Biological diversity of prokaryotic type IV secretion systems

Microbiology and Molecular Biology Reviews

Volumn 73, Issue 4, 2009, Pages 775-808

  Alvarez Martinez, Cristina E ^a   Christie, Peter J ^a  

^a UNIVERSITY OF TEXAS MEDICAL SCHOOL   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

ADAPTOR PROTEIN; ADHESIN; BACTERIAL PROTEIN; CARRIER PROTEIN; CHAPERONE; GLYCOSYLTRANSFERASE; PROTEIN VIRB1; PROTEIN VIRB10; PROTEIN VIRB11; PROTEIN VIRB2; PROTEIN VIRB3; PROTEIN VIRB5; PROTEIN VIRB6; PROTEIN VIRB7; PROTEIN VIRB8; PROTEIN VIRB9; UNCLASSIFIED DRUG;

BACTERIAL GENE; BACTERIAL MEMBRANE; BACTERIUM CONJUGATION; BACTERIUM PILUS; BIODIVERSITY; GENE CLUSTER; GENE SEQUENCE; GRAM NEGATIVE BACTERIUM; GRAM POSITIVE BACTERIUM; HORIZONTAL GENE TRANSFER; MOLECULAR RECOGNITION; NONHUMAN; NUCLEOTIDE SEQUENCE; PLASMID; PROKARYOTE; PROTEIN SECRETION; REVIEW; SIGNAL TRANSDUCTION; TYPE III SECRETION SYSTEM; TYPE IV SECRETION SYSTEM;

ANIMALS; BACTERIAL INFECTIONS; BACTERIAL PROTEINS; BIODIVERSITY; CRENARCHAEOTA; DNA; EVOLUTION, MOLECULAR; GRAM-NEGATIVE BACTERIA; GRAM-POSITIVE BACTERIA; HUMANS; SECRETORY PATHWAY;

AGROBACTERIUM TUMEFACIENS; BACTERIA (MICROORGANISMS); EUKARYOTA; NEGIBACTERIA; PROKARYOTA;

EID: 71449116162     PISSN: 10922172     EISSN: None     Source Type: Journal    
DOI: 10.1128/MMBR.00023-09     Document Type: Review

References (300)

1. Abajy, M. Y., J. Kopec, K. Schiwon, M. Burzynski, M. Doring, C. Bohn, and E. Grohmann. 2007. A type IV-secretion-like system is required for conjugative DNA transport of broad-host-range plasmid pIP501 in gram-positive bacteria. J. Bacteriol. 189:2487-2496.
2. Abdallah, A. M., N. C. G. van Pittius, P. A. Champion, J. Cox, J. Luirink, C. M. Vandenbroucke-Grauls, B. J. Appelmelk, and W. Bitter. 2007. Type VII secretion - mycobacteria show the way. Nat. Rev. Microbiol. 5:883-891.
3. Al-Khodor, S., C. T. Price, F. Habyarimana, A. Kalia, and Y. Abu Kwaik. 2008. A Dot/Icm-translocated ankyrin protein of Legionella pneumophila is required for intracellular proliferation within human macrophages and protozoa. Mol. Microbiol. 70:908-923.
4. Aly, K. A., and C. Baron. 2007. The VirB5 protein localizes to the T-pilus tips in Agrobacterium tumefaciens. Microbiology 153:3766-3775.
5. Amor, J. C., J. Swails, X. Zhu, C. R. Roy, H. Nagai, A. Ingmundson, X. Cheng, and R. A. Kahn. 2005. The structure of RalF, an ADP-ribosylation factor guanine nucleotide exchange factor from Legionella pneumophila, reveals the presence of a cap over the active site. J. Biol. Chem. 280:1392-1400.
6. Anderson, L. B., A. V. Hertzel, and A. Das. 1996. Agrobacterium tumefaciens VirB7 and VirB9 form a disulfide-linked protein complex. Proc. Natl. Acad. Sci. USA 93:8889-8894.
7. Andersson, S. G., A. Zomorodipour, J. O. Andersson, T. Sicheritz-Ponten, U. C. Alsmark, R. M. Podowski, A. K. Naslund, A. S. Eriksson, H. H. Winkler, and C. G. Kurland. 1998. The genome sequence of Rickettsia prowazekii and the origin of mitochondria. Nature 396:133-140.
8. Aras, R. A., W. Fischer, G. I. Perez-Perez, M. Crosatti, T. Ando, R. Haas, and M. J. Blaser. 2003. Plasticity of repetitive DNA sequences within a bacterial (type IV) secretion system component. J. Exp. Med. 198:1349-1360.
9. Arechaga, I., A. Pena, S. Zunzunegui, M. del Carmen Fernandez-Alonso, G. Rivas, and F. de la Cruz. 2008. ATPase activity and oligomeric state of TrwK, the VirB4 homologue of the plasmid R388 type IV secretion system. J. Bacteriol. 190:5472-5479.
10. Atmakuri, K., E. Cascales, O. T. Burton, L. M. Banta, and P. J. Christie. 2007. Agrobacterium ParA/MinD-like VirC1 spatially coordinates early conjugative DNA transfer reactions. EMBO J. 26:2540-2551.
11. Atmakuri, K., E. Cascales, and P. J. Christie. 2004. Energetic components VirD4, VirB11 and VirB4 mediate early DNA transfer reactions required for bacterial type IV secretion. Mol. Microbiol. 54:1199-1211.
12. Atmakuri, K., Z. Ding, and P. J. Christie. 2003. VirE2, a type IV secretion substrate, interacts with the VirD4 transfer protein at cell poles of Agrobacterium tumefaciens. Mol. Microbiol. 49:1699-1713.
13. Auchtung, J. M., C. A. Lee, K. L. Garrison, and A. D. Grossman. 2007. Identification and characterization of the immunity repressor (ImmR) that controls the mobile genetic element ICEBs1 of Bacillus subtilis. Mol. Microbiol. 64:1515-1528.
14. Auchtung, J. M., C. A. Lee, R. E. Monson, A. P. Lehman, and A. D. Grossman. 2005. Regulation of a Bacillus subtilis mobile genetic element by intercellular signaling and the global DNA damage response. Proc. Natl. Acad. Sci. USA 102:12554-12559.
15. Audette, G. F., J. Manchak, P. Beatty, W. A. Klimke, and L. S. Frost. 2007. Entry exclusion in F-like plasmids requires intact TraG in the donor that recognizes its cognate TraS in the recipient. Microbiology 153:442-451.
16. Babic, A., A. B. Lindner, M. Vulic, E. J. Stewart, and M. Radman. 2008. Direct visualization of horizontal gene transfer. Science 319:1533-1536.
17. Backert, S., R. Fronzes, and G. Waksman. 2008. VirB2 and VirB5 proteins: specialized adhesins in bacterial type-IV secretion systems? Trends Microbiol. 16:409-413.
18. Backert, S., and T. F. Meyer. 2006. Type IV secretion systems and their effectors in bacterial pathogenesis. Curr. Opin. Microbiol. 9:207-217.
19. Backert, S., and M. Selbach. 2008. Role of type IV secretion in Helicobacter pylori pathogenesis. Cell. Microbiol. 10:1573-1581.
20. Bai, X., T. Fazzolari, and S. A. Hogenhout. 2004. Identification and characterization of traE genes of Spiroplasma kunkelii. Gene 336:81-91.
21. Bailey, S., D. Ward, R. Middleton, J. G. Grossmann, and P. C. Zambryski. 2006. Agrobacterium tumefaciens VirB8 structure reveals potential proteinprotein interaction sites. Proc. Natl. Acad. Sci. USA 103:2582-2587.
22. Bao, W., Y. Kumagai, H. Niu, M. Yamaguchi, K. Miura, and Y. Rikihisa. 2009. Four VirB6 paralogs and VirB9 are expressed and interact in Ehrlichia chaffeensis-containing vacuoles. J. Bacteriol. 191:278-286.
23. Bardill, J. P., J. L. Miller, and J. P. Vogel. 2005. IcmS-dependent translocation of SdeA into macrophages by the Legionella pneumophila type IV secretion system. Mol. Microbiol. 56:90-103.
24. Baron, C., and B. Coombes. 2007. Targeting bacterial secretion systems: benefits of disarmament in the microcosm. Infect. Disord. Drug Targets 7:19-27.
25. Barroso, G., and J. Labarere. 1988. Chromosomal gene transfer in Spiroplasma citri. Science 241:959-961.
26. Bayliss, R., R. Harris, L. Coutte, A. Monier, R. Fronzes, P. J. Christie, P. C. Driscoll, and G. Waksman. 2007. NMR structure of a complex between the VirB9/VirB7 interaction domains of the pKM101 type IV secretion system. Proc. Natl. Acad. Sci. USA 104:1673-1678.
27. Beare, P. A., D. Howe, D. C. Cockrell, A. Omsland, B. Hansen, and R. A. Heinzen. 2009. Characterization of a Coxiella burnetii ftsZ mutant generated by Himar1 transposon mutagenesis. J. Bacteriol. 191:1369-1381.
28. Beranek, A., M. Zettl, K. Lorenzoni, A. Schauer, M. Manhart, and G. Koraimann. 2004. Thirty-eight C-terminal amino acids of the coupling protein TraD of the F-like conjugative resistance plasmid R1 are required and sufficient to confer binding to the substrate selector protein TraM. J. Bacteriol. 186:6999-7006. (Pubitemid 39332138)
29. Berger, B. R., and P. J. Christie. 1994. Genetic complementation analysis of the Agrobacterium tumefaciens virB operon: virB2 through virB11 are essential virulence genes. J. Bacteriol. 176:3646-3660.
30. Bigot, S., V. Sivanathan, C. Possoz, F. X. Barre, and F. Cornet. 2007. FtsK, a literate chromosome segregation machine. Mol. Microbiol. 64:1434-1441.
31. Boer, R., S. Russi, A. Guasch, M. Lucas, A. G. Blanco, R. Perez-Luque, M. Coll, and F. de la Cruz. 2006. Unveiling the molecular mechanism of a conjugative relaxase: the structure of TrwC complexed with a 27-mer DNA comprising the recognition hairpin and the cleavage site. J. Mol. Biol. 358:857-869.
32. Bohne, J., A. Yim, and A. N. Binns. 1998. The Ti plasmid increases the efficiency of Agrobacterium tumefaciens as a recipient in virB-mediated conjugal transfer of an IncQ plasmid. Proc. Natl. Acad. Sci. USA 95:7057-7062.
33. Bonheyo, G., D. Graham, N. B. Shoemaker, and A. A. Salyers. 2001. Transfer region of a Bacteroides conjugative transposon, CTnDOT. Plasmid 45: 41-51.
34. Boschiroli, M. L., S. Ouahrani-Bettache, V. Foulongne, S. Michaux-Charachon, G. Bourg, A. Allardet-Servent, C. Cazevieille, J. P. Lavigne, J. P. Liautard, M. Ramuz, and D. O'Callaghan. 2002. Type IV secretion and Brucella virulence. Vet. Microbiol. 90:341-348.
35. Bourg, G., R. Sube, D. O'Callaghan, and G. Patey. 2009. Interactions between Brucella suis VirB8 and its homolog TraJ from the plasmid pSB102 underline the dynamic nature of type IV secretion systems. J. Bacteriol. 191:2985-2992.
36. Bowie, J. U., and R. T. Sauer. 1990. TraY proteins of F and related episomes are members of the Arc and Mnt repressor family. J. Mol. Biol. 211:5-6.
37. Brochet, M., V. Da Cunha, E. Couve, C. Rusniok, P. Trieu-Cuot, and P. Glaser. 2009. Atypical association of DDE transposition with conjugation specifies a new family of mobile elements. Mol. Microbiol. 71:948-959.
38. Brochet, M., C. Rusniok, E. Couve, S. Dramsi, C. Poyart, P. Trieu-Cuot, F. Kunst, and P. Glaser. 2008. Shaping a bacterial genome by large chromosomal replacements, the evolutionary history of Streptococcus agalactiae. Proc. Natl. Acad. Sci. USA 105:15961-15966.
39. Burns, D. L. 2003. Type IV transporters of pathogenic bacteria. Curr. Opin. Microbiol. 6:29-34.
40. Burrus, V., G. Pavlovic, B. Decaris, and G. Guedon. 2002. Conjugative transposons: the tip of the iceberg. Mol. Microbiol. 46:601-610.
41. Burrus, V., and M. K. Waldor. 2004. Shaping bacterial genomes with integrative and conjugative elements. Res. Microbiol. 155:376-386.
42. Cabezon, E., J. I. Sastre, and F. de la Cruz. 1997. Genetic evidence of a coupling role for the TraG protein family in bacterial conjugation. Mol. Gen. Genet. 254:400-406.
43. Calcutt, M. J., M. S. Lewis, and K. S. Wise. 2002. Molecular genetic analysis of ICEF, an integrative conjugal element that is present as a repetitive sequence in the chromosome of Mycoplasma fermentans PG18. J. Bacteriol. 184:6929-6941.
44. Cambronne, E. D., and C. R. Roy. 2007. The Legionella pneumophila IcmSW complex interacts with multiple Dot/Icm effectors to facilitate type IV translocation. PLoS Pathog. 3:e188.
45. Carle, A., C. Hoppner, K. A. Aly, Q. Yuan, A. den Dulk-Ras, A. Vergunst, D. O'Callaghan, and C. Baron. 2006. The Brucella suis type IV secretion system assembles in the cell envelope of the heterologous host Agrobacterium tumefaciens and increases IncQ plasmid pLS1 recipient competence. Infect. Immun. 74:108-117.
46. Caryl, J. A., M. C. Smith, and C. D. Thomas. 2004. Reconstitution of a staphylococcal plasmid-protein relaxation complex in vitro. J. Bacteriol. 186:3374-3383.
47. Caryl, J. A., and C. D. Thomas. 2006. Investigating the basis of substrate recognition in the pC221 relaxosome. Mol. Microbiol. 60:1302-1318.
48. Cascales, E., and P. J. Christie. 2004. Agrobacterium VirB10, an ATP energy sensor required for type IV secretion. Proc. Natl. Acad. Sci. USA 101:17228-17233.
49. Cascales, E., and P. J. Christie. 2004. Definition of a bacterial type IV secretion pathway for a DNA substrate. Science 304:1170-1173. (Pubitemid 38661856)
50. Cascales, E., and P. J. Christie. 2003. The versatile bacterial type IV secretion systems. Nat. Rev. Microbiol. 1:137-150.
51. Celli, J., and J. P. Gorvel. 2004. Organelle robbery: Brucella interactions with the endoplasmic reticulum. Curr. Opin. Microbiol. 7:93-97.
52. Cesar, C. E., C. Machon, F. de la Cruz, and M. Llosa. 2006. A new domain of conjugative relaxase TrwC responsible for efficient oriT-specific recombination on minimal target sequences. Mol. Microbiol. 62:984-996.
53. Cevallos, M. A., R. Cervantes-Rivera, and R. M. Gutierrez-Rios. 2008. The repABC plasmid family. Plasmid 60:19-37.
54. Chen, I., P. J. Christie, and D. Dubnau. 2005. The ins and outs of DNA transfer in bacteria. Science 310:1456-1460. (Pubitemid 41746335)
55. Chen, I., and D. Dubnau. 2003. DNA transport during transformation. Front. Biosci. 8:S544-S556.
56. Chen, I., R. Provvedi, and D. Dubnau. 2006. A macromolecular complex formed by a pilin-like protein in competent Bacillus subtilis. J. Biol. Chem. 281:21720-21727.
57. Chen, J., M. Reyes, M. Clarke, and H. A. Shuman. 2007. Host cell-dependent secretion and translocation of the LepA and LepB effectors of Legionella pneumophila. Cell. Microbiol. 9:1660-1671.
58. Chen, Y., J. H. Staddon, and G. M. Dunny. 2007. Specificity determinants of conjugative DNA processing in the Enterococcus faecalis plasmid pCF10 and the Lactococcus lactis plasmid pRS01. Mol. Microbiol. 63:1549-1564.
59. Chen, Y., X. Zhang, D. Manias, H. J. Yeo, G. M. Dunny, and P. J. Christie. 2008. Enterococcus faecalis PcfC, a spatially localized substrate receptor for type IV secretion of the pCF10 transfer intermediate. J. Bacteriol. 190: 3632-3645.
60. Cheng, Z., X. Wang, and Y. Rikihisa. 2008. Regulation of type IV secretion apparatus genes during Ehrlichia chaffeensis intracellular development by a previously unidentified protein. J. Bacteriol. 190:2096-2105.
61. Cho, N. H., H. R. Kim, J. H. Lee, S. Y. Kim, J. Kim, S. Cha, S. Y. Kim, A. C. Darby, H. H. Fuxelius, J. Yin, J. H. Kim, J. Kim, S. J. Lee, Y. S. Koh, W. J. Jang, K. H. Park, S. G. Andersson, M. S. Choi, and I. S. Kim. 2007. The Orientia tsutsugamushi genome reveals massive proliferation of conjugative type IV secretion system and host-cell interaction genes. Proc. Natl. Acad. Sci. USA 104:7981-7986.
62. Christie, P. J. 2004. Bacterial type IV secretion: the Agrobacterium VirB/D4 and related conjugation systems. Biochim. Biophys. Acta 1694:219-234.
63. Christie, P. J., K. Atmakuri, V. Krishnamoorthy, S. Jakubowski, and E. Cascales. 2005. Biogenesis, architecture, and function of bacterial type IV secretion systems. Annu. Rev. Microbiol. 59:451-485.
64. Christie, P. J., and E. Cascales. 2005. Structural and dynamic properties of bacterial type IV secretion systems. Mol. Membr. Biol. 22:51-61.
65. Chuang, O. N., P. M. Schlievert, C. L. Wells, D. A. Manias, T. J. Tripp, and G. M. Dunny. 2009. Multiple functional domains of Enterococcus faecalis aggregation substance Asc10 contribute to endocarditis virulence. Infect. Immun. 77:539-548.
66. Clarke, M., L. Maddera, R. L. Harris, and P. M. Silverman. 2008. F-pili dynamics by live-cell imaging. Proc. Natl. Acad. Sci. USA 105:17978-17981.
67. Clewell, D. B. 2007. Properties of Enterococcus faecalis plasmid pAD1, a member of a widely disseminated family of pheromone-responding, conjugative, virulence elements encoding cytolysin. Plasmid 58:205-227.
68. Clewell, D. B., S. E. Flannagan, and D. D. Jaworski. 1995. Unconstrained bacterial promiscuity: the Tn916-Tn1545 family of conjugative transposons. Trends Microbiol. 3:229-236.
69. Coros, A., B. Callahan, E. Battaglioli, and K. M. Derbyshire. 2008. The specialized secretory apparatus ESX-1 is essential for DNA transfer in Mycobacterium smegmatis. Mol. Microbiol. 69:794-808.
70. Couturier, M. R., E. Tasca, C. Montecucco, and M. Stein. 2006. Interaction with CagF is required for translocation of CagA into the host via the Helicobacter pylori type IV secretion system. Infect. Immun. 74:273-281.
71. Dang, T. A., and P. J. Christie. 1997. The VirB4 ATPase of Agrobacterium tumefaciens is a cytoplasmic membrane protein exposed at the periplasmic surface. J. Bacteriol. 179:453-462.
72. Datta, S., C. Larkin, and J. F. Schildbach. 2003. Structural insights into single-stranded DNA binding and cleavage by F factor TraI. Structure 11:1369-1379. (Pubitemid 37412419)
73. Davis, R. E., E. L. Dally, R. Jomantiene, Y. Zhao, B. Roe, S. Lin, and J. Shao. 2005. Cryptic plasmid pSKU146 from the wall-less plant pathogen Spiroplasma kunkelii encodes an adhesin and components of a type IV translocation-related conjugation system. Plasmid 53:179-190.
74. Degtyar, E., T. Zusman, M. Ehrlich, and G. Segal. 2009. A Legionella effector acquired from protozoa is involved in sphingolipids metabolism and is targeted to the host cell mitochondria. Cell. Microbiol. 11:1219-1235.
75. Dehio, C. 2008. Infection-associated type IV secretion systems of Bartonella and their diverse roles in host cell interaction. Cell. Microbiol. 10:1591-1598.
76. de Jong, M. F., Y. H. Sun, A. B. den Hartigh, J. M. van Dijl, and R. M. Tsolis. 2008. Identification of VceA and VceC, two members of the VjbR regulon that are translocated into macrophages by the Brucella type IV secretion system. Mol. Microbiol. 70:1378-1396.
77. Deng, W., L. Chen, W. T. Peng, X. Liang, S. Sekiguchi, M. P. Gordon, L. Comai, and E. W. Nester. 1999. VirE1 is a specific molecular chaperone for the exported single-stranded-DNA-binding protein VirE2 in Agrobacterium. Mol. Microbiol. 31:1795-1807. (Pubitemid 29151278)
78. den Hartigh, A. B., H. G. Rolan, M. F. de Jong, and R. M. Tsolis. 2008. VirB3 to VirB6 and VirB8 to VirB11, but not VirB7, are essential for mediating persistence of Brucella in the reticuloendothelial system. J. Bacteriol. 190:4427-4436.
79. de Paz, H. D., F. J. Sangari, S. Bolland, J. M. Garcia-Lobo, C. Dehio, F. de la Cruz, and M. Llosa. 2005. Functional interactions between type IV secretion systems involved in DNA transfer and virulence. Microbiology 151:3505-3516. (Pubitemid 41632064)
80. DiGiuseppe Champion, P. A., and J. S. Cox. 2007. Protein secretion systems in mycobacteria. Cell. Microbiol. 9:1376-1384.
81. Ding, Z., K. Atmakuri, and P. J. Christie. 2003. The outs and ins of bacterial type IV secretion substrates. Trends Microbiol. 11:527-535. (Pubitemid 37357206)
82. Disque-Kochem, C., and B. Dreiseikelmann. 1997. The cytoplasmic DNAbinding protein TraM binds to the inner membrane protein TraD in vitro. J. Bacteriol. 179:6133-6137. (Pubitemid 27419040)
83. Dorman, C. J., and K. A. Kane. 2009. DNA bridging and antibridging: a role for bacterial nucleoid-associated proteins in regulating the expression of laterally acquired genes. FEMS Microbiol. Rev. 33:587-592.
84. Draper, O., C. E. Cesar, C. Machon, F. de la Cruz, and M. Llosa. 2005. Site-specific recombinase and integrase activities of a conjugative relaxase in recipient cells. Proc. Natl. Acad. Sci. USA 102:16385-16390.
85. Driessen, A. J., and N. Nouwen. 2008. Protein translocation across the bacterial cytoplasmic membrane. Annu. Rev. Biochem. 77:643-667.
86. Dunny, G. M. 2007. The peptide pheromone-inducible conjugation system of Enterococcus faecalis plasmid pCF10: cell-cell signalling, gene transfer, complexity and evolution. Philos. Trans. R. Soc. Lond. B Biol. Sci. 362: 1185-1193.
87. Durrenberger, M. B., W. Villiger, and T. Bachi. 1991. Conjugational junctions: morphology of specific contacts in conjugating Escherichia coli bacteria. J. Struct. Biol. 107:146-156.
88. Dym, O., S. Albeck, T. Unger, J. Jacobovitch, A. Branzburg, Y. Michael, D. Frenkiel-Krispin, S. G. Wolf, and M. Elbaum. 2008. Crystal structure of the Agrobacterium virulence complex VirE1-VirE2 reveals a flexible protein that can accommodate different partners. Proc. Natl. Acad. Sci. USA 105: 11170-11175.
89. Economou, A., P. J. Christie, R. C. Fernandez, T. Palmer, G. V. Plano, and A. P. Pugsley. 2006. Secretion by numbers: protein traffic in prokaryotes. Mol. Microbiol. 62:308-319.
90. Eisenbrandt, R., M. Kalkum, R. Lurz, and E. Lanka. 2000. Maturation of IncP pilin precursors resembles the catalytic dyad-like mechanism of leader peptidases. J. Bacteriol. 182:6751-6761. (Pubitemid 32249259)
91. Erauso, G., K. M. Stedman, H. J. van de Werken, W. Zillig, and J. van der Oost. 2006. Two novel conjugative plasmids from a single strain of Sulfolobus. Microbiology 152:1951-1968. (Pubitemid 44070453)
92. Farias, M. E., E. Grohmann, and M. Espinosa. 1999. Expression of the mobM gene of the streptococcal plasmid pMV158 in Lactococcus lactis subsp. lactis. FEMS Microbiol. Lett. 176:403-410.
93. Farizo, K. M., T. G. Cafarella, and D. L. Burns. 1996. Evidence for a ninth gene, ptlI, in the locus encoding the pertussis toxin secretion system of Bordetella pertussis and formation of a PtlI-PtlF complex. J. Biol. Chem. 271:31643-31649.
94. Felek, S., H. Huang, and Y. Rikihisa. 2003. Sequence and expression analysis of virB9 of the type IV secretion system of Ehrlichia canis strains in ticks, dogs, and cultured cells. Infect. Immun. 71:6063-6067.
95. Felix, C., S. Pichon, C. Braquart-Varnier, H. Braig, L. Chen, R. A. Garrett, G. Martin, and P. Greve. 2008. Characterization and transcriptional analysis of two gene clusters for type IV secretion machinery in Wolbachia of Armadillidium vulgare. Res. Microbiol. 159:481-485.
96. Fernandez, D., T. A. Dang, G. M. Spudich, X. R. Zhou, B. R. Berger, and P. J. Christie. 1996. The Agrobacterium tumefaciens virB7 gene product, a proposed component of the T-complex transport apparatus, is a membraneassociated lipoprotein exposed at the periplasmic surface. J. Bacteriol. 178:3156-3167.
97. Fischer, W., R. Haas, and S. Odenbreit. 2002. Type IV secretion systems in pathogenic bacteria. Int. J. Med. Microbiol. 292:159-168.
98. Flint, J. L., J. C. Kowalski, P. K. Karnati, and K. M. Derbyshire. 2004. The RD1 virulence locus of Mycobacterium tuberculosis regulates DNA transfer in Mycobacterium smegmatis. Proc. Natl. Acad. Sci. USA 101:12598-12603.
99. Flower, A. M. 2007. The SecY translocation complex: convergence of genetics and structure. Trends Microbiol. 15:203-210.
100. Francia, M. V., A. Varsaki, M. P. Garcillan-Barcia, A. Latorre, C. Drainas, and F. de la Cruz. 2004. A classification scheme for mobilization regions of bacterial plasmids. FEMS Microbiol. Rev. 28:79-100.
101. Frenkiel-Krispin, D., S. G. Wolf, S. Albeck, T. Unger, Y. Peleg, J. Jacobovitch, Y. Michael, S. Daube, M. Sharon, C. V. Robinson, D. I. Svergun, D. Fass, T. Tzfira, and M. Elbaum. 2007. Plant transformation by Agrobacterium tumefaciens: modulation of single-stranded DNA-VirE2 complex assembly by VirE1. J. Biol. Chem. 282:3458-3464.
102. Fronzes, R., E. Schafer, L. Wang, H. R. Saibil, E. V. Orlova, and G. Waksman. 2009. Structure of a type IV secretion system core complex. Science 323:266-268.
103. Fuxelius, H. H., A. Darby, C. K. Min, N. H. Cho, and S. G. Andersson. 2007. The genomic and metabolic diversity of Rickettsia. Res. Microbiol. 158:745-753.
104. Garcillan-Barcia, M. P., M. V. Francia, and F. de la Cruz. 2009. The diversity of conjugative relaxases and its application in plasmid classification. FEMS Microbiol. Rev. 33:657-687.
105. Garcillán-Barcia, M. P., P. Jurado, B. Gonzalez-Pérez, G. Moncalián, L. A. Fernández, and F. de la Cruz. 2007. Conjugative transfer can be inhibited by blocking relaxase activity within recipient cells with intrabodies. Mol. Microbiol. 63:404-416.
106. Gillespie, J. J., N. C. Ammerman, S. M. Dreher-Lesnick, M. S. Rahman, M. J. Worley, J. C. Setubal, B. S. Sobral, and A. F. Azad. 2009. An anomalous type IV secretion system in Rickettsia is evolutionarily conserved. PLoS ONE 4:e4833.
107. Gillespie, J. J., M. S. Beier, M. S. Rahman, N. C. Ammerman, J. M. Shallom, A. Purkayastha, B. S. Sobral, and A. F. Azad. 2007. Plasmids and rickettsial evolution: insight from Rickettsia felis. PLoS ONE 2:e266.
108. Gillespie, J. J., K. Williams, M. Shukla, E. E. Snyder, E. K. Nordberg, S. M. Ceraul, C. Dharmanolla, D. Rainey, J. Soneja, J. M. Shallom, N. D. Vishnubhat, R. Wattam, A. Purkayastha, M. Czar, O. Crasta, J. C. Setubal, A. F. Azad, and B. S. Sobral. 2008. Rickettsia phylogenomics: unwinding the intricacies of obligate intracellular life. PLoS ONE 3:e2018.
109. Gilmour, M. W., T. D. Lawley, M. M. Rooker, P. J. Newnham, and D. E. Taylor. 2001. Cellular location and temperature-dependent assembly of IncHI1 plasmid R27-encoded TrhC-associated conjugative transfer protein complexes. Mol. Microbiol. 42:705-715.
110. Gomis-Ruth, F. X., and M. Coll. 2006. Cut and move: protein machinery for DNA processing in bacterial conjugation. Curr. Opin. Struct. Biol. 16:744-752.
111. Gomis-Ruth, F. X., and M. Coll. 2001. Structure of TrwB, a gatekeeper in bacterial conjugation. Int. J. Biochem. Cell Biol. 33:839-843.
112. Gomis-Ruth, F. X., F. de la Cruz, and M. Coll. 2002. Structure and role of coupling proteins in conjugal DNA transfer. Res. Microbiol. 153:199-204.
113. Gomis-Ruth, F. X., G. Moncalian, R. Perez-Luque, A. Gonzalez, E. Cabezon, F. de la Cruz, and M. Coll. 2001. The bacterial conjugation protein TrwB resembles ring helicases and F1-ATPase. Nature 409:637-641.
114. Gomis-Ruth, F. X., M. Sola, F. de la Cruz, and M. Coll. 2004. Coupling factors in macromolecular type-IV secretion machineries. Curr. Pharm. Des. 10:1551-1565.
115. Greve, B., S. Jensen, K. Brugger, W. Zillig, and R. A. Garrett. 2004. Genomic comparison of archaeal conjugative plasmids from Sulfolobus. Archaea 1:231-239. (Pubitemid 39520704)
116. Grohmann, E., G. Muth, and M. Espinosa. 2003. Conjugative plasmid transfer in gram-positive bacteria. Microbiol. Mol. Biol. Rev. 67:277-301.
117. Guasch, A., M. Lucas, G. Moncalian, M. Cabezas, R. Perez-Luque, F. X. Gomis-Ruth, F. de la Cruz, and M. Coll. 2003. Recognition and processing of the origin of transfer DNA by conjugative relaxase TrwC. Nat. Struct. Biol. 10:1002-1010.
118. Gunton, J. E., M. W. Gilmour, K. P. Baptista, T. D. Lawley, and D. E. Taylor. 2007. Interaction between the co-inherited TraG coupling protein and the TraJ membrane-associated protein of the H-plasmid conjugative DNA transfer system resembles chromosomal DNA translocases. Microbiology 153:428-441.
119. Guo, M., S. Jin, D. Sun, C. L. Hew, and S. Q. Pan. 2007. Recruitment of conjugative DNA transfer substrate to Agrobacterium type IV secretion apparatus. Proc. Natl. Acad. Sci. USA 104:20019-20024.
120. Guzman, L. M., and M. Espinosa. 1997. The mobilization protein, MobM, of the streptococcal plasmid pMV158 specifically cleaves supercoiled DNA at the plasmid oriT. J. Mol. Biol. 266:688-702.
121. Hamilton, C. M., H. Lee, P. L. Li, D. M. Cook, K. R. Piper, S. B. von Bodman, E. Lanka, W. Ream, and S. K. Farrand. 2000. TraG from RP4 and TraG and VirD4 from Ti plasmids confer relaxosome specificity to the conjugal transfer system of pTiC58. J. Bacteriol. 182:1541-1548.
122. Hamilton, H. L., N. M. Dominguez, K. J. Schwartz, K. T. Hackett, and J. P. Dillard. 2005. Neisseria gonorrhoeae secretes chromosomal DNA via a novel type IV secretion system. Mol. Microbiol. 55:1704-1721.
123. Hare, S., R. Bayliss, C. Baron, and G. Waksman. 2006. A large domain swap in the VirB11 ATPase of Brucella suis leaves the hexameric assembly intact. J. Mol. Biol. 360:56-66.
124. Haren, L., B. Ton-Hoang, and M. Chandler. 1999. Integrating DNA: transposases and retroviral integrases. Annu. Rev. Microbiol. 53:245-281.
125. Hazes, B., and L. Frost. 2008. Towards a systems biology approach to study type II/IV secretion systems. Biochim. Biophys. Acta 1778:1839-1850.
126. Hickman, A. B., D. R. Ronning, Z. N. Perez, R. M. Kotin, and F. Dyda. 2004. The nuclease domain of adeno-associated virus rep coordinates replication initiation using two distinct DNA recognition interfaces. Mol. Cell 13:403-414.
127. Hirt, H., D. A. Manias, E. M. Bryan, J. R. Klein, J. K. Marklund, J. H. Staddon, M. L. Paustian, V. Kapur, and G. M. Dunny. 2005. Characterization of the pheromone response of the Enterococcus faecalis conjugative plasmid pCF10: complete sequence and comparative analysis of the transcriptional and phenotypic responses of pCF10-containing cells to pheromone induction. J. Bacteriol. 187:1044-1054.
128. Hodges, L. D., A. C. Vergunst, J. Neal-McKinney, A. den Dulk-Ras, D. M. Moyer, P. J. Hooykaas, and W. Ream. 2006. Agrobacterium rhizogenes GALLS protein contains domains for ATP binding, nuclear localization, and type IV secretion. J. Bacteriol. 188:8222-8230.
129. Hofreuter, D., A. Karnholz, and R. Haas. 2003. Topology and membrane interaction of Helicobacter pylori ComB proteins involved in natural transformation competence. Int. J. Med. Microbiol. 293:153-165.
130. Hofreuter, D., S. Odenbreit, and R. Haas. 2001. Natural transformation competence in Helicobacter pylori is mediated by the basic components of a type IV secretion system. Mol. Microbiol. 41:379-391.
131. Hofreuter, D., S. Odenbreit, G. Henke, and R. Haas. 1998. Natural competence for DNA transformation in Helicobacter pylori: identification and genetic characterization of the comB locus. Mol. Microbiol. 28:1027-1038.
132. Hohlfeld, S., I. Pattis, J. Puls, G. V. Plano, R. Haas, and W. Fischer. 2006. A C-terminal translocation signal is necessary, but not sufficient for type IV secretion of the Helicobacter pylori CagA protein. Mol. Microbiol. 59:1624-1637.
133. Hoppner, C., Z. Liu, N. Domke, A. N. Binns, and C. Baron. 2004. VirB1 orthologs from Brucella suis and pKM101 complement defects of the lytic transglycosylase required for efficient type IV secretion from Agrobacterium tumefaciens. J. Bacteriol. 186:1415-1422.
134. Hormaeche, I., I. Iloro, J. L. Arrondo, F. M. Goni, F. de la Cruz, and I. Alkorta. 2004. Role of the transmembrane domain in the stability of TrwB, an integral protein involved in bacterial conjugation. J. Biol. Chem. 279: 10955-10961.
135. Hormaeche, I., R. L. Segura, A. J. Vecino, F. M. Goni, F. de la Cruz, and I. Alkorta. 2006. The transmembrane domain provides nucleotide binding specificity to the bacterial conjugation protein TrwB. FEBS Lett. 580:3075-3082.
136. Hotopp, J. C., M. Lin, R. Madupu, J. Crabtree, S. V. Angiuoli, J. A. Eisen, R. Seshadri, Q. Ren, M. Wu, T. R. Utterback, S. Smith, M. Lewis, H. Khouri, C. Zhang, H. Niu, Q. Lin, N. Ohashi, N. Zhi, W. Nelson, L. M. Brinkac, R. J. Dodson, M. J. Rosovitz, J. Sundaram, S. C. Daugherty, T. Davidsen, A. S. Durkin, M. Gwinn, D. H. Haft, J. D. Selengut, S. A. Sullivan, N. Zafar, L. Zhou, F. Benahmed, H. Forberger, R. Halpin, S. Mulligan, J. Robinson, O. White, Y. Rikihisa, and H. Tettelin. 2006. Comparative genomics of emerging human ehrlichiosis agents. PLoS Genet. 2:e21.
137. Hu, X., G. Van der Auwera, S. Timmery, L. Zhu, and J. Mahillon. 2009. Distribution, diversity, and potential mobility of extrachromosomal elements related to the Bacillus anthracis pXO1 and pXO2 virulence plasmids. Appl. Environ. Microbiol. 75:3016-3028.
138. Iannelli, F., M. R. Oggioni, and G. Pozzi. 2002. Allelic variation in the highly polymorphic locus pspC of Streptococcus pneumoniae. Gene 284: 63-71.
139. Ilyina, T. V., and E. V. Koonin. 1992. Conserved sequence motifs in the initiator proteins for rolling circle DNA replication encoded by diverse replicons from eubacteria, eucaryotes and archaebacteria. Nucleic Acids Res. 20:3279-3285.
140. Ishmael, N., J. C. D. Hotopp, P. Ioannidis, S. Biber, J. Sakamoto, S. Siozios, V. Nene, J. Werren, K. Bourtzis, S. R. Bordenstein, and H. Tettelin. 2009. Extensive genomic diversity of closely related Wolbachia strains. Microbiology 9:1252-1266.
141. Jackson, C. R., J. A. Boylan, J. G. Frye, and F. C. Gherardini. 2007. Evidence of a conjugal erythromycin resistance element in the Lyme disease spirochete Borrelia burgdorferi. Int. J. Antimicrob. Agents 30:496-504.
142. Jakubowski, S. J., E. Cascales, V. Krishnamoorthy, and P. J. Christie. 2005. Agrobacterium tumefaciens VirB9, an outer-membrane-associated component of a type IV secretion system, regulates substrate selection and T-pilus biogenesis. J. Bacteriol. 187:3486-3495.
143. Jakubowski, S. J., J. E. Kerr, I. Garza, V. Krishnamoorthy, R. Bayliss, G. Waksman, and P. J. Christie. 2009. Agrobacterium VirB10 domain requirements for type IV secretion and T pilus biogenesis. Mol. Microbiol. 71:779-794.
144. Jakubowski, S. J., V. Krishnamoorthy, E. Cascales, and P. J. Christie. 2004. Agrobacterium tumefaciens VirB6 domains direct the ordered export of a DNA substrate through a type IV secretion system. J. Mol. Biol. 341:961-977.
145. Jakubowski, S. J., V. Krishnamoorthy, and P. J. Christie. 2003. Agrobacterium tumefaciens VirB6 protein participates in formation of VirB7 and VirB9 complexes required for type IV secretion. J. Bacteriol. 185:2867-2878.
146. Jones, A. L., K. Shirasu, and C. I. Kado. 1994. The product of the virB4 gene of Agrobacterium tumefaciens promotes accumulation of VirB3 protein. J. Bacteriol. 176:5255-5261.
147. Joshi, B. D., M. Berg, J. Rogers, J. Fletcher, and U. Melcher. 2005. Sequence comparisons of plasmids pBJS-O of Spiroplasma citri and pSKU146 of S. kunkelii: implications for plasmid evolution. BMC Genomics 6:175.
148. Judd, P. K., R. B. Kumar, and A. Das. 2005. Spatial location and requirements for the assembly of the Agrobacterium tumefaciens type IV secretion apparatus. Proc. Natl. Acad. Sci. USA 102:11498-11503.
149. Judd, P. K., R. B. Kumar, and A. Das. 2005. The type IV secretion appa- ratus protein VirB6 of Agrobacterium tumefaciens localizes to a cell pole. Mol. Microbiol. 55:115-124.
150. Juhas, M., D. W. Crook, I. D. Dimopoulou, G. Lunter, R. M. Harding, D. J. Ferguson, and D. W. Hood. 2007. Novel type IV secretion system involved in propagation of genomic islands. J. Bacteriol. 189:761-771.
151. Juhas, M., D. W. Crook, and D. W. Hood. 2008. Type IV secretion systems: tools of bacterial horizontal gene transfer and virulence. Cell. Microbiol. 10:2377-2386.
152. Juhas, M., J. R. van der Meer, M. Gaillard, R. M. Harding, D. W. Hood, and D. W. Crook. 2009. Genomic islands: tools of bacterial horizontal gene transfer and evolution. FEMS Microbiol. Rev. 33:376-393.
153. Kalkum, M., R. Eisenbrandt, and E. Lanka. 2004. Protein circlets as sex pilus subunits. Curr. Protein Pept. Sci. 5:417-424.
154. Kalkum, M., R. Eisenbrandt, R. Lurz, and E. Lanka. 2002. Tying rings for sex. Trends Microbiol. 10:382-387.
155. Karnholz, A., C. Hoefler, S. Odenbreit, W. Fischer, D. Hofreuter, and R. Haas. 2006. Functional and topological characterization of novel components of the comB DNA transformation competence system in Helicobacter pylori. J. Bacteriol. 188:882-893. (Pubitemid 43146403)
156. Kay, B. K., M. P. Williamson, and M. Sudol. 2000. The importance of being proline: the interaction of proline-rich motifs in signaling proteins with their cognate domains. FASEB J. 14:231-241.
157. Koonin, E. V., and Y. I. Wolf. 2008. Genomics of bacteria and archaea: the emerging dynamic view of the prokaryotic world. Nucleic Acids Res. 36: 6688-6719.
158. Kopec, J., A. Bergmann, G. Fritz, E. Grohmann, and W. Keller. 2005. TraA and its N-terminal relaxase domain of the gram-positive plasmid pIP501 show specific oriT binding and behave as dimers in solution. Biochem. J. 387:401-409.
159. Krause, S., M. Barcena, W. Pansegrau, R. Lurz, J. M. Carazo, and E. Lanka. 2000. Sequence-related protein export NTPases encoded by the conjugative transfer region of RP4 and by the cag pathogenicity island of Helicobacter pylori share similar hexameric ring structures. Proc. Natl. Acad. Sci. USA 97:3067-3072.
160. Krause, S., W. Pansegrau, R. Lurz, F. de la Cruz, and E. Lanka. 2000. Enzymology of type IV macromolecule secretion systems: the conjugative transfer regions of plasmids RP4 and R388 and the cag pathogenicity island of Helicobacter pylori encode structurally and functionally related nucleoside triphosphate hydrolases. J. Bacteriol. 182:2761-2770.
161. Kumar, R. B., and A. Das. 2001. Functional analysis of the Agrobacterium tumefaciens T-DNA transport pore protein VirB8. J. Bacteriol. 183:3636-3641.
162. Kumar, R. B., Y. H. Xie, and A. Das. 2000. Subcellular localization of the Agrobacterium tumefaciens T-DNA transport pore proteins: VirB8 is essential for the assembly of the transport pore. Mol. Microbiol. 36:608-617.
163. Kurenbach, B., J. Kopec, M. Magdefrau, K. Andreas, W. Keller, C. Bohn, M. Y. Abajy, and E. Grohmann. 2006. The TraA relaxase autoregulates the putative type IV secretion-like system encoded by the broad-host-range Streptococcus agalactiae plasmid pIP501. Microbiology 152:637-645.
164. Kwok, T., D. Zabler, S. Urman, M. Rohde, R. Hartig, S. Wessler, R. Misselwitz, J. Berger, N. Sewald, W. Konig, and S. Backert. 2007. Helicobacter exploits integrin for type IV secretion and kinase activation. Nature 449:862-866.
165. Lai, E. M., O. Chesnokova, L. M. Banta, and C. I. Kado. 2000. Genetic and environmental factors affecting T-pilin export and T-pilus biogenesis in relation to flagellation of Agrobacterium tumefaciens. J. Bacteriol. 182:3705-3716.
166. Larkin, C., S. Datta, A. Nezami, J. A. Dohm, and J. F. Schildbach. 2003. Crystallization and preliminary X-ray characterization of the relaxase domain of F factor TraI. Acta Crystallogr. D Biol. Crystallogr. 59:1514-1516.
167. Larkin, C., R. J. Haft, M. J. Harley, B. Traxler, and J. F. Schildbach. 2007. Roles of active site residues and the HUH motif of the F plasmid TraI relaxase. J. Biol. Chem. 282:33707-33713.
168. Lau-Wong, I. C., T. Locke, M. J. Ellison, T. L. Raivio, and L. S. Frost. 2008. Activation of the Cpx regulon destabilizes the F plasmid transfer activator, TraJ, via the HslVU protease in Escherichia coli. Mol. Microbiol. 67:516-527.
169. Lawley, T. D., W. A. Klimke, M. J. Gubbins, and L. S. Frost. 2003. F factor conjugation is a true type IV secretion system. FEMS Microbiol. Lett. 224:1-15.
170. Lee, C. A., J. M. Auchtung, R. E. Monson, and A. D. Grossman. 2007. Identification and characterization of int (integrase), xis (excisionase) and chromosomal attachment sites of the integrative and conjugative element ICEBs1 of Bacillus subtilis. Mol. Microbiol. 66:1356-1369.
171. Lee, C. A., and A. D. Grossman. 2007. Identification of the origin of transfer (oriT) and DNA relaxase required for conjugation of the integrative and conjugative element ICEBs1 of Bacillus subtilis. J. Bacteriol. 189:7254-7261.
172. Lillestol, R. K., S. A. Shah, K. Brugger, P. Redder, H. Phan, J. Christiansen, and R. A. Garrett. 2009. CRISPR families of the crenarchaeal genus Sulfolobus: bidirectional transcription and dynamic properties. Mol. Microbiol. 72:259-272.
173. Lin, M., A. den Dulk-Ras, P. J. Hooykaas, and Y. Rikihisa. 2007. Anaplasma phagocytophilum AnkA secreted by type IV secretion system is tyrosine phosphorylated by Abl-1 to facilitate infection. Cell. Microbiol. 9:2644-2657.
174. Lipps, G. 2006. Plasmids and viruses of the thermoacidophilic crenarchaeote Sulfolobus. Extremophiles 10:17-28.
175. Liu, Z., and A. N. Binns. 2003. Functional subsets of the VirB type IV transport complex proteins involved in the capacity of Agrobacterium tumefaciens to serve as a recipient in virB-mediated conjugal transfer of plasmid RSF1010. J. Bacteriol. 185:3259-3269.
176. Llosa, M., S. Zunzunegui, and F. de la Cruz. 2003. Conjugative coupling proteins interact with cognate and heterologous VirB10-like proteins while exhibiting specificity for cognate relaxosomes. Proc. Natl. Acad. Sci. USA 100:10465-10470.
177. Llosa, M., J. Zupan, C. Baron, and P. Zambryski. 2000. The N- and C-terminal portions of the Agrobacterium VirB1 protein independently enhance tumorigenesis. J. Bacteriol. 182:3437-3445.
178. Lopez, J. E., G. H. Palmer, K. A. Brayton, M. J. Dark, S. E. Leach, and W. C. Brown. 2007. Immunogenicity of Anaplasma marginale type IV secretion system proteins in a protective outer membrane vaccine. Infect. Immun. 75:2333-2342.
179. Lu, J., A. den Dulk-Ras, P. J. J. Hooykaas, and J. N. M. Glover. 2009. Agrobacterium tumefaciens VirC2 enhances T-DNA transfer and virulence through its C-terminal ribbon-helix-helix DNA-binding fold. Proc. Natl. Acad. Sci. USA. 106:9643-9648.
180. Lu, J., and L. S. Frost. 2005. Mutations in the C-terminal region of TraM provide evidence for in vivo TraM-TraD interactions during F-plasmid conjugation. J. Bacteriol. 187:4767-4773. (Pubitemid 40962194)
181. Lu, J., J. J. Wong, R. A. Edwards, J. Manchak, L. S. Frost, and J. N. Glover. 2008. Structural basis of specific TraD-TraM recognition during F plasmidmediated bacterial conjugation. Mol. Microbiol. 70:89-99.
182. Mahairas, G. G., J. A. Cao, and F. C. Minion. 1990. Genetic exchange of transposon and integrative plasmid markers in Mycoplasma pulmonis. J. Bacteriol. 172:2267-2272.
183. Marenda, M., V. Barbe, G. Gourgues, S. Mangenot, E. Sagne, and C. Citti. 2006. A new integrative conjugative element occurs in Mycoplasma agalactiae as chromosomal and free circular forms. J. Bacteriol. 188:4137-4141.
184. Marraffini, L. A., and E. J. Sontheimer. 2008. CRISPR interference limits horizontal gene transfer in staphylococci by targeting DNA. Science 322: 1843-1845.
185. Marrero, J., and M. K. Waldor. 2007. Determinants of entry exclusion within Eex and TraG are cytoplasmic. J. Bacteriol. 189:6469-6473. (Pubitemid 47378655)
186. Marrero, J., and M. K. Waldor. 2005. Interactions between inner membrane proteins in donor and recipient cells limit conjugal DNA transfer. Dev. Cell 8:963-970. (Pubitemid 40754066)
187. Massey, T. H., C. P. Mercogliano, J. Yates, D. J. Sherratt, and J. Lowe. 2006. Double-stranded DNA translocation: structure and mechanism of hexameric FtsK. Mol. Cell 23:457-469. (Pubitemid 44205486)
188. Masui, S., T. Sasaki, and H. Ishikawa. 2000. Genes for the type IV secretion system in an intracellular symbiont, Wolbachia, a causative agent of various sexual alterations in arthropods. J. Bacteriol. 182:6529-6531.
189. Meyer, R. 2009. The R1162 Mob proteins can promote conjugative transfer from cryptic origins in the bacterial chromosome. J. Bacteriol. 191:1574-1580.
190. Middleton, R., K. Sjolander, N. Krishamurthy, J. Foley, and P. Zambryski. 2005. Predicted hexameric structure of the Agrobacterium VirB4 C-terminus suggests VirB4 acts as a docking site during type IV secretion. Proc. Natl. Acad. Sci. USA 102:1685-1690.
191. Mojica, F. J., C. Diez-Villasenor, J. Garcia-Martinez, and C. Almendros. 2009. Short motif sequences determine the targets of the prokaryotic CRISPR defense system. Microbiology 155:733-740.
192. Moncalian, G., and F. de la Cruz. 2004. DNA binding properties of protein TrwA, a possible structural variant of the Arc repressor superfamily. Biochim. Biophys. Acta 1701:15-23.
193. Monzingo, A. F., A. Ozburn, S. Xia, R. J. Meyer, and J. D. Robertus. 2007. The structure of the minimal relaxase domain of MobA at 2.1 Å resolution. J. Mol. Biol. 366:165-178.
194. Nagai, H., E. D. Cambronne, J. C. Kagan, J. C. Amor, R. A. Kahn, and C. R. Roy. 2005. A C-terminal translocation signal required for Dot/Icm-dependent delivery of the Legionella RalF protein to host cells. Proc. Natl. Acad. Sci. USA 102:826-831.
195. Nagai, H., and C. R. Roy. 2003. Show me the substrates: modulation of host cell function by type IV secretion systems. Cell. Microbiol. 5:373-383.
196. Navarre, W. W., M. McClelland, S. J. Libby, and F. C. Fang. 2007. Silencing of xenogeneic DNA by H-NS-facilitation of lateral gene transfer in bacteria by a defense system that recognizes foreign DNA. Genes Dev. 21:1456-1471.
197. Navarre, W. W., S. Porwollik, Y. Wang, M. McClelland, H. Rosen, S. J. Libby, and F. C. Fang. 2006. Selective silencing of foreign DNA with low GC content by the H-NS protein in Salmonella. Science 313:236-238.
198. Navarre, W. W., and O. Schneewind. 1999. Surface proteins of grampositive bacteria and mechanisms of their targeting to the cell wall envelope. Microbiol. Mol. Biol. Rev. 63:174-229.
199. Nelson, C. M., M. J. Herron, R. F. Felsheim, B. R. Schloeder, S. M. Grindle, A. O. Chavez, T. J. Kurtti, and U. G. Munderloh. 2008. Whole genome transcription profiling of Anaplasma phagocytophilum in human and tick host cells by tiling array analysis. BMC Genomics 9:364.
200. Nguyen, K. T., K. Piastro, and K. M. Derbyshire. 2009. LpqM, a mycobacterial lipoprotein-metalloproteinase, is required for conjugal DNA transfer in Mycobacterium smegmatis. J. Bacteriol. 191:2721-2727.
201. Ninio, S., J. Celli, and C. R. Roy. 2009. A Legionella pneumophila effector protein encoded in a region of genomic plasticity binds to Dot/Icm-modi- fied vacuoles. PLoS Pathog. 5:e1000278.
202. Ninio, S., and C. R. Roy. 2007. Effector proteins translocated by Legionella pneumophila: strength in numbers. Trends Microbiol. 15:372-380.
203. Ninio, S., D. M. Zuckman-Cholon, E. D. Cambronne, and C. R. Roy. 2005. The Legionella IcmS-IcmW protein complex is important for Dot/Icmmediated protein translocation. Mol. Microbiol. 55:912-926.
204. Novick, R. P. 1987. Plasmid incompatibility. Microbiol. Rev. 51:3813-3895.
205. Ogata, H., B. La Scola, S. Audic, P. Renesto, G. Blanc, C. Robert, P. E. Fournier, J. M. Claverie, and D. Raoult. 2006. Genome sequence of Rickettsia bellii illuminates the role of amoebae in gene exchanges between intracellular pathogens. PLoS Genet. 2:e76.
206. Ogata, H., P. Renesto, S. Audic, C. Robert, G. Blanc, P. E. Fournier, H. Parinello, J. M. Claverie, and D. Raoult. 2005. The genome sequence of Rickettsia felis identifies the first putative conjugative plasmid in an obligate intracellular parasite. PLoS Biol. 3:e248.
207. Ohashi, N., N. Zhi, Q. Lin, and Y. Rikihisa. 2002. Characterization and transcriptional analysis of gene clusters for a type IV secretion machinery in human granulocytic and monocytic ehrlichiosis agents. Infect. Immun. 70: 2128-2138.
208. Omsland, A., D. C. Cockrell, D. Howe, E. R. Fischer, K. Virtaneva, D. E. Sturdevant, S. F. Porcella, and R. A. Heinzen. 2009. Host cell-free growth of the Q fever bacterium Coxiella burnetii. Proc. Natl. Acad. Sci. USA 106:4430-4434.
209. Osborn, A. M., and D. Boltner. 2002. When phage, plasmids, and transposons collide: genomic islands, and conjugative- and mobilizable- transposons as a mosaic continuum. Plasmid 48:202-212.
210. Ozbek, E., S. A. Miller, T. Meulia, and S. A. Hogenhout. 2003. Infection and replication sites of Spiroplasma kunkelii (class: Mollicutes) in midgut and Malpighian tubules of the leafhopper Dalbulus maidis. J. Invertebr. Pathol. 82:167-175. (Pubitemid 36435043)
211. Page, A. L., and C. Parsot. 2002. Chaperones of the type III secretion pathway: jacks of all trades. Mol. Microbiol. 46:1-11.
212. Pan, X., A. Luhrmann, A. Satoh, M. A. Laskowski-Arce, and C. R. Roy. 2008. Ankyrin repeat proteins comprise a diverse family of bacterial type IV effectors. Science 320:1651-1654. (Pubitemid 351931257)
213. Pansegrau, W., and E. Lanka. 1996. Enzymology of DNA transfer by conjugative mechanisms. Prog. Nucleic Acid Res. Mol. Biol. 54:197-251.
214. Parker, C., and R. J. Meyer. 2007. The R1162 relaxase/primase contains two, type IV transport signals that require the small plasmid protein MobB. Mol. Microbiol. 66:252-261.
215. Pattis, I., E. Weiss, R. Laugks, R. Haas, and W. Fischer. 2007. The Helicobacter pylori CagF protein is a type IV secretion chaperone-like molecule that binds close to the C-terminal secretion signal of the CagA effector protein. Microbiology 153:2896-2909.
216. Pettis, G. S., and S. N. Cohen. 2001. Unraveling the essential role in conjugation of the Tra protein of Streptomyces lividans plasmid pIJ101. Antonie van Leeuwenhoek 79:247-250.
217. Prangishvili, D., S. V. Albers, I. Holz, H. P. Arnold, K. Stedman, T. Klein, H. Singh, J. Hiort, A. Schweier, J. K. Kristjansson, and W. Zillig. 1998. Conjugation in archaea: frequent occurrence of conjugative plasmids in Sulfolobus. Plasmid 40:190-202. (Pubitemid 28527318)
218. Pulliainen, A. T., and C. Dehio. 2009. Bartonella henselae: subversion of vascular endothelial cell functions by translocated bacterial effector proteins. Int. J. Biochem. Cell Biol. 41:507-510.
219. Rabel, C., A. M. Grahn, R. Lurz, and E. Lanka. 2003. The VirB4 family of proposed traffic nucleoside triphosphatases: common motifs in plasmid RP4 TrbE are essential for conjugation and phage adsorption. J. Bacteriol. 185:1045-1058. (Pubitemid 36139464)
220. Raivio, T. L. 2005. Envelope stress responses and gram-negative bacterial pathogenesis. Mol. Microbiol. 56:1119-1128.
221. Rakita, R. M., N. N. Vanek, K. Jacques-Palaz, M. Mee, M. M. Mariscalco, G. M. Dunny, M. Snuggs, W. B. Van Winkle, and S. I. Simon. 1999. Enterococcus faecalis bearing aggregation substance is resistant to killing by human neutrophils despite phagocytosis and neutrophil activation. Infect. Immun. 67:6067-6075.
222. Rambow-Larsen, A. A., and A. A. Weiss. 2002. The PtlE protein of Bordetella pertussis has peptidoglycanase activity required for Ptl-mediated pertussis toxin secretion. J. Bacteriol. 184:2863-2869.
223. Rances, E., D. Voronin, V. Tran-Van, and P. Mavingui. 2008. Genetic and functional characterization of the type IV secretion system in Wolbachia. J. Bacteriol. 190:5020-5030. (Pubitemid 351991070)
224. Rashkova, S., G. M. Spudich, and P. J. Christie. 1997. Characterization of membrane and protein interaction determinants of the Agrobacterium tumefaciens VirB11 ATPase. J. Bacteriol. 179:583-591.
225. Roberts, M. C., and G. E. Kenny. 1987. Conjugal transfer of transposon Tn916 from Streptococcus faecalis to Mycoplasma hominis. J. Bacteriol. 169:3836-3839.
226. Rocco, J. M., and G. Churchward. 2006. The integrase of the conjugative transposon Tn916 directs strand- and sequence-specific cleavage of the origin of conjugal transfer, oriT, by the endonuclease Orf20. J. Bacteriol. 188:2207-2213.
227. Rohde, M., J. Puls, R. Buhrdorf, W. Fischer, and R. Haas. 2003. A novel sheathed surface organelle of the Helicobacter pylori cag type IV secretion system. Mol. Microbiol. 49:219-234.
228. Rosenshine, I., R. Tchelet, and M. Mevarech. 1989. The mechanism of DNA transfer in the mating system of an archaebacterium. Science 245: 1387-1389. (Pubitemid 19252606)
229. Sagulenko, V., E. Sagulenko, S. Jakubowski, E. Spudich, and P. J. Christie. 2001. VirB7 lipoprotein is exocellular and associates with the Agrobacterium tumefaciens T pilus. J. Bacteriol. 183:3642-3651.
230. Sagulenko, Y., V. Sagulenko, J. Chen, and P. J. Christie. 2001. Role of Agrobacterium VirB11 ATPase in T-pilus assembly and substrate selection. J. Bacteriol. 183:5813-5825.
231. Saillard, C., P. Carle, S. Duret-Nurbel, R. Henri, N. Killiny, S. Carrere, J. Gouzy, J. M. Bove, J. Renaudin, and X. Foissac. 2008. The abundant extrachromosomal DNA content of the Spiroplasma citri GII3-3X genome. BMC Genomics 9:195.
232. Salgado-Pabon, W., S. Jain, N. Turner, C. van der Does, and J. P. Dillard. 2007. A novel relaxase homologue is involved in chromosomal DNA processing for type IV secretion in Neisseria gonorrhoeae. Mol. Microbiol. 66:930-947. (Pubitemid 350050420)
233. Samuels, A. L., E. Lanka, and J. E. Davies. 2000. Conjugative junctions in RP4-mediated mating of Escherichia coli. J. Bacteriol. 182:2709-2715.
234. Schandel, K. A., M. M. Muller, and R. E. Webster. 1992. Localization of TraC, a protein involved in assembly of the F conjugative pilus. J. Bacteriol. 174:3800-3806.
235. Schleper, C., I. Holz, D. Janekovic, J. Murphy, and W. Zillig. 1995. A multicopy plasmid of the extremely thermophilic archaeon Sulfolobus effects its transfer to recipients by mating. J. Bacteriol. 177:4417-4426.
236. Schroder, G., S. Krause, E. L. Zechner, B. Traxler, H. J. Yeo, R. Lurz, G. Waksman, and E. Lanka. 2002. TraG-like proteins of DNA transfer systems and of the Helicobacter pylori type IV secretion system: inner membrane gate for exported substrates? J. Bacteriol. 184:2767-2779.
237. Schroder, G., and E. Lanka. 2005. The mating pair formation system of conjugative plasmids-a versatile secretion machinery for transfer of proteins and DNA. Plasmid 54:1-25. (Pubitemid 40720286)
238. Schroder, G., and E. Lanka. 2003. TraG-like proteins of type IV secretion systems: functional dissection of the multiple activities of TraG (RP4) and TrwB (R388). J. Bacteriol. 185:4371-4381.
239. Schulein, R., P. Guye, T. A. Rhomberg, M. C. Schmid, G. Schroder, A. C. Vergunst, I. Carena, and C. Dehio. 2005. A bipartite signal mediates the transfer of type IV secretion substrates of Bartonella henselae into human cells. Proc. Natl. Acad. Sci. USA 102:856-861.
240. Seth-Smith, H., and N. J. Croucher. 2009. Genome watch: breaking the ICE. Nat. Rev. Microbiol. 7:328-329.
241. Seubert, A., R. Hiestand, F. de la Cruz, and C. Dehio. 2003. A bacterial conjugation machinery recruited for pathogenesis. Mol. Microbiol. 49:1253-1266.
242. Shah, S. A., N. R. Hansen, and R. A. Garrett. 2009. Distribution of CRISPR spacer matches in viruses and plasmids of crenarchaeal acidothermophiles and implications for their inhibitory mechanism. Biochem. Soc. Trans. 37:23-28.
243. She, Q., H. Phan, R. A. Garrett, S. V. Albers, K. M. Stedman, and W. Zillig. 1998. Genetic profile of pNOB8 from Sulfolobus: the first conjugative plasmid from an archaeon. Extremophiles 2:417-425.
244. Siefert, J. L. 2009. Defining the mobilome. Methods Mol. Biol. 532:13-27.
245. Silverman, P. M. 1997. Towards a structural biology of bacterial conjugation. Mol. Microbiol. 23:423-429.
246. Simone, M., C. A. McCullen, L. E. Stahl, and A. N. Binns. 2001. The carboxy-terminus of VirE2 from Agrobacterium tumefaciens is required for its transport to host cells by the virB-encoded type IV transport system. Mol. Microbiol. 41:1283-1293.
247. Sirand-Pugnet, P., C. Lartigue, M. Marenda, D. Jacob, A. Barre, V. Barbe, C. Schenowitz, S. Mangenot, A. Couloux, B. Segurens, A. de Daruvar, A. Blanchard, and C. Citti. 2007. Being pathogenic, plastic, and sexual while living with a nearly minimal bacterial genome. PLoS Genet. 3:e75.
248. Soppa, J., A. Baumann, M. Brenneis, M. Dambeck, O. Hering, and C. Lange. 2008. Genomics and functional genomics with haloarchaea. Arch. Microbiol. 190:197-215.
249. Spudich, G. M., D. Fernandez, X. R. Zhou, and P. J. Christie. 1996. Intermolecular disulfide bonds stabilize VirB7 homodimers and VirB7/ VirB9 heterodimers during biogenesis of the Agrobacterium tumefaciens T-complex transport apparatus. Proc. Natl. Acad. Sci. USA 93:7512-7517.
250. Stedman, K. M., Q. She, H. Phan, I. Holz, H. Singh, D. Prangishvili, R. Garrett, and W. Zillig. 2000. pING family of conjugative plasmids from the extremely thermophilic archaeon Sulfolobus islandicus: insights into recombination and conjugation in crenarchaeota. J. Bacteriol. 182:7014-7020.
251. Steen, J. A., T. L. Bannam, W. L. Teng, R. J. Devenish, and J. I. Rood. 2009. The putative coupling protein TcpA interacts with other pCW3-encoded proteins to form an essential part of the conjugation complex. J. Bacteriol. 191:2926-2933.
252. Stoebel, D. M., A. Free, and C. J. Dorman. 2008. Anti-silencing: overcoming H-NS-mediated repression of transcription in gram-negative enteric bacteria. Microbiology 154:2533-2545.
253. Street, L. M., M. J. Harley, J. C. Stern, C. Larkin, S. L. Williams, D. L. Miller, J. A. Dohm, M. E. Rodgers, and J. F. Schildbach. 2003. Subdomain organization and catalytic residues of the F factor TraI relaxase domain. Biochim. Biophys. Acta 1646:86-99.
254. Szpirer, C. Y., M. Faelen, and M. Couturier. 2000. Interaction between the RP4 coupling protein TraG and the pBHR1 mobilization protein Mob. Mol. Microbiol. 37:1283-1292.
255. Szpirer, C. Y., M. Faelen, and M. Couturier. 2001. Mobilization function of the pBHR1 plasmid, a derivative of the broad-host-range plasmid pBBR1. J. Bacteriol. 183:2101-2110.
256. Tanaka, J., T. Suzuki, H. Mimuro, and C. Sasakawa. 2003. Structural definition on the surface of Helicobacter pylori type IV secretion apparatus. Cell. Microbiol. 5:395-404.
257. Tato, I., I. Matilla, I. Arechaga, S. Zunzunegui, F. de la Cruz, and E. Cabezon. 2007. The ATPase activity of the DNA transporter TrwB is modulated by protein TrwA: implications for a common assembly mechanism of DNA translocating motors. J. Biol. Chem. 282:25569-25576.
258. Teachman, A. M., C. T. French, H. Yu, W. L. Simmons, and K. Dybvig. 2002. Gene transfer in Mycoplasma pulmonis. J. Bacteriol. 184:947-951.
259. Teng, W. L., T. L. Bannam, J. A. Parsons, and J. I. Rood. 2008. Functional characterization and localization of the TcpH conjugation protein from Clostridium perfringens. J. Bacteriol. 190:5075-5086.
260. te Poele, E. M., H. Bolhuis, and L. Dijkhuizen. 2008. Actinomycete integrative and conjugative elements. Antonie van Leeuwenhoek 94:127-143.
261. Terradot, L., R. Bayliss, C. Oomen, G. A. Leonard, C. Baron, and G. Waksman. 2005. Structures of two core subunits of the bacterial type IV secretion system, VirB8 from Brucella suis and ComB10 from Helicobacter pylori. Proc. Natl. Acad. Sci. USA 102:4596-4601.
262. Thomas, J., and D. W. Hecht. 2007. Interaction of Bacteroides fragilis pLV22a relaxase and transfer DNA with Escherichia coli RP4-TraG coupling protein. Mol. Microbiol. 66:948-960.
263. Tomich, M., P. J. Planet, and D. H. Figurski. 2007. The tad locus: postcards from the widespread colonization island. Nat. Rev. Microbiol. 5:363-375.
264. Ton-That, H., L. A. Marraffini, and O. Schneewind. 2004. Protein sorting to the cell wall envelope of gram-positive bacteria. Biochim. Biophys. Acta 1694:269-278.
265. Toro, N., A. Datta, O. A. Carmi, C. Young, R. K. Prusti, and E. W. Nester. 1989. The Agrobacterium tumefaciens virC1 gene product binds to overdrive, a T-DNA transfer enhancer. J. Bacteriol. 171:6845-6849.
266. Vanek, N. N., S. I. Simon, K. Jacques-Palaz, M. M. Mariscalco, G. M. Dunny, and R. M. Rakita. 1999. Enterococcus faecalis aggregation substance promotes opsonin-independent binding to human neutrophils via a complement receptor type 3-mediated mechanism. FEMS Immunol. Med. Microbiol. 26:49-60.
267. VanRheenen, S. M., Z. Q. Luo, T. O'Connor, and R. R. Isberg. 2006. Members of a Legionella pneumophila family of proteins with ExoU (phospholipase A) active sites are translocated to target cells. Infect. Immun. 74:3597-3606.
268. Varsaki, A., G. Moncalián, M. del Pilar Garcillán-Barcia, C. Drainas, and F. de la Cruz. 2009. Analysis of ColE1 MbeC unveils an extended ribbonhelix- helix family of nicking accessory proteins. J. Bacteriol. 191:1446-1455.
269. Vergunst, A. C., M. C. van Lier, A. den Dulk-Ras, T. A. G. Stuve, A. Ouwehand, and P. J. Hooykaas. 2005. Positive charge is an important feature of the C-terminal transport signal of the VirB/D4-translocated proteins of Agrobacterium. Proc. Natl. Acad. Sci. USA 102:832-837.
270. Vergunst, A. C., M. C. Van Lier, A. Den Dulk-Ras, and P. J. Hooykaas. 2003. Recognition of the Agrobacterium tumefaciens VirE2 translocation signal by the VirB/D4 transport system does not require VirE1. Plant Physiol. 133:978-988.
271. Vincent, C. D., and J. P. Vogel. 2006. The Legionella pneumophila IcmSLvgA protein complex is important for Dot/Icm-dependent intracellular growth. Mol. Microbiol. 61:596-613.
272. Voelker, L. L., and K. Dybvig. 1996. Gene transfer in Mycoplasma arthritidis: transformation, conjugal transfer of Tn916, and evidence for a restriction system recognizing AGCT. J. Bacteriol. 178:6078-6081.
273. Voth, D. E., and R. A. Heinzen. 2009. Coxiella type IV secretion and cellular microbiology. Curr. Opin. Microbiol. 12:74-80.
274. Voth, D. E., D. Howe, P. A. Beare, J. P. Vogel, N. Unsworth, J. E. Samuel, and R. A. Heinzen. 2009. The Coxiella burnetii ankyrin repeat domaincontaining protein family is heterogeneous, with C-terminal truncations that influence Dot/Icm-mediated secretion. J. Bacteriol. 191:4232-4242.
275. Walker, D. H., and N. Ismail. 2008. Emerging and re-emerging rickettsioses: endothelial cell infection and early disease events. Nat. Rev. Microbiol. 6:375-386.
276. Wang, A., and F. L. Macrina. 1995. Characterization of six linked open reading frames necessary for pIP501-mediated conjugation. Plasmid 34:206-210. (Pubitemid 126737933)
277. Wang, J., and K. M. Derbyshire. 2004. Plasmid DNA transfer in Mycobacterium smegmatis involves novel DNA rearrangements in the recipient, which can be exploited for molecular genetic studies. Mol. Microbiol. 53:1233-1241.
278. Wang, J., P. K. Karnati, C. M. Takacs, J. C. Kowalski, and K. M. Derbyshire. 2005. Chromosomal DNA transfer in Mycobacterium smegmatis is mechanistically different from classical Hfr chromosomal DNA transfer. Mol. Microbiol. 58:280-288.
279. Wang, J., L. M. Parsons, and K. M. Derbyshire. 2003. Unconventional conjugal DNA transfer in mycobacteria. Nat. Genet. 34:80-84.
280. Wang, Y. A., X. Yu, P. M. Silverman, R. L. Harris, and E. H. Egelman. 2009. The structure of F-pili. J. Mol. Biol. 385:22-29.
281. Ward, D. V., O. Draper, J. R. Zupan, and P. C. Zambryski. 2002. Peptide linkage mapping of the Agrobacterium tumefaciens vir-encoded type IV secretion system reveals protein subassemblies. Proc. Natl. Acad. Sci. USA 99:11493-11500.
282. Ward, J. E., Jr., E. M. Dale, and A. N. Binns. 1991. Activity of the Agrobacterium T-DNA transfer machinery is affected by virB gene products. Proc. Natl. Acad. Sci. USA 88:9350-9354.
283. Watarai, M., H. L. Andrews, and R. Isberg. 2000. Formation of a fibrous structure on the surface of Legionella pneumophila associated with exposure of DotH and DotO proteins after intracellular growth. Mol. Microbiol. 39:313-329.
284. Waters, C. M., and G. M. Dunny. 2001. Analysis of functional domains of the Enterococcus faecalis pheromone-induced surface protein aggregation substance. J. Bacteriol. 183:5659-5667.
285. Waters, C. M., H. Hirt, J. K. McCormick, P. M. Schlievert, C. L. Wells, and G. M. Dunny. 2004. An amino-terminal domain of Enterococcus faecalis aggregation substance is required for aggregation, bacterial internalization by epithelial cells and binding to lipoteichoic acid. Mol. Microbiol. 52:1159-1171.
286. Waters, C. M., C. L. Wells, and G. M. Dunny. 2003. The aggregation domain of aggregation substance, not the RGD motifs, is critical for effi- cient internalization by HT-29 enterocytes. Infect. Immun. 71:5682-5689.
287. Weiss, A. A., F. D. Johnson, and D. L. Burns. 1993. Molecular characterization of an operon required for pertussis toxin secretion. Proc. Natl. Acad. Sci. USA 90:2970-2974.
288. Winans, S. C., and G. C. Walker. 1985. Conjugal transfer system of the IncN plasmid pKM101. J. Bacteriol. 161:402-410. (Pubitemid 15157357)
289. Yeo, H. J., S. N. Savvides, A. B. Herr, E. Lanka, and G. Waksman. 2000. Crystal structure of the hexameric traffic ATPase of the Helicobacter pylori type IV secretion system. Mol. Cell 6:1461-1472.
290. Yeo, H. J., Q. Yuan, M. R. Beck, C. Baron, and G. Waksman. 2003. Structural and functional characterization of the VirB5 protein from the type IV secretion system encoded by the conjugative plasmid pKM101. Proc. Natl. Acad. Sci. USA 100:15947-15952.
291. Yoshida, H., N. Furuya, Y. J. Lin, P. Guntert, T. Komano, and M. Kainosho. 2008. Structural basis of the role of the NikA ribbon-helix-helix domain in initiating bacterial conjugation. J. Mol. Biol. 384:690-701.
292. Yuan, Q., A. Carle, C. Gao, D. Sivanesan, K. A. Aly, C. Hoppner, L. Krall, N. Domke, and C. Baron. 2005. Identification of the VirB4-VirB8-VirB5- VirB2 pilus assembly sequence of type IV secretion systems. J. Biol. Chem. 280:26349-26359.
293. Zahrl, D., A. Wagner, M. Tscherner, and G. Koraimann. 2007. GroEL plays a central role in stress-induced negative regulation of bacterial conjugation by promoting proteolytic degradation of the activator protein TraJ. J. Bacteriol. 189:5885-5894. (Pubitemid 47236141)
294. Zahrl, D., M. Wagner, K. Bischof, M. Bayer, B. Zavecz, A. Beranek, C. Ruckenstuhl, G. E. Zarfel, and G. Koraimann. 2005. Peptidoglycan degradation by specialized lytic transglycosylases associated with type III and type IV secretion systems. Microbiology 151:3455-3467. (Pubitemid 41632060)
295. Zahrl, D., M. Wagner, K. Bischof, and G. Koraimann. 2006. Expression and assembly of a functional type IV secretion system elicit extracytoplasmic and cytoplasmic stress responses in Escherichia coli. J. Bacteriol. 188:6611-6621. (Pubitemid 44412072)
296. Zamboni, D. S., S. McGrath, M. Rabinovitch, and C. R. Roy. 2003. Coxiella burnetii express type IV secretion system proteins that function similarly to components of the Legionella pneumophila Dot/Icm system. Mol. Microbiol. 49:965-976.
297. Zhang, S., and R. Meyer. 1997. The relaxosome protein MobC promotes conjugal plasmid mobilization by extending DNA strand separation to the nick site at the origin of transfer. Mol. Microbiol. 25:509-516.
298. Zhao, Z., E. Sagulenko, Z. Ding, and P. J. Christie. 2001. Activities of virE1 and the VirE1 secretion chaperone in export of the multifunctional VirE2 effector via an Agrobacterium type IV secretion pathway. J. Bacteriol. 183: 3855-3865.
299. Ziegelin, G., J. P. Furste, and E. Lanka. 1989. TraJ protein of plasmid RP4 binds to a 19-base pair invert sequence repetition within the transfer origin. J. Biol. Chem. 264:11989-11994.
300. Zupan, J., C. A. Hackworth, J. Aguilar, D. Ward, and P. Zambryski. 2007. VirB1* promotes T-pilus formation in the vir-type IV secretion system of Agrobacterium tumefaciens. J. Bacteriol. 189:6551-6563.