5. Perspectives
5.3 To identify the phosphorylated and non-phosphorylated RcsB
5.3.1 Identifying the RcsB interacting protein(s) using immunoprecipitation (IP) and mass spectrometry from the total proteins of the bacteria grown specifically under acidic conditions or the conditions optimal for type 3 fimbriae expression.
5.3.2 Study further for the co-regulator mechanisms.
53
6. References
1. Topley WWC, Wilson GS, Collier LH, Balows A, Sussman M, Topley WWC: Topley & Wilson's microbiology and microbial infections, 9th edn. London; New York; Arnold; Oxford University Press; 1998.
2. Bagley ST, Seidler RJ, Talbot HW, Jr., Morrow JE: Isolation of Klebsielleae from within living wood. Appl Environ Microbiol 1978, 36(1):178-185.
3. Brown C, Seidler RJ: Potential pathogens in the environment:
Klebsiella pneumoniae, a taxonomic and ecological enigma. Appl Microbiol 1973, 25(6):900-904.
4. Edberg SC, Piscitelli V, Cartter M: Phenotypic characteristics of coliform and noncoliform bacteria from a public water supply compared with regional and national clinical species. Appl Environ Microbiol 1986, 52(3):474-478.
5. Seidler RJ, Knittel MD, Brown C: Potential pathogens in the environment: cultural reactions and nucleic acid studies on Klebsiella pneumoniae from clinical and environmental sources.
Appl Microbiol 1975, 29(6):819-825.
6. Matsen JM, Spindler JA, Blosser RO: Characterization of
Klebsiella isolates from natural receiving waters and comparison with human isolates. Appl Microbiol 1974, 28(4):672-678.
7. Podschun R, Ullmann U: Klebsiella spp. as nosocomial pathogens:
epidemiology, taxonomy, typing methods, and pathogenicity factors. Clin Microbiol Rev 1998, 11(4):589-603.
54
8. Yinnon AM, Butnaru A, Raveh D, Jerassy Z, Rudensky B:
Klebsiella bacteraemia: community versus nosocomial infection.
QJM 1996, 89(12):933-941.
9. Bartlett JG, O'Keefe P, Tally FP, Louie TJ, Gorbach SL:
Bacteriology of hospital-acquired pneumonia. Arch Intern Med 1986, 146(5):868-871.
10. Jarvis WR, Munn VP, Highsmith AK, Culver DH, Hughes JM: The epidemiology of nosocomial infections caused by Klebsiella pneumoniae. Infect Control 1985, 6(2):68-74.
11. Ko WC, Paterson DL, Sagnimeni AJ, Hansen DS, Von Gottberg A, Mohapatra S, Casellas JM, Goossens H, Mulazimoglu L, Trenholme G et al: Community-acquired Klebsiella pneumoniae bacteremia:
global differences in clinical patterns. Emerg Infect Dis 2002, 8(2):160-166.
12. Paterson DL, Ko WC, Von Gottberg A, Mohapatra S, Casellas JM, Goossens H, Mulazimoglu L, Trenholme G, Klugman KP, Bonomo RA et al: International prospective study of Klebsiella
pneumoniae bacteremia: implications of extended-spectrum beta-lactamase production in nosocomial Infections. Ann Intern Med 2004, 140(1):26-32.
13. Davis TJ, Matsen JM: Prevalence and characteristics of Klebsiella species: relation to association with a hospital environment. J Infect Dis 1974, 130(4):402-405.
14. Selden R, Lee S, Wang WL, Bennett JV, Eickhoff TC: Nosocomial klebsiella infections: intestinal colonization as a reservoir. Ann Intern Med 1971, 74(5):657-664.
55
15. Pollack M, Charache P, Nieman RE, Jett MP, Reimhardt JA, Hardy PH, Jr.: Factors influencing colonisation and antibiotic-resistance patterns of gram-negative bacteria in hospital patients. Lancet 1972, 2(7779):668-671.
16. Rose HD, Schreier J: The effect of hospitalization and antibiotic therapy on the gram-negative fecal flora. Am J Med Sci 1968, 255:228-236.
17. Flidel-Rimon O, Leibovitz E, Juster-Reicher A, Amitay M, Miskin A, Barak Y, Mogilner B: An outbreak of antibiotic multiresistant Klebsiella at the Neonatal Intensive Care Unit, Kaplan Hospital, Rehovot, Israel, November 1991 to April 1992. Am J Perinatol 1996, 13(2):99-102.
18. French GL, Shannon KP, Simmons N: Hospital outbreak of Klebsiella pneumoniae resistant to broad-spectrum
cephalosporins and beta-lactam-beta-lactamase inhibitor combinations by hyperproduction of SHV-5 beta-lactamase. J Clin Microbiol 1996, 34(2):358-363.
19. Keynan Y, Rubinstein E: The changing face of Klebsiella
pneumoniae infections in the community. Int J Antimicrob Agents 2007, 30(5):385-389.
20. Medeiros AA: Nosocomial outbreaks of multiresistant bacteria:
extended-spectrum beta-lactamases have arrived in North America. Ann Intern Med 1993, 119(5):428-430.
21. Meyer KS, Urban C, Eagan JA, Berger BJ, Rahal JJ: Nosocomial outbreak of Klebsiella infection resistant to late-generation cephalosporins. Ann Intern Med 1993, 119(5):353-358.
56
22. Colodner R, Raz R, Chazan B, Sakran W: Susceptibility pattern of extended-spectrum beta-lactamase producing bacteria isolated from inpatients to five antimicrobial drugs in a community hospital in Northern Israel. Int J Antimicrob Agents 2004, 24(4):409-410.
23. Hirsch EB, Tam VH: Detection and treatment options for Klebsiella pneumoniae carbapenemases (KPCs): an emerging cause of multidrug-resistant infection. J Antimicrob Chemother 2010, 65(6):1119-1125.
24. Kumarasamy KK, Toleman MA, Walsh TR, Bagaria J, Butt F, Balakrishnan R, Chaudhary U, Doumith M, Giske CG, Irfan S et al:
Emergence of a new antibiotic resistance mechanism in India, Pakistan, and the UK: a molecular, biological, and
epidemiological study. Lancet Infect Dis 2010, 10(9):597-602.
25. Nordmann P, Cuzon G, Naas T: The real threat of Klebsiella pneumoniae carbapenemase-producing bacteria. Lancet Infect Dis 2009, 9(4):228-236.
26. Yong D, Toleman MA, Giske CG, Cho HS, Sundman K, Lee K, Walsh TR: Characterization of a new metallo-beta-lactamase gene, bla(NDM-1), and a novel erythromycin esterase gene carried on a unique genetic structure in Klebsiella pneumoniae sequence type 14 from India. Antimicrob Agents Chemother 2009, 53(12):5046-5054.
27. [New Delhi-metallo-beta-lactamase 1 (NDM-1)-producing,
multiple drug resistant Enterobacteriaceae]. The Japanese journal of antibiotics 2010, 63(5):365-367.
57
28. McDonald MI: Pyogenic liver abscess: diagnosis, bacteriology and treatment. Eur J Clin Microbiol 1984, 3(6):506-509.
29. Lee CH, Leu HS, Wu TS, Su LH, Liu JW: Risk factors for spontaneous rupture of liver abscess caused by Klebsiella pneumoniae. Diagn Microbiol Infect Dis 2005, 52(2):79-84.
30. Lindstrom ST, Healey PR, Chen SC: Metastatic septic
endophthalmitis complicating pyogenic liver abscess caused by Klebsiella pneumoniae. Aust N Z J Med 1997, 27(1):77-78.
31. Wang JH, Liu YC, Lee SS, Yen MY, Chen YS, Wang JH, Wann SR, Lin HH: Primary liver abscess due to Klebsiella pneumoniae in Taiwan. Clin Infect Dis 1998, 26(6):1434-1438.
32. Chuang YP, Fang CT, Lai SY, Chang SC, Wang JT: Genetic
determinants of capsular serotype K1 of Klebsiella pneumoniae causing primary pyogenic liver abscess. J Infect Dis 2006,
193(5):645-654.
33. Evrard B, Balestrino D, Dosgilbert A, Bouya-Gachancard JL, Charbonnel N, Forestier C, Tridon A: Roles of capsule and lipopolysaccharide O antigen in interactions of human
monocyte-derived dendritic cells and Klebsiella pneumoniae.
Infect Immun 2010, 78(1):210-219.
34. Flannagan RS, Cosio G, Grinstein S: Antimicrobial mechanisms of phagocytes and bacterial evasion strategies. Nat Rev Microbiol 2009, 7(5):355-366.
35. Lin JC, Chang FY, Fung CP, Xu JZ, Cheng HP, Wang JJ, Huang LY, Siu LK: High prevalence of phagocytic-resistant capsular
58
serotypes of Klebsiella pneumoniae in liver abscess. Microbes Infect 2004, 6(13):1191-1198.
36. Campos MA, Vargas MA, Regueiro V, Llompart CM, Alberti S, Bengoechea JA: Capsule polysaccharide mediates bacterial resistance to antimicrobial peptides. Infect Immun 2004, 72(12):7107-7114.
37. Llobet E, Tomas JM, Bengoechea JA: Capsule polysaccharide is a bacterial decoy for antimicrobial peptides. Microbiology 2008, 154(Pt 12):3877-3886.
38. Mizuta K, Ohta M, Mori M, Hasegawa T, Nakashima I, Kato N:
Virulence for mice of Klebsiella strains belonging to the O1
group: relationship to their capsular (K) types. Infect Immun 1983, 40(1):56-61.
39. Lee HC, Chuang YC, Yu WL, Lee NY, Chang CM, Ko NY, Wang LR, Ko WC: Clinical implications of hypermucoviscosity
phenotype in Klebsiella pneumoniae isolates: association with invasive syndrome in patients with community-acquired bacteraemia. J Intern Med 2006, 259(6):606-614.
40. Merino S, Camprubi S, Alberti S, Benedi VJ, Tomas JM:
Mechanisms of Klebsiella pneumoniae resistance to complement-mediated killing. Infect Immun 1992, 60(6):2529-2535.
41. Shankar-Sinha S, Valencia GA, Janes BK, Rosenberg JK, Whitfield C, Bender RA, Standiford TJ, Younger JG: The Klebsiella
pneumoniae O antigen contributes to bacteremia and lethality during murine pneumonia. Infection and immunity 2004,
72(3):1423-1430.
59
42. Medzhitov R, Janeway C, Jr.: Innate immune recognition:
mechanisms and pathways. Immunological reviews 2000, 173:89-97.
43. Stahlhut SG, Tchesnokova V, Struve C, Weissman SJ,
Chattopadhyay S, Yakovenko O, Aprikian P, Sokurenko EV, Krogfelt KA: Comparative structure-function analysis of mannose-specific FimH adhesins from Klebsiella pneumoniae and Escherichia coli. J Bacteriol 2009, 191(21):6592-6601.
44. Struve C, Bojer M, Krogfelt KA: Characterization of Klebsiella pneumoniae type 1 fimbriae by detection of phase variation during colonization and infection and impact on virulence. Infect Immun 2008, 76(9):4055-4065.
45. Struve C, Bojer M, Krogfelt KA: Identification of a conserved chromosomal region encoding Klebsiella pneumoniae type 1 and type 3 fimbriae and assessment of the role of fimbriae in
pathogenicity. Infect Immun 2009, 77(11):5016-5024.
46. Huang YJ, Liao HW, Wu CC, Peng HL: MrkF is a component of type 3 fimbriae in Klebsiella pneumoniae. Res Microbiol 2009, 160(1):71-79.
47. Langstraat J, Bohse M, Clegg S: Type 3 fimbrial shaft (MrkA) of Klebsiella pneumoniae, but not the fimbrial adhesin (MrkD), facilitates biofilm formation. Infect Immun 2001, 69(9):5805-5812.
48. Stahlhut SG, Chattopadhyay S, Struve C, Weissman SJ, Aprikian P, Libby SJ, Fang FC, Krogfelt KA, Sokurenko EV: Population
variability of the FimH type 1 fimbrial adhesin in Klebsiella pneumoniae. J Bacteriol 2009, 191(6):1941-1950.
60
49. Di Martino P, Bertin Y, Girardeau JP, Livrelli V, Joly B, Darfeuille-Michaud A: Molecular characterization and adhesive properties of CF29K, an adhesin of Klebsiella pneumoniae strains involved in nosocomial infections. Infect Immun 1995, 63(11):4336-4344.
50. Di Martino P, Livrelli V, Sirot D, Joly B, Darfeuille-Michaud A: A new fimbrial antigen harbored by CAZ-5/SHV-4-producing Klebsiella pneumoniae strains involved in nosocomial infections.
Infect Immun 1996, 64(6):2266-2273.
51. Di Martino P, Sirot D, Joly B, Rich C, Darfeuille-Michaud A:
Relationship between adhesion to intestinal Caco-2 cells and multidrug resistance in Klebsiella pneumoniae clinical isolates. J Clin Microbiol 1997, 35(6):1499-1503.
52. Lawlor MS, Hsu J, Rick PD, Miller VL: Identification of Klebsiella pneumoniae virulence determinants using an intranasal infection model. Molecular microbiology 2005, 58(4):1054-1073.
53. Clegg S, Purcell BK, Pruckler J: Characterization of genes
encoding type 1 fimbriae of Klebsiella pneumoniae, Salmonella typhimurium, and Serratia marcescens. Infection and immunity 1987, 55(2):281-287.
54. Gerlach GF, Clegg S, Allen BL: Identification and
characterization of the genes encoding the type 3 and type 1 fimbrial adhesins of Klebsiella pneumoniae. Journal of bacteriology 1989, 171(3):1262-1270.
55. Schembri MA, Blom J, Krogfelt KA, Klemm P: Capsule and fimbria interaction in Klebsiella pneumoniae. Infection and immunity 2005, 73(8):4626-4633.
61
56. Ryan RP, Fouhy Y, Lucey JF, Crossman LC, Spiro S, He YW, Zhang LH, Heeb S, Camara M, Williams P et al: Cell-cell signaling in Xanthomonas campestris involves an HD-GYP domain
protein that functions in cyclic di-GMP turnover. Proceedings of the National Academy of Sciences of the United States of America 2006, 103(17):6712-6717.
57. Matatov R, Goldhar J, Skutelsky E, Sechter I, Perry R, Podschun R, Sahly H, Thankavel K, Abraham SN, Ofek I: Inability of
encapsulated Klebsiella pneumoniae to assemble functional type 1 fimbriae on their surface. FEMS microbiology letters 1999, 179(1):123-130.
58. Sahly H, Podschun R, Oelschlaeger TA, Greiwe M, Parolis H, Hasty D, Kekow J, Ullmann U, Ofek I, Sela S: Capsule impedes adhesion to and invasion of epithelial cells by Klebsiella pneumoniae.
Infection and immunity 2000, 68(12):6744-6749.
59. Boddicker JD, Anderson RA, Jagnow J, Clegg S: Signature-tagged mutagenesis of Klebsiella pneumoniae to identify genes that influence biofilm formation on extracellular matrix material.
Infection and immunity 2006, 74(8):4590-4597.
60. Burmolle M, Bahl MI, Jensen LB, Sorensen SJ, Hansen LH: Type 3 fimbriae, encoded by the conjugative plasmid pOLA52, enhance biofilm formation and transfer frequencies in
Enterobacteriaceae strains. Microbiology 2008, 154(Pt 1):187-195.
61. Di Martino P, Cafferini N, Joly B, Darfeuille-Michaud A: Klebsiella pneumoniae type 3 pili facilitate adherence and biofilm
62
formation on abiotic surfaces. Research in microbiology 2003, 154(1):9-16.
62. Jagnow J, Clegg S: Klebsiella pneumoniae MrkD-mediated biofilm formation on extracellular matrix- and collagen-coated surfaces. Microbiology 2003, 149(Pt 9):2397-2405.
63. Ong CL, Beatson SA, McEwan AG, Schembri MA: Conjugative plasmid transfer and adhesion dynamics in an Escherichia coli biofilm. Applied and environmental microbiology 2009,
75(21):6783-6791.
64. Schroll C, Barken KB, Krogfelt KA, Struve C: Role of type 1 and type 3 fimbriae in Klebsiella pneumoniae biofilm formation.
BMC microbiology 2010, 10:179.
65. Sutherland I: Biofilm exopolysaccharides: a strong and sticky framework. Microbiology 2001, 147(Pt 1):3-9.
66. Warren JW: Catheter-associated urinary tract infections.
International journal of antimicrobial agents 2001, 17(4):299-303.
67. Maki DG, Tambyah PA: Engineering out the risk for infection with urinary catheters. Emerging infectious diseases 2001, 7(2):342-347.
68. Ronald A: The etiology of urinary tract infection: traditional and emerging pathogens. Disease-a-month : DM 2003, 49(2):71-82.
69. Hornick DB, Allen BL, Horn MA, Clegg S: Adherence to
respiratory epithelia by recombinant Escherichia coli expressing Klebsiella pneumoniae type 3 fimbrial gene products. Infect Immun 1992, 60(4):1577-1588.
63
70. Huang YJ, Wu CC, Chen MC, Fung CP, Peng HL:
Characterization of the type 3 fimbriae with different MrkD adhesins: possible role of the MrkD containing an RGD motif.
Biochemical and biophysical research communications 2006, 350(3):537-542.
71. Sebghati TA, Clegg S: Construction and characterization of
mutations within the Klebsiella mrkD1P gene that affect binding to collagen type V. Infection and immunity 1999, 67(4):1672-1676.
72. Tarkkanen AM, Allen BL, Westerlund B, Holthofer H, Kuusela P, Risteli L, Clegg S, Korhonen TK: Type V collagen as the target for type-3 fimbriae, enterobacterial adherence organelles. Mol
Microbiol 1990, 4(8):1353-1361.
73. Tarkkanen AM, Virkola R, Clegg S, Korhonen TK: Binding of the type 3 fimbriae of Klebsiella pneumoniae to human endothelial and urinary bladder cells. Infect Immun 1997, 65(4):1546-1549.
74. Allen BL, Gerlach GF, Clegg S: Nucleotide sequence and
functions of mrk determinants necessary for expression of type 3 fimbriae in Klebsiella pneumoniae. J Bacteriol 1991, 173(2):916-920.
75. Duguid JP: Fimbriae and adhesive properties in Klebsiella strains.
J Gen Microbiol 1959, 21:271-286.
76. Castanie-Cornet MP, Treffandier H, Francez-Charlot A, Gutierrez C, Cam K: The glutamate-dependent acid resistance system in
Escherichia coli: essential and dual role of the His-Asp
phosphorelay RcsCDB/AF. Microbiology 2007, 153(Pt 1):238-246.
64
77. Imlay JA: Pathways of oxidative damage. Annu Rev Microbiol 2003, 57:395-418.
78. Galperin MY, Higdon R, Kolker E: Interplay of heritage and habitat in the distribution of bacterial signal transduction systems. Mol Biosyst 2010, 6(4):721-728.
79. Perry J, Koteva K, Wright G: Receptor domains of two-component signal transduction systems. Mol Biosyst 2011, 7(5):1388-1398.
80. Laub MT, Goulian M: Specificity in two-component signal transduction pathways. Annu Rev Genet 2007, 41:121-145.
81. Majdalani N, Gottesman S: The Rcs phosphorelay: a complex signal transduction system. Annu Rev Microbiol 2005, 59:379-405.
82. Virlogeux I, Waxin H, Ecobichon C, Lee JO, Popoff MY:
Characterization of the rcsA and rcsB genes from Salmonella typhi: rcsB through tviA is involved in regulation of Vi antigen synthesis. J Bacteriol 1996, 178(6):1691-1698.
83. McCallum KL, Whitfield C: The rcsA gene of Klebsiella
pneumoniae O1:K20 is involved in expression of the serotype-specific K (capsular) antigen. Infect Immun 1991, 59(2):494-502.
84. Bernhard F, Poetter K, Geider K, Coplin DL: The rcsA gene from Erwinia amylovora: identification, nucleotide sequence, and regulation of exopolysaccharide biosynthesis. Mol Plant Microbe Interact 1990, 3(6):429-437.
85. Bereswill S, Geider K: Characterization of the rcsB gene from Erwinia amylovora and its influence on exoploysaccharide synthesis and virulence of the fire blight pathogen. J Bacteriol 1997, 179(4):1354-1361.
65
86. Poetter K, Coplin DL: Structural and functional analysis of the rcsA gene from Erwinia stewartii. Mol Gen Genet 1991,
229(1):155-160.
87. Stout V, Gottesman S: RcsB and RcsC: a two-component regulator of capsule synthesis in Escherichia coli. J Bacteriol 1990, 172(2):659-669.
88. Carballes F, Bertrand C, Bouche JP, Cam K: Regulation of Escherichia coli cell division genes ftsA and ftsZ by the two-component system rcsC-rcsB. Mol Microbiol 1999, 34(3):442-450.
89. Francez-Charlot A, Laugel B, Van Gemert A, Dubarry N, Wiorowski F, Castanie-Cornet MP, Gutierrez C, Cam K: RcsCDB His-Asp phosphorelay system negatively regulates the flhDC operon in Escherichia coli. Mol Microbiol 2003, 49(3):823-832.
90. Schwan WR, Shibata S, Aizawa S, Wolfe AJ: The two-component response regulator RcsB regulates type 1 piliation in Escherichia coli. J Bacteriol 2007, 189(19):7159-7163.
91. Cano DA, Dominguez-Bernal G, Tierrez A, Garcia-Del Portillo F, Casadesus J: Regulation of capsule synthesis and cell motility in Salmonella enterica by the essential gene igaA. Genetics 2002, 162(4):1513-1523.
92. Garcia-Calderon CB, Casadesus J, Ramos-Morales F: Rcs and PhoPQ regulatory overlap in the control of Salmonella enterica virulence. J Bacteriol 2007, 189(18):6635-6644.
93. Erickson KD, Detweiler CS: The Rcs phosphorelay system is specific to enteric pathogens/commensals and activates ydeI, a
66
gene important for persistent Salmonella infection of mice. Mol Microbiol 2006, 62(3):883-894.
94. Wacharotayankun R, Arakawa Y, Ohta M, Hasegawa T, Mori M, Horii T, Kato N: Involvement of rcsB in Klebsiella K2 capsule synthesis in Escherichia coli K-12. J Bacteriol 1992, 174(3):1063-1067.
95. Nassif X, Honore N, Vasselon T, Cole ST, Sansonetti PJ: Positive control of colanic acid synthesis in Escherichia coli by rmpA and rmpB, two virulence-plasmid genes of Klebsiella pneumoniae.
Mol Microbiol 1989, 3(10):1349-1359.
96. Nassif X, Fournier JM, Arondel J, Sansonetti PJ: Mucoid phenotype of Klebsiella pneumoniae is a plasmid-encoded virulence factor.
Infect Immun 1989, 57(2):546-552.
97. Jayaratne P, Keenleyside WJ, MacLachlan PR, Dodgson C,
Whitfield C: Characterization of rcsB and rcsC from Escherichia coli O9:K30:H12 and examination of the role of the rcs
regulatory system in expression of group I capsular polysaccharides. J Bacteriol 1993, 175(17):5384-5394.
98. Chen MH, Takeda S, Yamada H, Ishii Y, Yamashino T, Mizuno T:
Characterization of the RcsC-->YojN-->RcsB phosphorelay signaling pathway involved in capsular synthesis in Escherichia coli. Biosci Biotechnol Biochem 2001, 65(10):2364-2367.
99. Latasa C, Garcia B, Echeverz M, Toledo-Arana A, Valle J, Campoy S, Garcia-del Portillo F, Solano C, Lasa I: Salmonella biofilm development depends on the phosphorylation status of RcsB. J Bacteriol 2012, 194(14):3708-3722.
67
100. Wehland M, Kiecker C, Coplin DL, Kelm O, Saenger W, Bernhard F:
Identification of an RcsA/RcsB recognition motif in the promoters of exopolysaccharide biosynthetic operons from
Erwinia amylovora and Pantoea stewartii subspecies stewartii. J Biol Chem 1999, 274(6):3300-3307.
101. Ebel W, Trempy JE: Escherichia coli RcsA, a positive activator of colanic acid capsular polysaccharide synthesis, functions To activate its own expression. J Bacteriol 1999, 181(2):577-584.
102. Wehland M, Bernhard F: The RcsAB box. Characterization of a new operator essential for the regulation of exopolysaccharide biosynthesis in enteric bacteria. J Biol Chem 2000, 275(10):7013-7020.
103. Pristovsek P, Sengupta K, Lohr F, Schafer B, von Trebra MW, Ruterjans H, Bernhard F: Structural analysis of the DNA-binding domain of the Erwinia amylovora RcsB protein and its
interaction with the RcsAB box. J Biol Chem 2003, 278(20):17752-17759.
104. Rahn A, Whitfield C: Transcriptional organization and regulation of the Escherichia coli K30 group 1 capsule biosynthesis (cps) gene cluster. Mol Microbiol 2003, 47(4):1045-1060.
105. Russo TA, Singh G: An extraintestinal, pathogenic isolate of Escherichia coli (O4/K54/H5) can produce a group 1 capsule which is divergently regulated from its constitutively produced group 2, K54 capsular polysaccharide. J Bacteriol 1993,
175(23):7617-7623.
68
106. Whitfield C, Roberts IS: Structure, assembly and regulation of expression of capsules in Escherichia coli. Mol Microbiol 1999, 31(5):1307-1319.
107. Arricau N, Hermant D, Waxin H, Ecobichon C, Duffey PS, Popoff MY: The RcsB-RcsC regulatory system of Salmonella typhi differentially modulates the expression of invasion proteins, flagellin and Vi antigen in response to osmolarity. Mol Microbiol 1998, 29(3):835-850.
108. Johnson MD, Burton NA, Gutierrez B, Painter K, Lund PA: RcsB is required for inducible acid resistance in Escherichia coli and acts at gadE-dependent and -independent promoters. J Bacteriol 2011, 193(14):3653-3656.
109. Venkatesh GR, Kembou Koungni FC, Paukner A, Stratmann T, Blissenbach B, Schnetz K: BglJ-RcsB heterodimers relieve
repression of the Escherichia coli bgl operon by H-NS. J Bacteriol 2010, 192(24):6456-6464.
110. Cheng H-Y: Regulation of Two-component Systems in the Polymyxin B Resistance and Capsular Polysaccharide
Biosynthesis in Klebsiella pneumoniae CG43. PhD thesis. Taiwan R.O.C.: National Chiao Tung University; 2010.
111. Wu C-C: Prevalence Analysis and Expressional Control of Klebsiella pneumoniae Fimbriae. PhD thesis. Taiwan R.O.C.:
National Chiao Tung University; 2010.
112. Lin C-H: Role of RcsB in acid-resistance in Klebsiella
pneumoniae CG43. Master thesis. Taiwan R.O.C.: National Chiao Tung University; 2009.
69
113. Fang CT, Chuang YP, Shun CT, Chang SC, Wang JT: A novel virulence gene in Klebsiella pneumoniae strains causing primary liver abscess and septic metastatic complications. J Exp Med 2004, 199(5):697-705.
114. Lai YC, Peng HL, Chang HY: RmpA2, an activator of capsule biosynthesis in Klebsiella pneumoniae CG43, regulates K2 cps gene expression at the transcriptional level. J Bacteriol 2003, 185(3):788-800.
115. Cheng HY, Chen YS, Wu CY, Chang HY, Lai YC, Peng HL: RmpA regulation of capsular polysaccharide biosynthesis in Klebsiella pneumoniae CG43. J Bacteriol 2010, 192(12):3144-3158.
116. Gupte G, Woodward C, Stout V: Isolation and characterization of rcsB mutations that affect colanic acid capsule synthesis in Escherichia coli K-12. J Bacteriol 1997, 179(13):4328-4335.
117. Stock AM, Robinson VL, Goudreau PN: Two-component signal transduction. Annu Rev Biochem 2000, 69:183-215.
118. Volz K: Structural conservation in the CheY superfamily.
Biochemistry 1993, 32(44):11741-11753.
119. Lukat GS, Stock AM, Stock JB: Divalent metal ion binding to the CheY protein and its significance to phosphotransfer in bacterial chemotaxis. Biochemistry 1990, 29(23):5436-5442.
120. Chang HY, Lee JH, Deng WL, Fu TF, Peng HL: Virulence and outer membrane properties of a galU mutant of Klebsiella pneumoniae CG43. Microb Pathog 1996, 20(5):255-261.
70
121. Domenico P, Schwartz S, Cunha BA: Reduction of capsular polysaccharide production in Klebsiella pneumoniae by sodium salicylate. Infect Immun 1989, 57(12):3778-3782.
122. Blumer C, Kleefeld A, Lehnen D, Heintz M, Dobrindt U, Nagy G, Michaelis K, Emody L, Polen T, Rachel R et al: Regulation of type 1 fimbriae synthesis and biofilm formation by the transcriptional regulator LrhA of Escherichia coli. Microbiology 2005, 151(Pt 10):3287-3298.
123. Hall HK, Foster JW: The role of fur in the acid tolerance response of Salmonella typhimurium is physiologically and genetically separable from its role in iron acquisition. J Bacteriol 1996, 178(19):5683-5691.
124. Lin CT, Huang TY, Liang WC, Peng HL: Homologous response regulators KvgA, KvhA and KvhR regulate the synthesis of capsular polysaccharide in Klebsiella pneumoniae CG43 in a coordinated manner. J Biochem 2006, 140(3):429-438.
125. Miller JH: Experiments in Molecular Genetics. . Cold Spring Harbor Laboratory Press 1972.
126. Maris AE, Sawaya MR, Kaczor-Grzeskowiak M, Jarvis MR, Bearson SM, Kopka ML, Schroder I, Gunsalus RP, Dickerson RE:
Dimerization allows DNA target site recognition by the NarL response regulator. Nat Struct Biol 2002, 9(10):771-778.
127. Cheng HP, Walker GC: Succinoglycan production by Rhizobium meliloti is regulated through the ExoS-ChvI two-component regulatory system. J Bacteriol 1998, 180(1):20-26.
71
128. Lukat GS, McCleary WR, Stock AM, Stock JB: Phosphorylation of bacterial response regulator proteins by low molecular weight phospho-donors. Proc Natl Acad Sci U S A 1992, 89(2):718-722.
129. Schroder I, Wolin CD, Cavicchioli R, Gunsalus RP:
Phosphorylation and dephosphorylation of the NarQ, NarX, and NarL proteins of the nitrate-dependent two-component
regulatory system of Escherichia coli. J Bacteriol 1994, 176(16):4985-4992.
130. McCleary WR, Stock JB, Ninfa AJ: Is acetyl phosphate a global signal in Escherichia coli? J Bacteriol 1993, 175(10):2793-2798.
131. Dahl MK, Msadek T, Kunst F, Rapoport G: Mutational analysis of the Bacillus subtilis DegU regulator and its phosphorylation by the DegS protein kinase. J Bacteriol 1991, 173(8):2539-2547.
132. Ma S, Selvaraj U, Ohman DE, Quarless R, Hassett DJ, Wozniak DJ:
Phosphorylation-independent activity of the response regulators AlgB and AlgR in promoting alginate biosynthesis in mucoid Pseudomonas aeruginosa. J Bacteriol 1998, 180(4):956-968.
133. Gueriri I, Bay S, Dubrac S, Cyncynatus C, Msadek T: The
133. Gueriri I, Bay S, Dubrac S, Cyncynatus C, Msadek T: The