurium mutants deficient in hookassociated proteins . J. Bacteriol. 159 : 1056 - 1059 .
(
10.1128/JB.159.3.1056-1059.1984
) / J. Bacteriol. / Excretion of unassembled flagellin by Salmonella typhimurium mutants deficient in hookassociated proteins by Homma M. (1984) -
Homma , M. , and T. Iino . 1985 . Locations of hook-associated proteins in flagellar structures of Salmonella typhimurium . J. Bacteriol. 162 : 183 - 189 .
(
10.1128/JB.162.1.183-189.1985
) / J. Bacteriol. / Locations of hook-associated proteins in flagellar structures of Salmonella typhimurium by Homma M. (1985) -
Hughes , K. T. , K. L. Gillen , M. J. Semon , and J. E. Karlinsey . 1993 . Sensing structural intermediates in bacterial flagellar assembly by export of a negative regulator . Science 262 : 1277 - 1280 .
(
10.1126/science.8235660
) / Science / Sensing structural intermediates in bacterial flagellar assembly by export of a negative regulator by Hughes K. T. (1993) -
Hyman , H. C. , and S. Trachtenberg . 1991 . Point mutations that lock Salmonella typhimurium flagellar filaments in the straight right-handed and lefthanded forms and their relation to filament superhelicity . J. Mol. Biol. 220 : 79 - 88 .
(
10.1016/0022-2836(91)90382-G
) / J. Mol. Biol. / Point mutations that lock Salmonella typhimurium flagellar filaments in the straight right-handed and lefthanded forms and their relation to filament superhelicity by Hyman H. C. (1991) -
Ikeda , T. , M. Homma , T. Iino , S. Asakura , and R. Kamiya . 1987 . Localization and stoichiometry of hook-associated proteins within Salmonella typhimurium flagella . J. Bacteriol. 169 : 1168 - 1173 .
(
10.1128/JB.169.3.1168-1173.1987
) / J. Bacteriol. / Localization and stoichiometry of hook-associated proteins within Salmonella typhimurium flagella by Ikeda T. (1987) - Jones , C. J. , and S. -I. Aizawa . 1991 . The bacterial flagellum and flagellar motor: structure, assembly and function . Adv. Microb. Physiol. 32 : 110 - 163 . / Adv. Microb. Physiol. / The bacterial flagellum and flagellar motor: structure, assembly and function by Jones C. J. (1991)
-
Joys , T. M. 1988 . The flagellar filament protein . Can. J. Microbiol. 34 : 452 - 458 .
(
10.1139/m88-078
) / Can. J. Microbiol. / The flagellar filament protein by Joys T. M. (1988) -
Kalmokoff , M. L. , and K. F. Jarrell . 1991 . Cloning and sequencing of a multigene family encoding the flagellins of Methanococcus voltae . J. Bacteriol. 173 : 7113 - 7125 .
(
10.1128/JB.173.22.7113-7125.1991
) / J. Bacteriol. / Cloning and sequencing of a multigene family encoding the flagellins of Methanococcus voltae by Kalmokoff M. L. (1991) -
Keen , N. T. , S. Tamaki , D. Kobayashi , and D. Trollinger . 1988 . Improved broad-host-range plasmids for DNA cloning in gram-negative bacteria . Gene 70 : 191 - 197 .
(
10.1016/0378-1119(88)90117-5
) / Gene / Improved broad-host-range plasmids for DNA cloning in gram-negative bacteria by Keen N. T. (1988) -
Kelly-Wintenberg , K. , S. L. South , and T. C. Montie . 1993 . Tyrosine phosphate in a- and b-type flagellins of Pseudomonas aeruginosa . J. Bacteriol. 172 : 2458 - 2461 .
(
10.1128/JB.175.8.2458-2461.1993
) / J. Bacteriol. / Tyrosine phosphate in a- and b-type flagellins of Pseudomonas aeruginosa by Kelly-Wintenberg K. (1993) -
Kutsukake , K. , Y. Ohya , and T. Iino . 1990 . Transcriptional analysis of the flagellar regulon of Salmonella typhimurium . J. Bacteriol. 172 : 741 - 747 .
(
10.1128/JB.172.2.741-747.1990
) / J. Bacteriol. / Transcriptional analysis of the flagellar regulon of Salmonella typhimurium by Kutsukake K. (1990) -
Logan , S. M. , T. J. Trust , and P. Guerry . 1989 . Evidence for posttranslational modification and gene duplication of Campylobacter flagellin . J. Bacteriol. 171 : 3031 - 3038 .
(
10.1128/jb.171.6.3031-3038.1989
) / J. Bacteriol. / Evidence for posttranslational modification and gene duplication of Campylobacter flagellin by Logan S. M. (1989) - Macnab R. M. 1987. Flagella p. 70-83. In F. C. Neidhardt J. L. Ingraham K. B. Low B. Magasanik M. Schaechter and H. E. Umbarger (ed.) Escherichia coli and Salmonella typhimurium: cellular and molecular biology. American Society for Microbiology Washington D.C.
-
Manoil , C. , and J. Beckwith . 1985 . TnphoA: a transposon probe for protein export signals . Proc. Natl. Acad. Sci. USA 82 : 8129 - 8133 .
(
10.1073/pnas.82.23.8129
) / Proc. Natl. Acad. Sci. USA / TnphoA: a transposon probe for protein export signals by Manoil C. (1985) -
Martin , M. , R. Showalter , and M. Silverman . 1989 . Identification of a locus controlling expression of Iuminescence genes in Vibrio harveyi . J. Bacteriol. 171 : 2406 - 2414 .
(
10.1128/jb.171.5.2406-2414.1989
) / J. Bacteriol. / Identification of a locus controlling expression of Iuminescence genes in Vibrio harveyi by Martin M. (1989) - .McCarter L. M. Unpublished observation.
-
McCarter , L. , M. Hilmen , and M. Silverman . 1988 . Flagellar dynamometer controls swarmer cell differentiation of V. parahaemolyticus . Cell 54 : 345 - 351 .
(
10.1016/0092-8674(88)90197-3
) / Cell / Flagellar dynamometer controls swarmer cell differentiation of V. parahaemolyticus by McCarter L. (1988) -
McCarter , L. , and M. Silverman . 1989 . Iron regulation of swarmer cell differentiation of Vibrio parahaemolyticus . J. Bacteriol. 171 : 731 - 736 .
(
10.1128/jb.171.2.731-736.1989
) / J. Bacteriol. / Iron regulation of swarmer cell differentiation of Vibrio parahaemolyticus by McCarter L. (1989) -
McCarter , L. , and M. Silverman . 1990 . Surface-induced swarmer cell differentiation of Vibrio parahaemolyticus . Mol. Microbiol. 4 : 1057 - 1062 .
(
10.1111/j.1365-2958.1990.tb00678.x
) / Mol. Microbiol. / Surface-induced swarmer cell differentiation of Vibrio parahaemolyticus by McCarter L. (1990) -
McCarter , L. L. , and M. Silverman . 1987 . Phosphate regulation of gene expression in Vibrio parahaemolyticus . J. Bacteriol. 169 : 3441 - 3449 .
(
10.1128/JB.169.8.3441-3449.1987
) / J. Bacteriol. / Phosphate regulation of gene expression in Vibrio parahaemolyticus by McCarter L. L. (1987) -
McCarter , L. L. , and M. E. Wright . 1993 . Identification of genes encoding components of the swarmer cell flagellar motor and propeller and a sigma factor controlling differentiation of Vibrio parahaemolyticus . J. Bacteriol. 175 : 3361 - 3371 .
(
10.1128/JB.175.11.3361-3371.1993
) / J. Bacteriol. / Identification of genes encoding components of the swarmer cell flagellar motor and propeller and a sigma factor controlling differentiation of Vibrio parahaemolyticus by McCarter L. L. (1993) -
Messing , J. 1983 . New M13 vectors for cloning . Methods Enzymol. 101 : 20 - 78 .
(
10.1016/0076-6879(83)01005-8
) / Methods Enzymol. / New M13 vectors for cloning by Messing J. (1983) -
Minnich , S. , and A. Newton . 1987 . Promoter mapping and cell cycle regulation of flagellin gene transcription in Caulobacter crescentus . Proc. Natl. Acad. Sci. USA 84 : 1142 - 1146 .
(
10.1073/pnas.84.5.1142
) / Proc. Natl. Acad. Sci. USA / Promoter mapping and cell cycle regulation of flagellin gene transcription in Caulobacter crescentus by Minnich S. (1987) -
Minnich , S. A. , N. Ohta , N. Taylor , and A. Newton . 1988 . Role of the 25-, 27-, and 29-kilodalton flagellins in Caulobacter crescentus cell motility: method for construction of deletion and Tn5 insertion mutants by gene replacement . J. Bacteriol. 170 : 3953 - 3960
(
10.1128/JB.170.9.3953-3960.1988
) / J. Bacteriol. / Role of the 25-, 27-, and 29-kilodalton flagellins in Caulobacter crescentus cell motility: method for construction of deletion and Tn5 insertion mutants by gene replacement by Minnich S. A. (1988) -
Mullin , D. , S. Minnich , L. -S. Chen , and A. Newton . 1987 . A set of positively regulated flagellar gene promoters in Caulobacter crescentus with sequence homology to the nif gene promoters of Klebsiella pneumoniae . J. Mol. Biol. 195 : 939 - 943 .
(
10.1016/0022-2836(87)90497-9
) / J. Mol. Biol. / A set of positively regulated flagellar gene promoters in Caulobacter crescentus with sequence homology to the nif gene promoters of Klebsiella pneumoniae by Mullin D. (1987) -
Pleier , E. , and R. Schmitt . 1991 . Expression of two Rhizobium meliloti flagellin genes and their contribution to the complex filament structure . J. Bacteriol. 173 : 2077 - 2085 .
(
10.1128/JB.173.6.2077-2085.1991
) / J. Bacteriol. / Expression of two Rhizobium meliloti flagellin genes and their contribution to the complex filament structure by Pleier E. (1991) - Sambrook J. E. F. Fritsch and T. Maniatis. 1989. Molecular cloning: a laboratory manual 2nd ed. Cold Spring Harbor Laboratory Cold Spring Harbor N.Y.
-
Sanger , F. , S. Nicklen , and A. R. Coulson . 1977 . DNA sequencing with chain-terminating inhibitors . Proc. Natl. Acad. Sci. USA 74 : 5463 - 5467 .
(
10.1073/pnas.74.12.5463
) / Proc. Natl. Acad. Sci. USA / DNA sequencing with chain-terminating inhibitors by Sanger F. (1977) - Schweizer , H. P. 1990 . The pUC18CM plasmids: a chloramphenicol resistance gene cassette for site-directed insertion and deletion mutagenesis in Escherichia coli . BioTechniques 8 : 612 - 616 . / BioTechniques / The pUC18CM plasmids: a chloramphenicol resistance gene cassette for site-directed insertion and deletion mutagenesis in Escherichia coli by Schweizer H. P. (1990)
-
Shinoda , S. , and K. Okamoto . 1977 . Formation and function of Vibrio parahaemolyticus lateral flagella . J. Bacteriol. 129 : 1266 - 1271 .
(
10.1128/JB.129.3.1266-1271.1977
) / J. Bacteriol. / Formation and function of Vibrio parahaemolyticus lateral flagella by Shinoda S. (1977) -
Showalter , R. E. , M. O. Martin , and M. R. Silverman . 1990 . Cloning and nucleotide sequence of luxR, a regulatory gene controlling bioluminescence in Vibrio harveyi . J. Bacteriol. 172 : 2946 - 2954 .
(
10.1128/JB.172.6.2946-2954.1990
) / J. Bacteriol. / Cloning and nucleotide sequence of luxR, a regulatory gene controlling bioluminescence in Vibrio harveyi by Showalter R. E. (1990) -
Silverman , M. , R. Showalter , and L. McCarter . 1991 . Genetic analysis in Vibrio . Methods Enzymol. 204 : 515 - 536 .
(
10.1016/0076-6879(91)04026-K
) / Methods Enzymol. / Genetic analysis in Vibrio by Silverman M. (1991) -
Silverman , M. , and M. Simon . 1974 . Flagellar rotation and the mechanism of bacterial motility . Nature (London) 249 : 73 - 74 .
(
10.1038/249073a0
) / Nature (London) / Flagellar rotation and the mechanism of bacterial motility by Silverman M. (1974) -
Sjoblad , R. D. , C. W. Emala , and R. N. Doetsch . 1983 . Bacterial flagellar sheaths: structures in search of a function . Cell Motil. 3 : 93 - 103 .
(
10.1002/cm.970030108
) / Cell Motil. / Bacterial flagellar sheaths: structures in search of a function by Sjoblad R. D. (1983) - Slot , J. W. , and H. J. Geuze . 1985 . A new method of preparing gold probes for multiple-labeling cytochemistry . Eur. J. Cell Biol. 38 : 87 - 93 . / Eur. J. Cell Biol. / A new method of preparing gold probes for multiple-labeling cytochemistry by Slot J. W. (1985)
-
Suerbaum , S. , C. Josenhans , and A. Labigne . 1993 . Cloning and genetic characterization of the Helicobacter pylori and Helicobacter mustelae flaB flagellin genes and construction of H. pylori flaA- and flaB-negative mutants by electroporation-mediated allelic exchange . J. Bacteriol. 175 : 3278 - 3288 .
(
10.1128/JB.175.11.3278-3288.1993
) / J. Bacteriol. / Cloning and genetic characterization of the Helicobacter pylori and Helicobacter mustelae flaB flagellin genes and construction of H. pylori flaA- and flaB-negative mutants by electroporation-mediated allelic exchange by Suerbaum S. (1993) -
Tomashow , L. S. , and S. C. Rittenberg . 1985 . Waveform analysis and structure of flagella and basal complexes from Bdellovibrio bacteriovorus 109J . J. Bacteriol. 163 : 1038 - 1046 .
(
10.1128/JB.163.3.1038-1046.1985
) / J. Bacteriol. / Waveform analysis and structure of flagella and basal complexes from Bdellovibrio bacteriovorus 109J by Tomashow L. S. (1985) -
Tomashow , L. S. , and S. C. Rittenberg . 1985 . Isolation and composition of sheathed flagella from Bdellovibrio bacteriovorus 109J . J. Bacteriol. 163 : 1047 - 1054 .
(
10.1128/JB.163.3.1047-1054.1985
) / J. Bacteriol. / Isolation and composition of sheathed flagella from Bdellovibrio bacteriovorus 109J by Tomashow L. S. (1985) -
Trachtenberg , S. , D. J. DeRosier , and R. M. Macnab . 1987 . Three-dimensional structure of the complex flagellar filament of Rhizobium Iupini and its relation to the structure of the plain filament . J. Mol. Biol. 195 : 603 - 620 .
(
10.1016/0022-2836(87)90185-9
) / J. Mol. Biol. / Three-dimensional structure of the complex flagellar filament of Rhizobium Iupini and its relation to the structure of the plain filament by Trachtenberg S. (1987) -
Wilson , D. R. , and T. J. Beveridge . 1993 . Bacterial flagellar filaments and their component flagellins . Can. J. Microbiol. 39 : 451 - 472 .
(
10.1139/m93-066
) / Can. J. Microbiol. / Bacterial flagellar filaments and their component flagellins by Wilson D. R. (1993)
Dates
Type | When |
---|---|
Created | 8 years, 9 months ago (Nov. 9, 2016, 11:23 a.m.) |
Deposited | 4 years, 1 month ago (July 29, 2021, 1:55 p.m.) |
Indexed | 3 months, 2 weeks ago (May 22, 2025, 11:07 a.m.) |
Issued | 30 years, 6 months ago (March 1, 1995) |
Published | 30 years, 6 months ago (March 1, 1995) |
Published Print | 30 years, 6 months ago (March 1, 1995) |
@article{McCarter_1995, title={Genetic and molecular characterization of the polar flagellum of Vibrio parahaemolyticus}, volume={177}, ISSN={1098-5530}, url={http://dx.doi.org/10.1128/jb.177.6.1595-1609.1995}, DOI={10.1128/jb.177.6.1595-1609.1995}, number={6}, journal={Journal of Bacteriology}, publisher={American Society for Microbiology}, author={McCarter, L L}, year={1995}, month=mar, pages={1595–1609} }