• [1]
    Solnick, J.V., Schauer, D.B. (2001) Emergence of diverse Helicobacter species in the pathogenesis of gastric and enterohepatic diseases. Clin. Microbiol. Rev. 14, 5997.
  • [2]
    Blaser, M.J., Atherton, J.C. (2004) Helicobacter pylori persistence: biology and disease. J. Clin. Invest. 113, 321333.
  • [3]
    Monack, D.M., Mueller, A., Falkow, S. (2004) Persistent bacterial infections: the interface of the pathogen and the host immune system. Nat. Rev. Microbiol. 2, 747765.
  • [4]
    Falush, D., Wirth, T., Linz, B., Pritchard, J.K., Stephens, M., Kidd, M., Blaser, M.J., Graham, D.Y., Vacher, S., Perez-Perez, G.I., Yamaoka, Y., Megraud, F., Otto, K., Reichard, U., Katzowitsch, E., Wang, X., Achtman, M., Suerbaum, S. (2003) Traces of human migrations in Helicobacter pylori populations. Science 299, 15821585.
  • [5]
    Salama, N., Guillemin, K., McDaniel, T.K., Sherlock, G., Tompkins, L.S., Falkow, S. (2000) A whole-genome microarray reveals genetic diversity among Helicobacter pylori strains. Proc. Natl. Acad. Sci. USA 97, 1466814673.
  • [6]
    Alm, R.A., Ling, L.L., Moir, D.T., King, B.L., Brown, E.D., Doig, P.C., Smith, D.R., Noonan, B., Guild, B.C., DeJonge, B.L., Carmel, G., Tummino, P.J., Caruso, A., Uria-Nickelsen, M., Mills, D.M., Ives, C., Gibson, R., Merberg, D., Mills, S.D., Jiang, Q., Taylor, D.E., Vovis, G.F., Trust, T.J. (1999) Genomic-sequence comparison of two unrelated isolates of the human gastric pathogen Helicobacter pylori. Nature 397, 176180.
  • [7]
    Bjorkholm, B., Sjolund, M., Falk, P.G., Berg, O.G., Engstrand, L., Andersson, D.I. (2001) Mutation frequency and biological cost of antibiotic resistance in Helicobacter pylori. Proc. Natl. Acad. Sci. USA 98, 1460714612.
  • [8]
    Suerbaum, S., Smith, J.M., Bapumia, K., Morelli, G., Smith, N.H., Kunstmann, E., Dyrek, I., Achtman, M. (1998) Free recombination within Helicobacter pylori. Proc. Natl. Acad. Sci. USA 95, 1261912624.
  • [9]
    Aras, R.A., Kang, J., Tschumi, A.I., Harasaki, Y., Blaser, M.J. (2003) Extensive repetitive DNA facilitates prokaryotic genome plasticity. Proc. Natl. Acad. Sci. USA 100, 1357913584.
  • [10]
    Salaun, L., Linz, B., Suerbaum, S., Saunders, N.J. (2004) The diversity within an expanded and redefined repertoire of phase-variable genes in Helicobacter pylori. Microbiology 150, 817830.
  • [11]
    van der Ende, A., Hopman, C.T., Zaat, S., Essink, B.B., Berkhout, B., Dankert, J. (1995) Variable expression of class 1 outer membrane protein in Neisseria meningitidis is caused by variation in the spacing between the +10 and +35 regions of the promoter. J. Bacteriol. 177, 24752480.
  • [12]
    Israel, D.A., Salama, N., Krishna, U., Rieger, U.M., Atherton, J.C., Falkow, S., Peek, R.M. (2001) Helicobacter pylori genetic diversity within the gastric niche of a single human host. Proc. Natl. Acad. Sci. USA 98, 1462514630.
  • [13]
    Lundin, A., Bjorkholm, B., Kupershmidt, I., Unemo, M., Nilsson, P., Andersson, D.I. and Engstrand, L. (2004) When and where is the Helicobacter pylori genome variability generated? In: General Meeting of the American Society of Microbiology, New Orleans, LA (Abstract R-075)
  • [14]
    Kuipers, E.J., Israel, D.A., Kusters, J.G., Gerrits, M.M., Weel, J., van Der Ende, A., van Der Hulst, R.W., Wirth, H.P., Hook-Nikanne, J., Thompson, S.A., Blaser, M.J. (2000) Quasispecies development of Helicobacter pylori observed in paired isolates obtained years apart from the same host. J. Infect. Dis. 181, 273282.
  • [15]
    Cases, I., de Lorenzo, V., Ouzounis, C.A. (2003) Transcription regulation and environmental adaptation in bacteria. Trends Microbiol. 11, 248253.
  • [16]
    Tomb, J.F., White, O., Kerlavage, A.R., Clayton, R.A., Sutton, G.G., Fleischmann, R.D., Ketchum, K.A., Klenk, H.P., Gill, S., Dougherty, B.A., Nelson, K., Quackenbush, J., Zhou, L., Kirkness, E.F., Peterson, S., Loftus, B., Richardson, D., Dodson, R., Khalak, H.G., Glodek, A., McKenney, K., Fitzegerald, L.M., Lee, N., Adams, M.D., Venter, J.C. et-al (1997) The complete genome sequence of the gastric pathogen Helicobacter pylori. Nature 388, 539547.
  • [17]
    Aras, R.A., Fischer, W., Perez-Perez, G.I., Crosatti, M., Ando, T., Haas, R., Blaser, M.J. (2003) Plasticity of repetitive DNA sequences within a bacterial (Type IV) secretion system component. J. Exp. Med. 198, 13491360.
  • [18]
    Robinson, K., Loughlin, M.F., Potter, R., Jenks, P.J. (2005) Host adaptation and immune modulation are mediated by homologous recombination in Helicobacter pylori. J. Infect. Dis. 191, 579587.
  • [19]
    Norris, T.L., Baumler, A.J. (1999) Phase variation of the lpf operon is a mechanism to evade cross-immunity between Salmonella serotypes. Proc. Natl. Acad. Sci. USA 96, 1339313398.
  • [20]
    Nicholson, T.L., Baumler, A.J. (2001) Salmonella enterica serotype typhimurium elicits cross-immunity against a Salmonella enterica serotype enteritidis strain expressing LP fimbriae from the lac promoter. Infect. Immun. 69, 204212.
  • [21]
    Solnick, J.V., Hansen, L.M., Salama, N., Boonjakuakul, J.K., Syvanen, M. (2004) Modification of Helicobacter pylori outer membrane protein expression during experimental infection of rhesus macaques. Proc. Natl. Acad. Sci. USA 101, 21062111.
  • [22]
    Ilver, D., Arnqvist, A., Ogren, J., Frick, I.M., Kersulyte, D., Incecik, E.T., Berg, D.E., Covacci, A., Engstrand, L., Boren, T. (1998) Helicobacter pylori adhesin binding fucosylated histo-blood group antigens revealed by retagging. Science 279, 373377.
  • [23]
    Aspholm-Hurtig, Boren, Dailide, G., Lahmann, M., Kalia, A., Ilver, D., Roche, N., Vikstrom, S., Sjostrom, R., Linden, S., Backstrom, A., Lundberg, C., Arnqvist, A., Mahdavi, J., Nilsson, U.J., Velapatino, B., Gilman, R.H., Gerhard, M., Alarcon, T., Lopez-Brea, M., Nakazawa, T., Fox, J.G., Correa, P., Dominguez-Bello, M.G., Perez-Perez, G.I., Blaser, M.J., Normark, S., Carlstedt, I., Oscarson, S., Teneberg, S., Berg, D.E., Boren, T. (2004) Functional adaptation of BabA, the H. pylori ABO blood group antigen binding adhesin. Science 305, 519522.
  • [24]
    Mahdavi, J., Sonden, B., Hurtig, M., Olfat, F.O., Forsberg, L., Roche, N., Angstrom, J., Larsson, T., Teneberg, S., Karlsson, K.A., Altraja, S., Wadstrom, T., Kersulyte, D., Berg, D.E., Dubois, A., Petersson, C., Magnusson, K.E., Norberg, T., Lindh, F., Lundskog, B.B., Arnqvist, A., Hammarstrom, L., Boren, T. (2002) Helicobacter pylori SabA adhesin in persistent infection and chronic inflammation. Science 297, 573578.
  • [25]
    Backstrom, A., Lundberg, C., Kersulyte, D., Berg, D.E., Boren, T., Arnqvist, A. (2004) Metastability of Helicobacter pylori bab adhesin genes and dynamics in Lewis b antigen binding. Proc. Natl. Acad. Sci. USA 101, 1692316928.
  • [26]
    Viala, J., Chaput, C., Boneca, I.G., Cardona, A., Girardin, S.E., Moran, A.P., Athman, R., Memet, S., Huerre, M.R., Coyle, A.J., DiStefano, P.S., Sansonetti, P.J., Labigne, A., Bertin, J., Philpott, D.J., Ferrero, R.L. (2004) Nod1 responds to peptidoglycan delivered by the Helicobacter pylori cag pathogenicity island. Nat. Immunol. 5, 11661174.
  • [27]
    Aviles-Jimenez, F., Letley, D.P., Gonzalez-Valencia, G., Salama, N., Torres, J., Atherton, J.C. (2004) Evolution of the Helicobacter pylori vacuolating cytotoxin in a human stomach. J. Bacteriol. 186, 51825185.
  • [28]
    Gerhard, M., Lehn, N., Neumayer, N., Borén, T., Rad, R., Schepp, W., Miehlke, S., Classen, M., Prinz, C. (1999) Clinical relevance of the Helicobacter pylori gene for blood-group antigen-binding adhesin. Proc. Natl. Acad. Sci. USA 96, 1277812783.
  • [29]
    Camorlinga-Ponce, M., Romo, C., Gonzalez-Valencia, G., Munoz, O., Torres, J. (2004) Topographical localisation of cagA positive and cagA negative Helicobacter pylori strains in the gastric mucosa; an in situ hybridisation study. J. Clin. Pathol. 57, 822828.
  • [30]
    Garhart, C.A., Redline, R.W., Nedrud, J.G., Czinn, S.J. (2002) Clearance of Helicobacter pylori infection and resolution of postimmunization gastritis in a kinetic study of prophylactically immunized mice. Infect. Immun. 70, 35293538.
  • [31]
    Eaton, K.A., Mefford, M.E. (2001) Cure of Helicobacter pylori infection and resolution of gastritis by adoptive transfer of splenocytes in mice. Infect. Immun. 69, 10251031.
  • [32]
    Ermak, T.H., Giannasca, P.J., Nichols, R., Myers, G.A., Nedrud, J., Weltzin, R., Lee, C.K., Kleanthous, H., Monath, T.P. (1998) Immunization of mice with urease vaccine affords protection against Helicobacter pylori infection in the absence of antibodies and is mediated by MHC class II-restricted responses. J. Exp. Med. 188, 22772288.
  • [33]
    Gewirtz, A.T., Yu, Y., Krishna, U.S., Israel, D.A., Lyons, S.L., Peek Jr, R.M. (2004) Helicobacter pylori flagellin evades toll-like receptor 5-mediated innate immunity. J. Infect. Dis. 189, 19141920.
  • [34]
    Mandell, L., Moran, A.P., Cocchiarella, A., Houghton, J., Taylor, N., Fox, J.G., Wang, T.C., Kurt-Jones, E.A. (2004) Intact Gram-negative Helicobacter pylori, Helicobacter felis, and Helicobacter hepaticus bacteria activate innate immunity via toll-like receptor 2 but not toll-like receptor 4. Infect. Immun. 72, 64466454.
  • [35]
    Netea, M.G., Van der Meer, J.W., Kullberg, B.J. (2004) Toll-like receptors as an escape mechanism from the host defense. Trends Microbiol. 12, 484488.
  • [36]
    Bergman, M.P., Engering, A., Smits, H.H., van Vliet, S.J., van Bodegraven, A.A., Wirth, H.P., Kapsenberg, M.L., Vandenbroucke-Grauls, C.M., van Kooyk, Y., Appelmelk, B.J. (2004) Helicobacter pylori modulates the T helper cell 1/T helper cell 2 balance through phase-variable interaction between lipopolysaccharide and DC-SIGN. J. Exp. Med. 200, 979990.
  • [37]
    Ramarao, N., Gray-Owen, S.D., Backert, S., Meyer, T.F. (2000) Helicobacter pylori inhibits phagocytosis by professional phagocytes involving type IV secretion components. Mol. Microbiol. 37, 13891404.
  • [38]
    Allen, L.A., Schlesinger, L.S., Kang, B. (2000) Virulent strains of Helicobacter pylori demonstrate delayed phagocytosis and stimulate homotypic phagosome fusion in macrophages. J. Exp. Med. 191, 115128.
  • [39]
    Menaker, R.J., Ceponis, P.J., Jones, N.L. (2004) Helicobacter pylori induces apoptosis of macrophages in association with alterations in the mitochondrial pathway. Infect. Immun. 72, 28892898.
  • [40]
    Gobert, A.P., McGee, D.J., Akhtar, M., Mendz, G.L., Newton, J.C., Cheng, Y., Mobley, H.L., Wilson, K.T. (2001) Helicobacter pylori arginase inhibits nitric oxide production by eukaryotic cells: a strategy for bacterial survival. Proc. Natl. Acad. Sci. USA 98, 1384413849.
  • [41]
    Bussiere, F.I., Chaturvedi, R., Cheng, Y., Gobert, A.P., Asim, M., Blumberg, D.R., Xu, H., Kim, P.Y., Hacker, A., Casero Jr, R.A., Wilson, K.T. (2005) Spermine causes loss of innate immune response to Helicobacter pylori by inhibition of inducible nitric-oxide synthase translation. J. Biol. Chem. 280, 24092412.
  • [42]
    Bryk, R., Griffin, P., Nathan, C. (2000) Peroxynitrite reductase activity of bacterial peroxiredoxins. Nature 407, 211215.
  • [43]
    Petersen, A.M., Krogfelt, K.A. (2003) Helicobacter pylori: an invading microorganism. A review. FEMS Immunol. Med. Microbiol. 36, 117126.
  • [44]
    Amieva, M.R., Salama, N.R., Tompkins, L.S., Falkow, S. (2002) Helicobacter pylori enter and survive within multivesicular vacuoles of epithelial cells. Cell Microbiol. 4, 677690.
  • [45]
    Anderson, G.G., Palermo, J.J., Schilling, J.D., Roth, R., Heuser, J., Hultgren, S.J. (2003) Intracellular bacterial biofilm-like pods in urinary tract infections. Science 301, 105107.
  • [46]
    Lundgren, A., Suri-Payer, E., Enarsson, K., Svennerholm, A.M., Lundin, B.S. (2003) Helicobacter pylori-specific CD4+ CD25high regulatory T cells suppress memory T-cell responses to H. pylori in infected individuals. Infect. Immun. 71, 17551762.
  • [47]
    Raghavan, S., Fredriksson, M., Svennerholm, A.M., Holmgren, J., Suri-Payer, E. (2003) Absence of CD4+CD25+ regulatory T cells is associated with a loss of regulation leading to increased pathology in Helicobacter pylori-infected mice. Clin. Exp. Immunol. 132, 393400.
  • [48]
    Lundgren, A., Stromberg, E., Sjoling, A., Lindholm, C., Enarsson, K., Edebo, A., Johnsson, E., Suri-Payer, E., Larsson, P., Rudin, A., Svennerholm, A.M., Lundin, B.S. (2005) Mucosal FOXP3-expressing CD4+CD25 high regulatory T cells in Helicobacter pylori-infected patients. Infect. Immun. 73, 523531.
  • [49]
    Mills, K.H. (2004) Regulatory T cells: friend or foe in immunity to infection. Nat. Rev. Immunol. 4, 841855.
  • [50]
    Boncristiano, M., Paccani, S.R., Barone, S., Ulivieri, C., Patrussi, L., Ilver, D., Amedei, A., D'Elios, M.M., Telford, J.L., Baldari, C.T. (2003) The Helicobacter pylori vacuolating toxin inhibits T cell activation by two independent mechanisms. J. Exp. Med. 198, 18871897.
  • [51]
    Gebert, B., Fischer, W., Weiss, E., Hoffmann, R., Haas, R. (2003) Helicobacter pylori vacuolating cytotoxin inhibits T lymphocyte activation. Science 301, 10991102.
  • [52]
    Sundrud, M.S., Torres, V.J., Unutmaz, D., Cover, T.L. (2004) Inhibition of primary human T cell proliferation by Helicobacter pylori vacuolating toxin (VacA) is independent of VacA effects on IL-2 secretion. Proc. Natl. Acad. Sci. USA 101, 77277732.
  • [53]
    Salama, N., Otto, G., Tompkins, L.S., Falkow, S. (2001) Vacuolating cytotoxin of Helicobacter pylori plays a role during colonization in a mouse model of infection. Infect. Immun. 69, 730736.
  • [54]
    Thomas, L. (1974) The lives of a cell: notes of a biology watcher. p. 76, Viking Press, New York.
  • [55]
    de Martel, C., Llosa, A.E., Farr, S.M., Friedman, G.D., Vogelman, J.H., Orentreich, N., Corley, D.A., Parsonnet, J. (2005) Helicobacter pylori infection and the risk of development of esophageal adenocarcinoma. J. Infect. Dis. 191, 761767.