Chapter 9 : Bacterial Conjugation in Gram-Negative Bacteria

MyBook is a cheap paperback edition of the original book and will be sold at uniform, low price.

Preview this chapter:
Zoom in

Bacterial Conjugation in Gram-Negative Bacteria , Page 1 of 2

| /docserver/preview/fulltext/10.1128/9781555817732/9781555812652_Chap09-1.gif /docserver/preview/fulltext/10.1128/9781555817732/9781555812652_Chap09-2.gif


Bacterial conjugation is one of the fundamental processes for gene dissemination or horizontal gene transfer in nature and involves the transfer of DNA between bacteria in close apposition with one another. This chapter pulls together the information currently available on conjugative systems in gram-negative bacteria; reassess the information available before 1994, the year the sequences of two important gram-negative conjugation systems, F and RP4, appeared; and begin to construct a way of classifying and naming the genes from a plethora of conjugative systems, many of which appear to be chimeras of F and RP4. The junction between mating cells requires that the outer membranes of the two cells come into close apposition. The mechanism of active disaggregation of mating cells is unknown but might be related to the finding that transfer of IncI1 plasmids is terminated by a process requiring de novo protein synthesis in the new transconjugant. The relaxase, which is asymmetric in shape, is situated at the entry portal (base) of the pilus and is translocated as a first step in the conjugative process during mating-pair stabilization. Its inclusion in the conjugative pore could be in response to the triggering of conjugative DNA processing in the donor cell. The chapter discusses DNA processing and transport, the nature of the "signal" that triggers transfer, and mobilizable plasmids. It talks about F-like systems and P-like systems for which detailed analyses of gene regulation exist.

Citation: Lawley T, Frost L, Wilkins B. 2004. Bacterial Conjugation in Gram-Negative Bacteria , p 203-226. In Funnell B, Phillips G (ed), Plasmid Biology. ASM Press, Washington, DC. doi: 10.1128/9781555817732.ch9
Highlighted Text: Show | Hide
Loading full text...

Full text loading...


Image of Figure 1
Figure 1

A summary of the five families of known sites at the origin of transfer in conjugative and mohilizable plasmids. Conserved nucleotides are in black boxes whereas semiconserved nucleotides are in gray boxes. The position of the site is indicated with a wedge above the sequence. The I family of conjugative systems shares the same sequence as the P family. pMV158 is a mobilizable plasmid found in gram-positive bacteria ( ).

Citation: Lawley T, Frost L, Wilkins B. 2004. Bacterial Conjugation in Gram-Negative Bacteria , p 203-226. In Funnell B, Phillips G (ed), Plasmid Biology. ASM Press, Washington, DC. doi: 10.1128/9781555817732.ch9
Permissions and Reprints Request Permissions
Download as Powerpoint


1. Abuladze, N. K.,, M. Gingery,, J . Tsai,, and F. A. Eiserling. 1994. Tail length determination in bacteriophage T4. Virology 199: 301 310.
2. Achtman, M.,, N. Kennedy,, and R. Skurray. 1977. Cell-cell interactions in conjugating Escherichia coli: role of TraT protein in surface exclusion. Proc. Natl. Acad. Sci. USA 74: 5104 5108.
3. Achtman, M.,, G. Morelli,, and S. Schwuchow. 1978. Cell-cell interactions in conjugating Escherichia coli: role of F pili and fate of mating aggregates. J. Bacteriol. 135: 1053 1061.
4. Achtman, M.,, and R. Skurray,. 1977. A redefinition of the mating phenomenon in bacteria, p. 233 279. In J. L. Reissig (ed.), Microbial Interactions: Receptors and Recognition, vol. 3. Chapman and Hall, London, United Kingdom.
5. Althorpe, N. J . , P. M. Chilley, A. T. Thomas, W. J . Bram mar, and B. M. Wilkins. 1999. Transient transcriptional activation of the Incll plasmid anti-restriction gene ( ardA) and SOS inhibition gene ( psiB) early in conjugating recipient bacteria. Mol. Microbiol. 31: 133 142.
6. Anthony, K. G.,, W. A. Klimke,, J . Manchak,, and L. S. Frost. 1999. Comparison of proteins involved in pilus synthesis and mating pair stabilization from the related plasmids F and R100-1: insights into the mechanism of conjugation. J. Bacteriol. 181: 5149 5159.
7.Anthony, K G., C. Sherburne, R. Sherburne, and L. S. Frost. 1994. The role of the pilus in recipient cell recognition during bacterial conjugation mediated by F-like plasmids. Mol. Microbiol. 13: 939953.
8. Armstrong, G. D.,, L. S. Frost,, P. A. Sastry,, and W. Paranchych. 1980. Comparative biochemical studies on F and EDP208 conjugative pili. J. Bacteriol. 141: 333 341.
8a.. Arthur, D. C.,, A. F. Ghetu,, M. J. Gubbins,, R. A. Edwards,, L. S. Frost,, and J. N. Glover. 2003. FinO is an RNA chaperone that facilitates sense-antisensc RNA interactions. EMBO J. 22: 6346 6355.
9. Bagdasarian, M.,, A. Bailone,, J. F. Angulo,, P. Scholz,, and R. Devoret. 1992. PsiB, and anti-SOS protein, is transiently expressed by the F sex factor during its transmission to an Escherichia coli K-12 recipient. Mol. Microbiol. 6: 885 893.
10. Bailey, M. J . , C. Hughes, and V. Koronakis. 1996. Increased distal gene transcription by the elongation factor RfaH, a specialized homologue of NusG. Mol. Microbiol. 22: 729 737.
11. Bardwell, J. C.,, K. McGovern,, and J. Beckwith. 1991. Identification of a protein required for disulfide bond formation in vivo. Cell 67: 581 589.
12. Bates, S.,, A. M. Cash more,, and B. M. Wilkins. 1998. IncP plasmids are unusually effective in mediating conjugation of Escherichia coli and Saccharomyces cerevisiae: involvement of the tra2 mating system. J. Bacteriol. 180: 6538 6543.
13. Bates, S.,, R. A. Roscoe,, N. J. Althorpe,, W. J. Brammar,, and B. M. Wilkins. 1999. Expression of leading region genes on IncI1 plasmid ColIb-P9: genetic evidence for single-stranded DNA transcription. Microbiology 145: 2655 2662.
14. Bayer, M.,, R. Eferl,, G. Zellnig,, K. Teferle,, A. Dijkstra,, G. Koraimann,, and G. Hogenauer. 1995. Gene 19 of plasmid R1 is required for both efficient conjugative DNA transfer and bacteriophage R17 infection. J . Bacteriol. 177: 4279 4288.
15. Becker, E. C , and R. J. Meyer. 2002. MobA, the DNA strand transferase of plasmid R1162: the minimal domain required for DNA processing at the origin of transfer. J. Biol. Chem. 277: 14575 14580.
16. Beijersbergen, A.,, A. D. Dulk-Ras,, R. A. Schilperoort,, and P. J. Hooykaas. 1992. Conjugative transfer by the virulence system of Agrobacterium tumefaciens. Science 256: 1324 1327.
17. Bicbricher, C. K.,, and E. M. Düker. 1984. F and type 1 piliation of Escherichia coli. J. Gen. Microbiol. 130( Pt. 4): 951 957.
18. Bignell, C. R.,, A. S. Haines,, D. Khare,, and C. M. Thomas. 1999. Effect of growth rate and incC mutation on symmetric plasmid distribution by the IncP-1 partitioning apparatus. Mol. Microbiol. 34: 205 216.
19. Bingle, L. E.,, and C. M. Thomas. 2001. Regulatory circuits for plasmid survival. Curr. Opin. Microbiol. 4: 194 200.
20. 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.
21. Bonheyo, G.,, D. Graham,, N. B. Shoemaker,, and A. A. Salvers, 2001. Transfer region of a bacteroides conjugative transposon, CTnDOT. Plasmid 45: 41 51.
22. Borrebaeck, C. A. K.,, E. Soderlind,, L. S. Frost,, and A. C. Malmborg,. 1997. Combining bacterial and phage display: tapping the potential of molecular libraries for gene identification, p. 197 208. In P. Guttry (ed.), Display Technologies: Novel Targets and Strategies. IBC, Inc., Southborough, Mass..
23. Boulnois, G. J.,, and B. M. Wilkins. 1978. A colI-specified product, synthesized in newly infected recipients, limits the amount of DNA transferred during conjugation of Escherichia coli K-12. J. Bacteriol. 133: 1 9.
24. Boyd, A. C.,, J. A. Archer,, and D. J. Sherratt. 1989. Characterization of the ColE1 mobilization region and its protein products. Mol. Gen. Genet. 217: 488 498.
25. Bradley, D. E. 1984. Characteristics and function of thick and thin conjugative pili determined by transfer-dcrepressed plasmids of incompatibility groups 11, 12,15, B, K and Z. J. Gen. Microbial. 130: 1489 1502.
26. Bradley, D. E. 1989. Interaction of drug resistance plasmids and bacteriophages with diarrheagenic strains of Escherichia coli. Infect. Immun. 57: 2331 2338.
27. Bradley, D. E. 1980. Morphological and serological relationships of conjugative pili. Plasmid 4: 155 169.
28. Bradley, D. E.,, D. E. Taylor,, and D. R. Cohen. 1980. Specification of surface mating systems among conjugative drug resistance plasmids in Escherichia coli K-12. J. Bacteriol. 143: 1466 1470.
29. Bradley, D. E.,, and J. Whelan. 1985. Conjugation systems of IncT plasmids. J. Gen. Microbiol. 131( Pt 10): 2665 2671.
30. Bradley, D. E.,, and J. Whelan. 1989. Escherichia coli tolQ mutants are resistant to filamentous bacteriophages that adsorb to the tips, not the shafts, of conjugative pili. J. Gen. Microbiol. 135( Pt. 7): 1857 1863.
31. Buchanan-Wollaston, V.,, J. E. Passiatore,, and F. Cannon. 1987. The mob and oriT mobilization functions of a bacterial plasmid promote its transfer into plants. Nature 328: 172 174.
32.Cabezon, E,, E. Lanka, and F. de la Cruz. 1994. Requirements for mobilization of plasmids RSF1010 and ColE1 by the IncW plasmid R388: trwB and RP4 traG are interchangeable. J. Bacteriol. 176: 44554458.
33. 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.
34. Camacho, E. M.,, and J . Casadesus. 2002. Conjugal transfer of the virulence plasmid of Salmonella enterica is regulated by the leucine-responsive regulatory protein and DNA adenine methylation. Mol. Microbiol. 44: 1589 1598.
35. Chan, P. T., H, Ohmori, J., Tomizawa, and J. Lebowitz. 1985. Nucleotide sequence and gene organization of ColE1 DNA. J. Biol. Chem. 260: 8925 8935.
36. Cheah, K. C.,, and R. Skurray. 1986. The F plasmid carries an IS3 insertion within finO. J. Gen. Microbiol. 132(Pt. 121: 3269 3275.
37. Chen, C. Y.,, and C. I. Kado. 1994. Inhibition of Agrobacterium tumefaciens oncogenicity by the osa gene of pSa. J. Bacteriol. 176: 5696 5703.
38. Chilley, P. M.,, and B. M. Wilkins. 1995. Distribution of the ardA family of antirestriction genes on conjugative plasmids. Microbiology 141( Pt. 9): 2157 2164.
39. Christie, P. J . 2001. Type IV secretion: intercellular transfer of macromolecules by systems ancestrally related to conjugation machines. Mol. Microbiol. 40: 294 305.
40. Christie, P. J.,, and J. P. Vogel. 2000. Bacterial type IV secretion: conjugation systems adapted to deliver effector molecules to host cells. Trends Microbiol. 8: 354 360.
41. Citovsky, V.,, J. Zupan,, D. Warnick,, and P. Zambryski. 1992. Nuclear localization of Agrobacterium VirE2 protein in plant cells. Science 256: 1802 1805.
42. Clewell, D. 1993. Bacterial Conjugation. Plenum Press, New York, N.Y..
43. Click, E. M. and R. E. Webster. 1997. Filamentous phage infection: required interactions with the TolA protein. J. Bacteriol. 179: 6464 6471.
44. Danesc, P. N.,, and T. J . Silhavy. 1997. The sigma(E) and the Cpx signal transduction systems control the synthesis of periplasmic protein-folding enzymes in Escherichia coli. Genes Dev. 11: 1183 1193.
45. Daugelavicius, R.,, J. K. Bamford,, A. M. Grahn,, E. Lanka,, and D. H. Bamford. 1997. The IncP plasmid-encoded cell envelope-associated DNA transfer complex increases cell permeability. J. Bacteriol. 179: 5195 5202.
46. Davis, B. M.,, E. H. Lawson,, M. Sandkvist,, A. Ali,, S. Sozhamannan,, and M. K. Waldor. 2000. Convergence of the secretory pathways for cholera toxin and the filamentous phage, CTXphi. Science 288: 333 335.
47. Davison, J. 1999. Genetic exchange between bacteria in the environment. Plasmid 42: 73 91.
48. Deng, L. W.,, and R. N. Perham. 2002. Delineating the site of interaction on the pill protein of filamentous bacteriophage fd with the F-pilus of Escherichia coli. J. Mol. Biol. 319: 603 614.
49. Di Laurenzio, L.,, D. G. Scraba,, W. Paranchych,, and L. S. Frost, 1995. Studies on the binding of integration host factor (IHF) and TraM to the origin of transfer of the IncFV plasmid pED208. MoL Gen. Genet. 247: 726 734.
50.Disque-Kocltem, C , and B. Dreiseikelmann. 1997. The cytoplasmic DNA-binding protein TraM binds to the inner membrane protein TraD in vitro. J. Bacteriol. 179: 61336137.
51. Doran, T. J.,, S. M. Loh,, N. Firth,, and R. A. Skurray. 1994. Molecular analysis of the F plasmid traVR region: traV encodes a lipoprotein. J. Bacteriol. 176: 4182 4186.
52. Dreiseikelmann, B. 1994. Translocation of DNA across bacterial membranes. Microbiol. Rev. 58: 293 316.
53. Dumas, F.,, M. Duckely,, P. Pclczar,, P. Van Gelder,, and B. Hohn. 2001. An Agrobacterium VirE2 channel for transferred-DNA transport into plant cells. Proc. Natl. Acad. Sci. USA 98: 485 490.
54. Dürrenberger, 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.
55. Eisenbrandt, R.,, M. Kalkum,, E. M. Lai,, R. Lurz,, C. I. Kado,, and E. Lanka. 1999. Conjugative pili of IncP plasmids, and the Ti plasmid T pilus are composed of cyclic subunits. J. Biol. Chem. 274: 22548 22555.
56. 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.
57. 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.
58. Fekete, R.,, and L. S. Frost. 2002. Characterizing the DNA contacts and cooperative binding of F plasmid TraM to its cognate sites at oriT. J. Biol. Chem. 277: 16705 16711.
59. Fekete, R. A.,, and L. S. Frost. 2000. Mobilization of chimeric oriT plasmids by F and R100-1: role of relaxosome formation in defining plasmid specificity, J. Bacteriol. 182: 4022 4027.
60. Feng, J. N.,, P. Model,, and M. Russel. 1999. A trans-envelope protein complex needed for filamentous phage assembly and export. Mol. Microbiol. 34: 745 755.
61. Firth, N.,, K. Ippen-Ihler,, and R. A. Skurray,. 1996. Structure and function of the F factor and mechanism of conjugation, p. 2377 2401. In F. C. Neidhardt (ed.), Escherichia coli and Salmonella: Cellular and Molecular Biology, 2nd ed., vol. 2. American Society for Microbiology, Washington, D.C..
62. Firth, N.,, and R. Skurray. 1992. Characterization of the F plasmid bifunctional conjugation gene, traG. Mol. Gen. Genet. 232: 145 153.
63. Frost, L. S., 1993. Conjugative pili and pilus-specific phages, p. 189 221. In D. B. Clewell (ed.), Bacterial Conjugation. Plenum Press, New York, N.Y..
64. Frost, L. S.,, K. Ippen-Ihler,, and R. A. Skurray. 1994. Analysis of the sequence and gene products of the transfer region of the F sex factor. Microbiol. Rev. 58: 162 210.
65. Frost, L. S.,, J. S. Lee,, D. G. Scraba,, and W. Paranchych. 1986. Two monoclonal antibodies specific for different epitopes within the ammo-terminal region of F pilin. J. Bacteriol 168: 192 198.
66. Frost, L. S.,, and J. Manchak. 1998. F- phenocopics: characterization of expression of the F transfer region in stationary phase. Microbiology 144( Pt. 9): 2579 2587.
67. Frost, L. S.,, and J. Simon,. 1993. Studies on the pili of promiscuous plasmid RP4, p. 47 65. In C. I. Kado, and J. H. Crosa (ed.), Molecular Mechanisms of Bacterial Virulence. Kluwer Academic Publishers, Amsterdam, The Netherlands.
68. Fu, Y. H.,, M. M. Tsai,, Y. N. Luo,, and R. C. Deonier. 1991. Deletion analysis of the F plasmid oriT locus. J . Bacteriol, 173: 1012 1020.
69. Gao, Q.,, Y. Luo,, and R. C. Deonier. 1994. Initiation and termination of DNA transfer at F plasmid oriT. Mol. Microbiol. 11: 449 158.
70. Garcia, L. R.,, and I. J. Molineux. 1995. Incomplete entry of bacteriophage T7 DNA into F plasmid-containing Escherichia coli. J. Bacteriol. 177: 4077 4083.
71. Ghetu, A. F.,, M. J , Gubbins, L. S. Frost, and J . N. Glover. 2000. Crystal structure of the bacterial conjugation repressor finO. Nat. Struct. Biol. 7: 565 569.
72. Ghigo, J. M. 2001. Natural conjugative plasmids induce bacterial biofilm development. Nature 412: 442 445.
73. Giebelhaus, L. A.,, L. Frost,, E. Lanka,, E. P. Gormley,, J. E. Davies,, and B. Leskiw. 1996. The Tra2 core of the IncP(alpha) plasmid RP4 is required for intergeneric mating between Escherichia coli and Streptomyces lividans. J. Bacteriol. 178: 6378 6381.
75. 73a. Gilmour, M. W., J. E. Gunton, T. D. Lawley, and D. E. Taylor. 2003. Interaction between the IncHI1 plasmid R27 coupling protein and type IV secretion system: TraG associates with the coiled-coil mating pair formation protein TrhB. Mol. Microbiol. 49: 105116.
74. Gilmour, M. W.,, T. D. Lawlcy,, M. M. Rookcr,, P. J. Newnham,, and D. E. Taylor. 2001. Cellular location and temperature-dependent assembly of IncHI1 plasmid R27-encoded TrhC-associatcd conjugative transfer protein complexes. Mol. Microbiol. 42: 705 715.
75. Golub, E.,, A. Bailonce, and R. Devoret. 1988. A gene encoding an SOS inhibitor is present in different conjugative plasmids. J. Bacteriol. 170: 4392 4394.
76. Gomis-Ruth, F. X.,, F. de la Cruz,, and M. ColI. 2002. Structure and role of coupling proteins in conjugal DNA transfer. Res. Microbiol. 153: 199 204.
77. Gomis-Ruth, F. X.,, G. Moncalian,, R. Perez-Luque,, A. Gonzalez,, E. Cabezon,, F. de la Cruz,, and M. ColI. 2001. The bacterial conjugation protein TrwB resembles ring helicases and F1-ATPase. Nature 409: 637 641.
78. Gordon, G. S.,, D. Sitnikov,, C. D. Webb,, A. Teleman,, A. Straight,, R. Losick,, A. W. Murray,, and A. Wright. 1997. Chromosome and low copy plasmid segregation in E. coli: visual evidence for distinct mechanisms. Cell 90: 1113 1121.
79. Gordon, G. S.,, and A. Wright. 2000. DNA segregation in bacteria. Annu. Rev. Microbiol. 54: 681 708.
80. Grahn, A. M.,, J. Haase,, D. H. Bamford,, and E. Lanka. 2000. Components of the RP4 conjugative transfer apparatus form an envelope structure bridging inner and outer membranes of donor celts: implications for related macromolecule transport systems. J. Bacteriol. 182: 1564 1574.
81. Grahn, A. M.,, J. Haase,, E. Lanka,, and D. H. Bamford, 1997. Assembly of a functional phage PRD1 receptor depends on 11 genes of the IncP plasmid mating pair formation complex. J. Bacteriol. 179: 4733 4740.
82. Grandoso, G.,, P. Avila,, A. Cayon,, M. A. Hernando,, M. Llosa,, and F. dela Cruz. 2000. Two active-site tyrosyl residues of protein TrwC act sequentially at the origin of transfer during plasmid R388 conjugation. J. Mol. Biol. 295: 1163 l 172.
83. Grossman, T. H.,, and P. M. Silverman. 1989. Structure and function of conjugative pili: inducible synthesis of functional F pili by Eschericim coli K-12 containing a lac-tra operon fusion. J. Bacteriol. 171: 650 656.
84. Guiney, D. G. 1982. Host range of conjugation and replication functions of the Escherichia coli sex plasmid Flac. Comparison with the broad host-range plasmid RK2. J. Mol. Biol. 162: 699 703.
85. Gyohda, A.,, N. Funayama,, and T. Komano. 1997. Analysis of DNA inversions in the shufflon of plasmid R64. J. Bacteriol. 179: 1867 1871.
86. Haase, J.,, M. Kalkum,, and E. Lanka. 1996. TrbK, a small cytoplasmic membrane lipoprotein, functions in entry exclusion of the IncP alpha plasmid RP4. J. Bacteriol. 178: 6720 6729.
87. Haase, J.,, and E. Lanka. 1997. A specific protease encoded by the conjugative DNA transfer systems of IncP and Ti plasmids is essential for pilus synthesis. J. Bacteriol. 179: 5728 5735.
88. Haase, J.,, R. Lurz,, A. M. Grahn,, D. H. Bamford,, and E. Lanka. 1995. Bacterial conjugation mediated by plasmid RP4: RSF1010 mobilization, donor-specific phage propagation, and pilus production require the same Tra2 core components of a proposed DNA transport complex. J. Bacteriol. 177: 4779 4791.
89. Hall, M. C.,, and S. W. Matson. 1999. Helicase motifs: the engine that powers DNA unwinding. Mol. Microbiol. 34: 867 877.
90. Hamilton, H. L.,, K. J. Schwartz,, and J. P. Dillard. 2001. Insertion-duplication mutagenesis of Neisseria: use in characterization of DNA transfer genes in the gonococcal genetic island. J. Bacteriol. 183: 4718 4726.
91. Hapfelmeier, S.,, N. Domke,, P. C. Zambryski,, and C. Baron. 2000. VirB6 is required for stabilization of VirB5 and VirB3 and formation of VirB7 homodimers in Agrobacterium tumefaciens. J. Bacteriol. 182: 4505 4511.
92. Harris, R. L.,, V. Hombs,, and P. M. Silverman. 2001. Evidence that F-plasmid proteins TraV, TraK and TraB assemble into an envelope-spanning structure in Escherichia coli. Mol. Microbiol. 42: 757 766.
93. Hartskeerl, R. A.,, and W. P. Hoekstra. 1984. Exclusion in Incl-type Escberichi coli conjugations: the stage of conjugation at which exclusion operates. Anionic Leeuwenhoek 50: 113 124.
94. Heincmann, J. A.,, and G. F. Sprague Jr., 1989. Bacterial conjugative plasmids mobilize DNA transfer between bacteria and yeast. Nature 340: 205 209.
95. Henderson, D., and R, Meyer. 1999. The MobA-linked primase is the only replication protein of R1162 required for conjugal mobilization. J. Bacteriol. 181: 2973 2978.
96. Hendrix, R. W. 1998. Bacteriophage DNA packaging: RNA gears in a DNA transport machine. Cell 94: 147 150.
97. Hochhut, B.,, and M. K. Waldor. 1999. Site-specific integration of the conjugal Vibrio cholerae SXT element into prfC. Mol. Microbiol. 32: 99 110.
98. Hofreuter, D.,, and R. Haas. 2002. Natural transformation in Helicobacter pylori: DNA transport in an unexpected way. Trends Microbiol. 10: 159 162.
99. 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.
100. Holloway, B. W. 1979. Plasmids that mobilize bacterial chromosome. Plasmid 2: 1 19.
101. Howard, M. T.,, W. C. Nelson,, and S. W. Matson. 1995. Stepwise assembly of a relaxosome at the F plasmid origin of transfer. J. Biol. Chem. 270: 28381 28386.
102. Jacobson, A. 1972. Role of F pili in the penetration of bacteriophage fl. J. Virol. 10: 835 843.
103. Jagura-Burdzy, G., and C, M. Thomas. 1997. Dissection of the switch between genes for replication and transfer of promiscuous plasmid RK2: basis of the dominance of trfAp over trbAp and specificity for KorA in controlling the switch. J. Mol. Biol. 265: 507 518.
104. Jerome, L. J.,, and L. S. Frost. 1999. In vitro analysis of the interaction between the FinO protein and FinP antisense RNA of F-like conjugative plasmids. J . Biol. Chem. 274: 10356 10362.
105. Jerome, L. J.,, T. van Biesen,, and L. S. Frost. 1999. Degradation of FinP antisense RNA from F-like plasmids: the RNA-binding protein, FinO, protects FinP from rihonucleasc E. J. Mol. Biol. 285: 1457 1473.
106. Jones, A. L.,, P. T. Barth,, and B. M. Wilkins. 1992. Zygotic induction of plasmid ssb and psiB genes following conjugative transfer of Incl1 plasmid ColIb-P9. Mol. Microbiol. 6: 605 613.
107. Kim, S. R.,, and T. Komano. 1997. The plasmid R64 thin pilus identified as a type IV pilus. J. Bacteriol. 179: 3594 3603.
108. Kingsman, A.,, and N. Willetts. 1978. The requirements for conjugal DNA synthesis in the donor strain during flat transfer. J. Mol. Biol. 122: 287 300.
109.Klimke, W, A., and L. S. Frost. 1998. Genetic analysis of the role of the transfer gene, traN. of the F and R100-1 plasmids in mating pair stabilization during conjugation. J. Bacteriol. 180: 40364043.
110. Koraimann, G.,, K. Teferle,, G. Markolin,, W. Woger,, and G. Hogenauer. 1996. The FinOP repressor system of plasmid Rl: analysis of the antisense RNA control of traj expression and conjugative DNA transfer. Mol. Microbiol. 21: 811 821.
111. Kostelidou, K.,, A. C. Jones,, and C. M. Thomas. 1999. Conserved C-terminal region of global repressor KorA of broad-host-range plasmid RK2 is required for co-operativity between KorA and a second RK2 global regulator, KorB. J. Mol. Biol. 289: 211 221.
112. Kotilaincn, M. M.,, A. M. Grahn,, J. K. Bamford,, and D. H. Bamford. 1993. Binding of an Escherichia coli double-stranded DNA virus PRD1 to a receptor coded by an IncP-type plasmid. J. Bacteriol. 175: 3089 3095.
113. Krause, S.,, W. Pansegrau,, R. Lurz,, F. de la Cruz,, and E. Lanka. 2000. Enzymology of type IV macromolcculc 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.
114. Kumar, J. K.,, S. Tabor,, and C. C. Richardson. 2001. Role of the C-terminal residue of the DNA polymerase of bacteriophage T7, J. Biol. Chem. 276: 34905 34912.
115. Kupelwieser, G.,, M. Schwab,, G. Hogenauer,, G. Koraimann,, and E. L. Zechner. 1998. Transfer protein TraM stimulates Tral-catalyzed cleavage of the transfer origin of plasmid R1 in vivo. J. Mol. Biol. 275: 81 94.
116. Lanka, E.,, and B. M. Wilkins. 1995. DNA processing reactions in bacterial conjugation. Annu. Rev. Biochem. 64: 141 169.
117. Lawlcy, T. D.,, M. W. Gilmour,, J. E. Gunton,, D. Tracz,, and D. E. Taylor. 2003. Functional and mutational analysis of the conjugative transfer region 2 (Tra2) of the IncHI1plasmid R27. J. Bacteriol. 185: 581 591,
118. Lawley, T. D.,, G. S. Gordon,, A. Wright,, and D. E. Taylor. 2002. Bacterial conjugative transfer: visualization of successful mating pairs and plasmid establishment in live Escherichia coli. Mol. Microbiol. 44: 947 956.
119. Lessl, M.,, D. Balzer,, R. Lurz,, V. L. Waters,, D. G. Guiney,, and E. Lanka. 1992. Dissection of IncP conjugative plasmid transfer: definition of the transfer region Tra2 by mobilization of the Tra1 region in trans. J. Bacteriol. 174: 2493 2500,
120. Lessl, M.,, D. Balzer,, K. Weyrauch,, and E. Lanka. 1993. The mating pair formation system of plasmid RP4 defined by RSF1010 mobilization and donor-specific phage propagation. J. Bacteriol. 175: 6415 6425.
121. Llosa, M. F. X. Gomis-Ruth, M. Coll, and L. Cruz Fd Fde. 2002. Bacterial conjugation: a two-step mechanism for DNA transport. Mol. Microbiol. 45: 1 8.
122. Loh, S. M.,, D. S. Cram, and R A. Skurray. 1988. Nucleotide sequence and transcriptional analysis of a third function (Flm) involved in F-plasmid maintenance. Gene 66: 259 268.
123.Lorenz, M, G., and W. Wackernagel. 1994. Bacterial gene transfer by natural genetic transformation in the environment. Microbiol. Rev. 58: 563602.
124. Lovett, M. A.,, and D. R. Helinski. 1975. Relaxation complexes of plasmid DNA and protein. II. Characterization of the proteins associated with the unrelaxed and relaxed complexes of plasmid ColE1. J. Biol. Chem. 250: 8790 8795.
125. Low, K. B., 1996. Hfr strains of Escherichia coli K-12, p. 2402 2405. In F. C. Neidhardt (ed.), Escherichia coli and Salmonella: Cellular and Molecular Biology, 2nd ed., vol. 2. American Society for Microbiology, Washington, D.C..
126. Lu, J.,, J. Manchak,, W. Klimke,, C. Davidson,, N. Firth,, R. A. Skurray,, and L. S. Frost. 2002. Analysis and characterization of IncFV plasmid pED208 transfer region. Plasmid 48: 24 34.
127. Lubkowski, J.,, F. Hennecke,, A. Pluckthun,, and A. Wlodawer, 1999. Filamentous phage infection: crystal structure of g3p in complex with its coreceptor, the C-terminal domain of To1A. Structure Fold Des. 7: 711 722.
128. Lum, P. L.,, and J. F. Schildbach. 1999. Specific DNA recognition by F factorTraY involves beta-sheet residues. J. Biol. Chem. 274: 19644 19648.
129. Macnab, R. M.. 2000. Microbiology. Action at a distance— bacterial flagellar assembly. Science 290: 2086 2097.
130. Maher, D.,, R. Sherburne,, and D. E. Taylor. 1993. H-pilus assembly kinetics determined by electron microscopy. J. Bacteriol. 175: 2175 2183.
131. Majdalani, N.,, and K. Ippen-lhler. 1996. Membrane insertion of the F-pilin subunit is Sec independent but requires leader peptidase B and the proton motive force. J. Bacteriol. 178: 3742 3747.
132. Majdalani, N.,, D. Moore,, S. Maneewannakul,, and K. Ippen-Ihler. 1996. Role of the propilin leader peptide in the maturation of F pilin. J. Bacteriol. 178: 3748 3754.
133. Manchak, J.,, K. G. Anthony,, and L. S. Frost. 2002. Mutational analysis of F-pilin reveals domains for pilus assembly, phage infection and DNA transfer. Mol. Microbiol. 43: 195 205.
134. Maneewannakul, K.,, P. Kathir,, S. Endley,, D. Moore,, J. Manchak,, L. Frost,, and K. Ippen-lhler. 1996. Construction of derivatives of the F plasmid pOX-tra715: characterization of traY and traD mutants that can be complemented in trans. Mol. Microbiol. 22: 197 205.
135. Maneewannakul, K.,, S. Maneewannakul,, and K. Ippen-lhler. 1992. Sequence alterations affecting F plasmid transfer gene expression: a conjugation system dependent on transcription by the RNA polymerase of phage T7. Mol. Microbiol. 6: 2961 2973.
136. Maneewannakul, S.,, P. Kathir,, and K. Ippen-Ihler. 1992. Characterization of the F plasmid mating aggregation gene traN and of a new F transfer region locus trbE. J . Mol. Biol. 225: 299 311.
137. Manning, P. A.,, G. Morell,, and M. Achtman. 1981. traG protein of the F sex factor of Escherichia coli K-12 and its role in conjugation. Proc. Natl. Acad. Sci. USA 78: 7487 7491.
138. Manwaring, N. P.,, R. A. Skurray,, and N. Firth. 1999. Nucleotide sequence of the F plasmid leading region. Plasmid 41: 219 225.
139. Marvin, D. A.,, and W. Folkhard. 1986. Structure of F-pili: reassessment of the symmetry. J. Mol. Biol. 191: 299 300.
140. Marzec, C. J.,, and L. A. Day. 1988. A theory of the symmetries of filamentous bacteriophages. Biopbys. J. 53: 425 440.
141. Meinhardt, H.,, and P. A. de Boer. 2001. Pattern formation in Escherichia coli: a model for the pole-to-pole oscillations of Min proteins and the localization of the division site. Proc. Natl. Acad. Sci. USA 98: 14202 14207.
142. Meyer, R. 2000. Identification of the mob genes of plasmid pSC101 and characterization of a hybrid pSC101-R1162 system for conjugal mobilization. J. Bacteriol. 182: 4875 4881.
143. Mileykovskaya, E.,, and W. Dowhan. 1997. The Cpx two-component signal transduction pathway is activated in Escherichia coli mutant strains lacking phosphatidyl-ethanolamine. J. Bacteriol. 179: 1029 1034.
144. Moncalian, G.,, M. Valle,, J. M. Valpuesta,, and F. de la Cruz. 1999. IHF protein inhibits cleavage but not assembly of plasmid R388 relaxosomes. Mol. Microbiol. 31: 1643 1652.
145. Moore, D., C M. Hamilton, K. Maneewannakul, Y, Mintz, L. S. Frost, and K, Ippen-Ihler. 1993. The Escherichia coli K-12 F plasmid gene traX is required for acctylation of F pilin. J. Bacteriol. 175: 1375 1383.
146. Moore, D.,, K. Maneewannakul,, S. Maneewannakul,, J. H. Wu,, K. Ippen-lhler,, and D. E. Bradley. 1990. Characterization of the F-plasmid conjugative transfer gene traU. J. Bacteriol. 172: 4263 4270.
147. Motallebi-Veshareh, M., D, Balzer, E. Lanka, G. Jagura-Burdzy, and C. M. Thomas. 1992. Conjugative transfer functions of broad-host-range plasmid RK2 arc coregulated with vegetative replication. Mol. Microbiol. 6: 907 920.
148. Murata, T.,, M. Ohnishi,, T. Ara,, J. Kaneko,, C. G. Han,, Y. F. Li,, K. Takashima,, H. Nojima,, K. Nakayama,, A. Kaji,, Y. Kamio,, T. Miki,, H. Mori,, E. Ohtsubo,, Y. Terawaki,, and T. Hayashi. 2002. Complete nucleotide sequence of plasmid Rts1: implications for evolution of large plasmid genomes. J. Bacteriol. 184: 3194 3202.
149. Nelson, W. C.,, and S. W. Matson. 1996. The F plasmid traY gene product binds DNA as a monomer or a dimer: structural and functional implications. Mol. Microbiol. 20: 1179 1187.
150. Nomura, N.,, H. Masai,, M. Inuzuka,, C. Miyazaki,, E. Ohtsubo,, T. Itoh,, S. Sasamoto,, M. Matsui,, R. Ishizaki,, and K. Arai. 1991. Identification of eleven single-strand initiation sequences (ssi) for priming of DNA replication in the F, R6K, R100 and ColE2 plasmids. Gene 108: 15 22.
151. Novotny, C. P.,, and P. Fives-Taylor. 1978. Effects of high temperature on Escherichia coli F pili. J. Bacteriol. 133: 459 464.
152. Novotny, C. P.,, and P. Fives-Taylor, 1974. Retraction of F pili. J. Bacteriol. 117: 1306 1311.
153. Nunez, B.,, and F. De La Cruz. 2001. Two atypical mobilization proteins are involved in plasmid CloDF13 relaxation. Mol. Microbiol. 39: 1088 1099.
154. Ottow, J.,, C. 1975. Ecology, physiology, and genetics of fimbriae and pili. Annu. Rev. Microbiol. 29: 79 108.
155. Ou, J. T. 1975. Mating signal and DNA penetration deficiency in conjugation between male Escherichia coli and minicells. Proc. Natl. Acad. Sci. USA 72: 3721 3725.
156. Ou, J. T. 1980. Role of surface exclusion genes in lethal zygosis in Escherichia coli K12 mating. Mol. Gen. Genet. 178: 573 581.
157. Paiva, W. D.,, T. Grossman,, and P. M. Silverman. 1992. Characterization of F-pilin as an inner membrane component of Escherichia coli K12 J. Biol. Chem. 267: 26191 26197.
158. Panicker, M. M.,, and E. G. Minklcy Jr., 1985. DNA transfer occurs during a cell surface contact stage of F sex factor-mediated bacterial conjugation. J. Bacteriol. 162: 584 590.
159. Pansegrau, W.,, and E. Lanka. 1996. Enzymology of DNA transfer by conjugative mechanisms. Prog. Nucleic Acid Res. Mol. Biol. 54: 197 251.
160. Pansegrau, W.,, E. Lanka,, P. T. Barth,, D. H. Figurski,, D. G. Guiney,, D. Haas,, D. R. Helinski,, H. Schwab,, V. A. Stanisich,, and C. M. Thomas. 1994. Complete nucleotide sequence of Birmingham IncP alpha plasmids. Compilation and comparative analysis. J. Mol. Biol. 239: 623 663.
161. Panscgrau, W.,, G. Ziegelin,, and E. Lanka. 1990. Covalent association of the tral gene product of plasmid RP4 with the 5'-terminal nucleotide at the relaxation nick site. J. Biol. Chem. 265: 10637 10644.
162. Paranchych, W.,, B. B. Finlay,, and L. S. Frost. 1986. Studies on the regulation of IncF plasmid transfer operon expression. Banbury Rep. 24: 117 129.
163. Paranchych, W., and L, S. Frost. 1988. The physiology and biochemistry of pili. Adv. Microb. Physiol. 29: 53 114.
164. Paul, J. H. 1999. Microbial gene transfer: an ecological perspective. J. Mol. Microbiol. Biotechnol. 1: 45 50.
165. Pembroke, J. T.,, and D. B. Murphy. 2000. Isolation and analysis of a circular form of the IncJ conjugative transposon-like elements, R391 and R997: implications for IncJ incompatibility. FEMS Microbiol. Lett. 187: 133 138.
166. Penfold, S. S.,, J . Simon,, and L. S. Frost. 1996. Regulation of the expression of the traM gene of the F sex factor of Escherichia coli. Mol. Microbiol. 20: 549 558.
167. Penfold, S. S.,, K. Usher,, and L. S. Frost. 1994. The nature of the traK4 mutation in the F sex factor of Escherichia coli. J. Bacteriol. 176: 1924 1931.
168. Perumal, N. B.,, and E. G. Minkley Jr., 1984. The product of the F sex factor traTsurface exclusion gene is a lipoprotein. J. Biol. Chem. 259: 5357 5360.
169. Pcrwcz, T.,, and R. J . Meyer, 1999. Stabilization of the relaxosome and stimulation of conjugal transfer are genetically distinct functions of the R1162 protein MobB. J. Bacteriol. 181: 2124 2131.
170. Planet, P. J.,, S. C. Kachlany,, R. DeSalle,, and D. H. Figurski. 2001. Phylogeny of genes for secretion NTPases: identification of the widespread tadA subfamily and development of a diagnostic key for gene classification. Proc. Natl. Acad. Sci. USA 98: 2503 2508.
171. Pogliano, J.,, T. Q. Ho,, Z. Zhong,, and D. R. Helinski. 2001. Multicopy plasmids are clustered and localized in Escherichia coli. Proc. Natl. Acad. Sci. USA 98: 4486 4491.
172. Pogliano, J.,, A. S. Lynch,, D. Belin,, E. C. Lin,, and J. Beckwith. 1997. Regulation of Escherichia coli cell envelope proteins involved in protein folding and degradation by the Cpx two-component system. Genes Dev. 11: 1169 1182.
173. Polzleitner, E.,, E. L. Zechner,, W. Renner,, R. Fratte,, B. Jauk,, G. Hogenauer,, and G. Koraimann. 1997. TraM of plasmid R1 controls transfer gene expression as an integrated control clement in a complex regulatory network. Mol. Microbiol. 25: 495 507.
174. Recs, C. E.,, and B. M. Wilkins. 1990. Protein transfer into the recipient cell during bacterial conjugation: studies with F and RP4. Mol. Microbiol. 4: 1199 1205.
175. Reimmann, C.,, and D. Haas,. 1993. Mobilization of chromosomes and nonconjugative plasmids by cointegrative mechanisms, p. 137 188. In D. B. Clewell (ed.), Bacterial Conjugation. Plenum Press, New York, N.Y..
176. Riechmann, L.,, and P. Holligcr. 1997. The C-terminal domain of TolA is the coreceptor for filamentous phage infection of E. coli. Cell 90: 351 360.
177. Riede, I.,, and M. L. Eschbach. 1986. Evidence that TraT interacts with OmpA of Escherichia coli. FEBS Lett. 205: 241 245.
178. Rondot, S.,, K. G. Anthony,, S. Dubel,, N. Ida,, S. Wiemann,, K. Beyrcuther,, L. S. Frost, M, Little, and F. Breitling. 1998. Epitopes fused to F-pilin are incorporated into functional recombinant pili. J. Mol. Biol. 279: 589 603.
179. Rosche, T. M.,, A. Siddique,, M. H. Larsen,, and D. H. Figurski. 2000. Incompatibility protein IncC and global regulator KorB interact in active partition of promiscuous plasmid RK2. J. Bacteriol. 182: 6014 6026.
180. Sakai, H.,, and T. Romano. 1996. DNA replication of IncQ broad-host-range plasmids in gram-negative bacteria. Biosci. Biotechnol. Biochem. 60: 377 382.
181. Salyers, A. A.,, N. B. Shoemaker,, A. M. Stevens,, and L. Y. Li. 1995. Conjugative transposons: an unusual and diverse set of integrated gene transfer elements. Microbiol. Rev. 59: 579 590.
182. Samuels, A. L.,, E. Lanka,, and J. E. Davies. 2000. Conjugative junctions in RP4-mediated mating of Escherichia coli. J. Bacteriol. 182: 2709 2715.
183. Santini, J . M. and V. A, Stanisich. 1998. Both the fipA gene of pKM101 and the pifC gene of F inhibit conjugal transfer of RP1 by an effect on traG. J. Bacteriol. 180: 4093 4101.
184. Sastre, J . I.,, E. Cabezon,, and F. de la Cruz. 1998. The carboxyl terminus of protein TraD adds specificity and efficiency to F-plasmid conjugative transfer. J. Bacteriol. 180: 6039 6042.
185. Schandel, K. A.,, S. Maneewannakul,, K. Ippen-Ihler,, and R. E. Webster. 1987. A traC mutant that retains sensitivity to f1 bacteriophage but lacks F pili. J. Bacteriol. 169: 3151 3159.
186. Scholz, P.,, V. Haring,, B. Wittmann-Liebold,, K. Ashman,, M. Bagdasarian,, and E. Scherzinger. 1989. Complete nucleotide sequence and gene organization of the broad-host-range plasmid RSFI010. Gene 75: 271 288.
187. 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.
188. Shapiro, J. A., 1977. Bacterial plasmids, p. 601 670. In A. I. Bukhari,, J. A. Shapiro,, and S. L. Adhya (ed.), DNA Insertion Elements, Plasmids, and Episomes. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, N.Y..
189. Silverman, P. M. 1997. Towards a structural biology of bacterial conjugation. Mol. Microbiol. 23: 423 429.
190. Silverman, P. M.,, S. Rother,, and H. Gaudin. 1991. Arc and Sfr functions of the Escherichia coli K-12 arcA gene product are genetically and physiologically separable. J. Bacteriol. 173: 5648 5652.
191. Silverman, P. M.,, and A. Sholl. 1996. Effect of traY amber mutations on F-plasmid traY promoter activity in vivo. J. Bacteriol. 178: 5787 5799.
192. Silverman, P. M.,, L. Tran,, R. Harris,, and H. M. Gaudin. 1993. Accumulation of the F plasmid TraJ protein in cpx mutants of Escherichia coli. J. Bacteriol. 175: 921 925.
193. Silverman, P. M., E, Wickersham, S. Rainwater, and R. Harris. 1991. Regulation of the F plasmid traY promoter in Escherichia coli K12 as a function of sequence context. J. Mol. Biol. 220: 271 279.
194. Simonsen, L. 1990. Dynamics of plasmid transfer on surfaces. J. Gen. Microbiol. 136( Pt. 6): 1001 1007.
195. Simpson, A. A.,, Y. Tao,, P. G. Leiman,, M. O. Badasso,, Y. He,, P. J . Jardine,, N. H. Olson,, M. C. Morais,, S. Grimes,, D. L. Anderson,, T. S. Baker,, and M. G. Rossmann. 2000. Structure of the bacteriophage phi29 DNA packaging motor. Nature 408: 745 750.
196. Skurray, R. A.,, and P. Reeves. 1974. F factor-mediated immunity to lethal zygosis in Escherichia coli K-12. J. Bacteriol. 117: 100 106.
197. Smeets, L. C , and J . G. Kusters. 2002. Natural transformation in Helicobacter pylori: DNA transport in an unexpected way. Trends Microbiol. 10: 159 162.
198. Snyder, W. B.,, L. J. Davis,, P. N. Danese,, C. L. Cosma,, and T. J. Silhavy. 1995. Overproduction of NlpE, a new outer membrane lipoprotein, suppresses the toxicity of periplasmic LacZ by activation of the Cpx signal transduction pathway. J. Bacteriol. 177: 4216 4223.
199. Sowa, B. A.,, D. Moore,, and K. Ippen-Ihler. 1983. Physiology of F-pilin synthesis and utilization. J. Bacteriol. 153: 962 968.
200.Stallions, D, R., and R. Curtiss, 3rd. 1972. Bacterial conjugation under anaerobic conditions. J. Bacteriol. 111: 294305.
201. Stockwell, D.,, V. Lelianova,, T. Thompson,, and W. B. Dempsey. 2000. Transcription of the transfer genes traY and traM of the antibiotic resistance plasmid R100-1 is linked. Plasmid 43: 35 48.
202. Strohmaier, H.,, R. Noiges,, S. Kotschan,, G. Sawcrs,, G. Hogenauer,, E. L. Zechner,, and G. Koraimann. 1998. Signal transduction and bacterial conjugation: characterization of the role of ArcA in regulating conjugative transfer of the resistance plasmid R1 J. Mol. Biol. 277: 309 316.
203. Sukupolvi, S.,, and C. D. O'Connor. 1990. TraT lipoprotein, a plasmid-specificd mediator of interactions between gram-negative bacteria and their environment. Microbiol. Rev. 54: 331 341.
204. Sundberg, C.,, L. Meek,, K. Carroll,, A. Das,, and W. Ream. 1996. VirE1 protein mediates export of the single-stranded DNA-binding protein VirE2 from Agrobacterium tumefaciens into plant cells. J. Bacteriol. 178: 1207 1212.
205. Taki, K.,, T. Abo,, and E. Ohtsubo. 1998. Regulatory mechanisms in expression of the traY-I operon of sex factor plasmid R100: involvement of traJ and traY gene products. Genes Cells 3: 331 345.
206. Thomas, C. M. 2000. The Horizontal Gene Pool: Bacterial Plasmids and Gene Spread. Harwood Academic, Amsterdam, The Netherlands.
207. Tomoeda, M.,, M. Inuzuka,, and T. Date. 1975. Bacterial sex pili. Prog. Biopbys. Mol. Biol. 30: 23 56.
208. Torreblanca, J.,, S. Marques,, and J. Casadesus. 1999. Synthesis of FinP RNA by plasmids F and pSLT is regulated by DNA adenine methylation. Genetics 152: 31 45.
209. van Biesen, T.,, and L. S. Frost. 1994. The FinO protein of IncF plasmids binds FinP antisense RNA and its target, traJ mRNA, and promotes duplex formation. Mol. Microbiol. 14: 427 436.
210. Waldor, M. K, 1998. Bacteriophage biology and bacterial virulence. Trends Microbiol. 6: 295 297.
211. Warren, G. J.,, M. W. Saul,, and D.J. Sherratt. 1979. ColEl plasmid mobility: essential and conditional functions. MoL Gen. Genet. 170: 103 107.
212. Waters, V. L. 2001. Conjugation between bacterial and mammalian cells. Nature Genetics 29: 375 376.
213. 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.
214. Wilkins, B., M, 1995. Gene transfer by bacterial conjugation: diversity of systems and functional specializations, p. 59 88. In S. Baumberg,, J. P. W. Young,, E. M. H. Wellington,, and J. R. Saunders (ed.), Population Genetics of Bacteria. Cambridge University Press, Cambridge, United Kingdom.
215. Wilkins, B. M.,, and L. S. Frost,. 2001. Mechanisms of gene exchange between bacteria, p. 355 400. In M. Sussman (ed.), Molecular Medical Microbiology. Academic Press, London, United Kingdom.
216. Wilkins, B. M.,, and A. T. Thomas. 2000. DNA-independent transport of plasmid primase protein between bacteria by the II conjugation system. Mol. Microbiol. 38: 650 657.
217. Willetts, N. 1980. Interactions between the F conjugal transfer system and CloDFl3::Tna plasmids. Mol. Gen. Genet. 180: 213 217.
218. Yamada, Y.,, M. Yamada,, and A. Nakazawa. 1995. A ColE1-encoded gene directs entry exclusion of the plasmid. J. Bacteriol. 177: 6064 6068.
219. Yoshida, T.,, S. R. Kim,, and T. Komano. 1999. Twelve pil genes arc required for biogenesis of the R64 thin pilus. J. Bacteriol. 181: 2038 2043.
220. Zatyka, M.,, L. Bingle,, A. C. Jones,, and C. M, Thomas. 2001. Cooperativity between KorB and TrbA repressors of broad-host-range plasmid RK2. J. Bacteriol. 183: 1022 1031.
221. Zatyka, M.,, G. Jagura-Burdzy,, and C. M. Thomas. 1994. Regulation of transfer genes of promiscuous IncP alpha plasmid RK2: repression of Tra1 region transcription both by relaxosome proteins and by the Tra2 regulator TrbA. Microbiology 140( Pt. 11): 2981 2990.
222. Zechner, E. L.,, F. de la Cruz,, R. Eisenbrandt,, A. M. Grahn,, G, Koraimann, E, Lanka, G. Muth, W. Pansegrau, C. M. Thomas, B. M. Wilkins, and M. Zatyka. 2000. Conjugative DNA transfer processes, p. 87 155. In C. M. Thomas (ed.), The Horizontal Gene Pool: Bacterial Plasmids and Gene Spread. Harwood Academic Publishers, Amsterdam, The Netherlands.
223. 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.
224. Ziegelin, G.,, W. Pansegrau,, R. Lurz,, and E. Lanka. 1992. TraK protein of conjugative plasmid RP4 forms a specialized nucleoprotein complex with the transfer origin. J. Biol. Chem. 267: 17279 17286.
225. Ziemienowicz, A.,, T. Merkle,, F. Schoumacher,, B. Hohn,, and L. Rossi. 2001. Import of Agrobacterium T-DNA into plant nuclei. Two distinct functions of VirD2 and VirE2 proteins. Plant Cell. 13: 369 384.


Generic image for table
Table 1

Characteristics of some self-transmissible plasmids

Citation: Lawley T, Frost L, Wilkins B. 2004. Bacterial Conjugation in Gram-Negative Bacteria , p 203-226. In Funnell B, Phillips G (ed), Plasmid Biology. ASM Press, Washington, DC. doi: 10.1128/9781555817732.ch9
Generic image for table
Table 2

Similarities/homologies shared by transfer and transport/virulence systems

Citation: Lawley T, Frost L, Wilkins B. 2004. Bacterial Conjugation in Gram-Negative Bacteria , p 203-226. In Funnell B, Phillips G (ed), Plasmid Biology. ASM Press, Washington, DC. doi: 10.1128/9781555817732.ch9

This is a required field
Please enter a valid email address
Please check the format of the address you have entered.
Approval was a Success
Invalid data
An Error Occurred
Approval was partially successful, following selected items could not be processed due to error