Timothy L. Cover, M.D.


Faculty Appointments
Professor of Medicine Professor of Pathology, Microbiology and Immunology
M.D., Duke University, Durham, North CarolinaB.S., Muhlenberg College, Allentown, Pennsylvania
Office Address
Division of Infectious Diseases
A2200 Medical Center North
Vanderbilt University School of Medicine
Nashville, TN 37232
Research Description
Research projects in the Cover laboratory are focused on investigating molecular mechanisms by which pathogenic bacteria cause disease in humans. We are particularly interested in the bacterium Helicobacter pylori, which contributes to the development of stomach cancer and peptic ulcer disease. H. pylori persistently colonizes the acidic gastric mucosa of humans, and persistent H. pylori colonization of the stomach results in chronic gastric inflammation. H. pylori is the only bacterial pathogen that commonly causes cancer in humans, and stomach cancer is the second leading cause of cancer mortality worldwide.

To investigate molecular mechanisms by which H. pylori causes disease, our laboratory uses a variety of molecular biology, cell biology, genomic, and immunologic methods. Current projects include (i) investigation of two H. pylori toxins (VacA and CagA) in cell culture and animal models, (ii) investigation of the H. pylori type IV secretion system that delivers CagA into gastric epithelial cells, (iii) investigation of mechanisms by which gene expression is regulated in H. pylori, and (iv) comparative genome sequence analysis of H. pylori strains isolated from patients with different disease states, with the goal of identifying new bacterial biomarkers that predict the development of gastric cancer.

Research Keywords
Bacteria-host interactions, bacterial toxins, Helicobacter pylori, gastric cancer, pathogenesis of bacterial infections, bacterial protein secretion
Clinical Research Keywords
Persistent bacterial infections, Immune evasion, gastric cancer, Helicobacter pylori, Bacterial toxins
Snider CA, Voss BJ, McDonald WH, Cover TL. Growth phase-dependent composition of the Helicobacter pylori exoproteome. J Proteomics [print-electronic]. 2016 Jan 1/1/2016; 130: 94-107. PMID: 26363098, PMCID: PMC4640983, PII: S1874-3919(15)30116-0, DOI: 10.1016/j.jprot.2015.08.025, ISSN: 1876-7737.

Frick-Cheng AE, Pyburn TM, Voss BJ, McDonald WH, Ohi MD, Cover TL. Molecular and Structural Analysis of the Helicobacter pylori cag Type IV Secretion System Core Complex. MBio. 2016; 7(1): PMID: 26758182, PMCID: PMC4725015, PII: mBio.02001-15, DOI: 10.1128/mBio.02001-15, ISSN: 2150-7511.

Cover TL. Helicobacter pylori Diversity and Gastric Cancer Risk. MBio. 2016; 7(1): PMID: 26814181, PII: mBio.01869-15, DOI: 10.1128/mBio.01869-15, ISSN: 2150-7511.

Loh JT, Gaddy JA, Algood HM, Gaudieri S, Mallal S, Cover TL. Helicobacter pylori Adaptation In Vivo in Response to a High-Salt Diet. Infect. Immun [print-electronic]. 2015 Dec; 83(12): 4871-83. PMID: 26438795, PMCID: PMC4645402, PII: IAI.00918-15, DOI: 10.1128/IAI.00918-15, ISSN: 1098-5522.

Voss BJ, Loh JT, Hill S, Rose KL, McDonald WH, Cover TL. Alteration of the Helicobacter pylori membrane proteome in response to changes in environmental salt concentration. Proteomics Clin Appl [print-electronic]. 2015 Dec; 9(11-12): 1021-34. PMID: 26109032, PMCID: PMC4690801, DOI: 10.1002/prca.201400176, ISSN: 1862-8354.

Noto JM, Lee JY, Gaddy JA, Cover TL, Amieva MR, Peek RM. Regulation of Helicobacter pylori Virulence Within the Context of Iron Deficiency. J. Infect. Dis [print-electronic]. 2015 Jun 6/1/2015; 211(11): 1790-4. PMID: 25505301, PMCID: PMC4447831, PII: jiu805, DOI: 10.1093/infdis/jiu805, ISSN: 1537-6613.

Gaddy JA, Radin JN, Loh JT, Piazuelo MB, Kehl-Fie TE, Delgado AG, Ilca FT, Peek RM, Cover TL, Chazin WJ, Skaar EP, Scott Algood HM. The host protein calprotectin modulates the Helicobacter pylori cag type IV secretion system via zinc sequestration. PLoS Pathog. 2014 Oct; 10(10): e1004450. PMID: 25330071, PMCID: PMC4199781, PII: PPATHOGENS-D-14-01130, DOI: 10.1371/journal.ppat.1004450, ISSN: 1553-7374.

Engler DB, Reuter S, van Wijck Y, Urban S, Kyburz A, Maxeiner J, Martin H, Yogev N, Waisman A, Gerhard M, Cover TL, Taube C, Müller A. Effective treatment of allergic airway inflammation with Helicobacter pylori immunomodulators requires BATF3-dependent dendritic cells and IL-10. Proc. Natl. Acad. Sci. U.S.A [print-electronic]. 2014 Aug 8/12/2014; 111(32): 11810-5. PMID: 25074917, PMCID: PMC4136625, PII: 1410579111, DOI: 10.1073/pnas.1410579111, ISSN: 1091-6490.

Johnson EM, Gaddy JA, Voss BJ, Hennig EE, Cover TL. Genes required for assembly of pili associated with the Helicobacter pylori cag type IV secretion system. Infect. Immun [print-electronic]. 2014 Aug; 82(8): 3457-70. PMID: 24891108, PMCID: PMC4136203, PII: IAI.01640-14, DOI: 10.1128/IAI.01640-14, ISSN: 1098-5522.

Voss BJ, Gaddy JA, McDonald WH, Cover TL. Analysis of surface-exposed outer membrane proteins in Helicobacter pylori. J. Bacteriol [print-electronic]. 2014 Jul; 196(13): 2455-71. PMID: 24769695, PMCID: PMC4054173, PII: JB.01768-14, DOI: 10.1128/JB.01768-14, ISSN: 1098-5530.

Kodaman N, Pazos A, Schneider BG, Piazuelo MB, Mera R, Sobota RS, Sicinschi LA, Shaffer CL, Romero-Gallo J, de Sablet T, Harder RH, Bravo LE, Peek RM, Wilson KT, Cover TL, Williams SM, Correa P. Human and Helicobacter pylori coevolution shapes the risk of gastric disease. Proc. Natl. Acad. Sci. U.S.A [print-electronic]. 2014 Jan 1/28/2014; 111(4): 1455-60. PMID: 24474772, PMCID: PMC3910595, PII: 1318093111, DOI: 10.1073/pnas.1318093111, ISSN: 1091-6490.

Radin JN, González-Rivera C, Frick-Cheng AE, Sheng J, Gaddy JA, Rubin DH, Algood HM, McClain MS, Cover TL. Role of connexin 43 in Helicobacter pylori VacA-induced cell death. Infect. Immun [print-electronic]. 2014 Jan; 82(1): 423-32. PMID: 24191302, PMCID: PMC3911829, PII: IAI.00827-13, DOI: 10.1128/IAI.00827-13, ISSN: 1098-5522.

McClain MS, Duncan SS, Gaddy JA, Cover TL. Control of gene expression in Helicobacter pylori using the Tet repressor. J. Microbiol. Methods [print-electronic]. 2013 Dec; 95(3): 336-41. PMID: 24113399, PMCID: PMC3856897, PII: S0167-7012(13)00302-3, DOI: 10.1016/j.mimet.2013.09.019, ISSN: 1872-8359.

Cover TL, Peek RM. Diet, microbial virulence, and Helicobacter pylori-induced gastric cancer. Gut Microbes [print-electronic]. 2013 Nov; 4(6): 482-93. PMID: 23989802, PMCID: PMC3928160, PII: 26262, DOI: 10.4161/gmic.26262, ISSN: 1949-0984.

Gaddy JA, Radin JN, Loh JT, Zhang F, Washington MK, Peek RM, Algood HM, Cover TL. High dietary salt intake exacerbates Helicobacter pylori-induced gastric carcinogenesis. Infect. Immun [print-electronic]. 2013 Jun; 81(6): 2258-67. PMID: 23569116, PMCID: PMC3676043, PII: IAI.01271-12, DOI: 10.1128/IAI.01271-12, ISSN: 1098-5522.

Parekh VV, Wu L, Boyd KL, Williams JA, Gaddy JA, Olivares-Villagómez D, Cover TL, Zong WX, Zhang J, Van Kaer L. Impaired autophagy, defective T cell homeostasis, and a wasting syndrome in mice with a T cell-specific deletion of Vps34. J. Immunol [print-electronic]. 2013 May 5/15/2013; 190(10): 5086-101. PMID: 23596309, PMCID: PMC3646937, PII: jimmunol.1202071, DOI: 10.4049/jimmunol.1202071, ISSN: 1550-6606.

Chambers MG, Pyburn TM, González-Rivera C, Collier SE, Eli I, Yip CK, Takizawa Y, Lacy DB, Cover TL, Ohi MD. Structural analysis of the oligomeric states of Helicobacter pylori VacA toxin. J. Mol. Biol [print-electronic]. 2013 Feb 2/8/2013; 425(3): 524-35. PMID: 23178866, PMCID: PMC3612943, PII: S0022-2836(12)00892-3, DOI: 10.1016/j.jmb.2012.11.020, ISSN: 1089-8638.

Barrozo RM, Cooke CL, Hansen LM, Lam AM, Gaddy JA, Johnson EM, Cariaga TA, Suarez G, Peek RM, Cover TL, Solnick JV. Functional plasticity in the type IV secretion system of Helicobacter pylori. PLoS Pathog [print-electronic]. 2013 Feb; 9(2): e1003189. PMID: 23468628, PMCID: PMC3585145, PII: PPATHOGENS-D-12-01776, DOI: 10.1371/journal.ppat.1003189, ISSN: 1553-7374.

Noto JM, Gaddy JA, Lee JY, Piazuelo MB, Friedman DB, Colvin DC, Romero-Gallo J, Suarez G, Loh J, Slaughter JC, Tan S, Morgan DR, Wilson KT, Bravo LE, Correa P, Cover TL, Amieva MR, Peek RM. Iron deficiency accelerates Helicobacter pylori-induced carcinogenesis in rodents and humans. J. Clin. Invest [print-electronic]. 2013 Jan; 123(1): 479-92. PMID: 23257361, PMCID: PMC3533289, PII: 64373, DOI: 10.1172/JCI64373, ISSN: 1558-8238.

Duncan SS, Valk PL, McClain MS, Shaffer CL, Metcalf JA, Bordenstein SR, Cover TL. Comparative genomic analysis of East Asian and non-Asian Helicobacter pylori strains identifies rapidly evolving genes. PLoS ONE [print-electronic]. 2013; 8(1): e55120. PMID: 23383074, PMCID: PMC3561388, PII: PONE-D-12-28737, DOI: 10.1371/journal.pone.0055120, ISSN: 1932-6203.

Duncan SS, Bertoli MT, Kersulyte D, Valk PL, Tamma S, Segal I, McClain MS, Cover TL, Berg DE. Genome Sequences of Three hpAfrica2 Strains of Helicobacter pylori. Genome Announc. 2013; 1(5): PMID: 24072860, PMCID: PMC3784780, PII: 1/5/e00729-13, DOI: 10.1128/genomeA.00729-13, ISSN: 2169-8287.

Radin JN, Gaddy JA, González-Rivera C, Loh JT, Algood HM, Cover TL. Flagellar localization of a Helicobacter pylori autotransporter protein. MBio. 2013; 4(2): e00613-12. PMID: 23572556, PMCID: PMC3622936, PII: mBio.00613-12, DOI: 10.1128/mBio.00613-12, ISSN: 2150-7511.

Loh JT, Friedman DB, Piazuelo MB, Bravo LE, Wilson KT, Peek RM, Correa P, Cover TL. Analysis of Helicobacter pylori cagA promoter elements required for salt-induced upregulation of CagA expression. Infect. Immun [print-electronic]. 2012 Sep; 80(9): 3094-106. PMID: 22710874, PMCID: PMC3418733, PII: IAI.00232-12, DOI: 10.1128/IAI.00232-12, ISSN: 1098-5522.

González-Rivera C, Algood HM, Radin JN, McClain MS, Cover TL. The intermediate region of Helicobacter pylori VacA is a determinant of toxin potency in a Jurkat T cell assay. Infect. Immun [print-electronic]. 2012 Aug; 80(8): 2578-88. PMID: 22585965, PMCID: PMC3434591, PII: IAI.00052-12, DOI: 10.1128/IAI.00052-12, ISSN: 1098-5522.

Cover TL. Role of Helicobacter pylori CagL in modulating gastrin expression. Gut [print-electronic]. 2012 Jul; 61(7): 965-6. PMID: 22406936, PMCID: PMC3919019, PII: gutjnl-2012-302142, DOI: 10.1136/gutjnl-2012-302142, ISSN: 1468-3288.

Sicinschi LA, Correa P, Bravo LE, Peek RM, Wilson KT, Loh JT, Yepez MC, Gold BD, Thompson DT, Cover TL, Schneider BG. Non-invasive genotyping of Helicobacter pylori cagA, vacA, and hopQ from asymptomatic children. Helicobacter. 2012 Apr; 17(2): 96-106. PMID: 22404439, PMCID: PMC3305281, DOI: 10.1111/j.1523-5378.2011.00919.x, ISSN: 1523-5378.

Duncan SS, Valk PL, Shaffer CL, Bordenstein SR, Cover TL. J-Western forms of Helicobacter pylori cagA constitute a distinct phylogenetic group with a widespread geographic distribution. J. Bacteriol [print-electronic]. 2012 Mar; 194(6): 1593-604. PMID: 22247512, PMCID: PMC3294863, PII: JB.06340-11, DOI: 10.1128/JB.06340-11, ISSN: 1098-5530.

Cover TL. Perspectives on methodology for in vitro culture of Helicobacter pylori. Methods Mol. Biol. 2012; 921: 11-5. PMID: 23015486, PMCID: PMC3921885, DOI: 10.1007/978-1-62703-005-2_3, ISSN: 1940-6029.

Johnson EM, Gaddy JA, Cover TL. Alterations in Helicobacter pylori triggered by contact with gastric epithelial cells. Front Cell Infect Microbiol. 2012; 2: 17. PMID: 22919609, PMCID: PMC3417513, DOI: 10.3389/fcimb.2012.00017, ISSN: 2235-2988.

Conradi J, Tegtmeyer N, Wozna M, Wissbrock M, Michalek C, Gagell C, Cover TL, Frank R, Sewald N, Backert S. An RGD helper sequence in CagL of Helicobacter pylori assists in interactions with integrins and injection of CagA. Front Cell Infect Microbiol. 2012; 2: 70. PMID: 22919661, PMCID: PMC3417467, DOI: 10.3389/fcimb.2012.00070, ISSN: 2235-2988.

Loh JT, Shaffer CL, Piazuelo MB, Bravo LE, McClain MS, Correa P, Cover TL. Analysis of cagA in Helicobacter pylori strains from Colombian populations with contrasting gastric cancer risk reveals a biomarker for disease severity. Cancer Epidemiol. Biomarkers Prev [print-electronic]. 2011 Oct; 20(10): 2237-49. PMID: 21859954, PMCID: PMC3189306, PII: 1055-9965.EPI-11-0548, DOI: 10.1158/1055-9965.EPI-11-0548, ISSN: 1538-7755.

Shaffer CL, Gaddy JA, Loh JT, Johnson EM, Hill S, Hennig EE, McClain MS, McDonald WH, Cover TL. Helicobacter pylori exploits a unique repertoire of type IV secretion system components for pilus assembly at the bacteria-host cell interface. PLoS Pathog [print-electronic]. 2011 Sep; 7(9): e1002237. PMID: 21909278, PMCID: PMC3164655, PII: PPATHOGENS-D-11-00533, DOI: 10.1371/journal.ppat.1002237, ISSN: 1553-7374.

de Sablet T, Piazuelo MB, Shaffer CL, Schneider BG, Asim M, Chaturvedi R, Bravo LE, Sicinschi LA, Delgado AG, Mera RM, Israel DA, Romero-Gallo J, Peek RM, Cover TL, Correa P, Wilson KT. Phylogeographic origin of Helicobacter pylori is a determinant of gastric cancer risk. Gut [print-electronic]. 2011 Sep; 60(9): 1189-95. PMID: 21357593, PMCID: PMC3133872, PII: gut.2010.234468, DOI: 10.1136/gut.2010.234468, ISSN: 1468-3288.

Nagy TA, Wroblewski LE, Wang D, Piazuelo MB, Delgado A, Romero-Gallo J, Noto J, Israel DA, Ogden SR, Correa P, Cover TL, Peek RM. ß-Catenin and p120 mediate PPARd-dependent proliferation induced by Helicobacter pylori in human and rodent epithelia. Gastroenterology [print-electronic]. 2011 Aug; 141(2): 553-64. PMID: 21704622, PMCID: PMC3152603, PII: S0016-5085(11)00621-4, DOI: 10.1053/j.gastro.2011.05.004, ISSN: 1528-0012.

Radin JN, González-Rivera C, Ivie SE, McClain MS, Cover TL. Helicobacter pylori VacA induces programmed necrosis in gastric epithelial cells. Infect. Immun [print-electronic]. 2011 Jul; 79(7): 2535-43. PMID: 21482684, PMCID: PMC3191986, PII: IAI.01370-10, DOI: 10.1128/IAI.01370-10, ISSN: 1098-5522.

Yan F, Cao H, Cover TL, Washington MK, Shi Y, Liu L, Chaturvedi R, Peek RM, Wilson KT, Polk DB. Colon-specific delivery of a probiotic-derived soluble protein ameliorates intestinal inflammation in mice through an EGFR-dependent mechanism. J. Clin. Invest [print-electronic]. 2011 Jun; 121(6): 2242-53. PMID: 21606592, PMCID: PMC3104743, PII: 44031, DOI: 10.1172/JCI44031, ISSN: 1558-8238.

Gangwer KA, Shaffer CL, Suerbaum S, Lacy DB, Cover TL, Bordenstein SR. Molecular evolution of the Helicobacter pylori vacuolating toxin gene vacA. J. Bacteriol [print-electronic]. 2010 Dec; 192(23): 6126-35. PMID: 20870762, PMCID: PMC2981223, PII: JB.01081-10, DOI: 10.1128/JB.01081-10, ISSN: 1098-5530.

González-Rivera C, Gangwer KA, McClain MS, Eli IM, Chambers MG, Ohi MD, Lacy DB, Cover TL. Reconstitution of Helicobacter pylori VacA toxin from purified components. Biochemistry. 2010 Jul 7/13/2010; 49(27): 5743-52. PMID: 20527875, PMCID: PMC2910095, DOI: 10.1021/bi100618g, ISSN: 1520-4995.

Loh JT, Gupta SS, Friedman DB, Krezel AM, Cover TL. Analysis of protein expression regulated by the Helicobacter pylori ArsRS two-component signal transduction system. J. Bacteriol [print-electronic]. 2010 Apr; 192(8): 2034-43. PMID: 20154125, PMCID: PMC2849440, PII: JB.01703-08, DOI: 10.1128/JB.01703-08, ISSN: 1098-5530.

Ivie SE, McClain MS, Algood HM, Lacy DB, Cover TL. Analysis of a beta-helical region in the p55 domain of Helicobacter pylori vacuolating toxin. BMC Microbiol. 2010; 10: 60. PMID: 20178613, PMCID: PMC2836311, PII: 1471-2180-10-60, DOI: 10.1186/1471-2180-10-60, ISSN: 1471-2180.

Shen F, Hobley L, Doherty N, Loh JT, Cover TL, Sockett RE, Hardie KR, Atherton JC. In Helicobacter pylori auto-inducer-2, but not LuxS/MccAB catalysed reverse transsulphuration, regulates motility through modulation of flagellar gene transcription. BMC Microbiol. 2010; 10: 210. PMID: 20691071, PMCID: PMC2923134, PII: 1471-2180-10-210, DOI: 10.1186/1471-2180-10-210, ISSN: 1471-2180.

Algood HM, Allen SS, Washington MK, Peek RM, Miller GG, Cover TL. Regulation of gastric B cell recruitment is dependent on IL-17 receptor A signaling in a model of chronic bacterial infection. J. Immunol [print-electronic]. 2009 Nov 11/1/2009; 183(9): 5837-46. PMID: 19812196, PMCID: PMC2834183, PII: jimmunol.0901206, DOI: 10.4049/jimmunol.0901206, ISSN: 1550-6606.

Cover TL, Blaser MJ. Helicobacter pylori in health and disease. Gastroenterology [print-electronic]. 2009 May; 136(6): 1863-73. PMID: 19457415, PMCID: PMC3644425, PII: S0016-5085(09)00339-4, DOI: 10.1053/j.gastro.2009.01.073, ISSN: 1528-0012.

Yan F, Cao H, Chaturvedi R, Krishna U, Hobbs SS, Dempsey PJ, Peek RM, Cover TL, Washington MK, Wilson KT, Polk DB. Epidermal growth factor receptor activation protects gastric epithelial cells from Helicobacter pylori-induced apoptosis. Gastroenterology [print-electronic]. 2009 Apr; 136(4): 1297-1307, e1. PMID: 19250983, PMCID: PMC2878739, PII: S0016-5085(08)02317-2, DOI: 10.1053/j.gastro.2008.12.059, ISSN: 1528-0012.

Gupta SS, Borin BN, Cover TL, Krezel AM. Structural analysis of the DNA-binding domain of the Helicobacter pylori response regulator ArsR. J. Biol. Chem [print-electronic]. 2009 Mar 3/6/2009; 284(10): 6536-45. PMID: 19117956, PMCID: PMC2649097, PII: M804592200, DOI: 10.1074/jbc.M804592200, ISSN: 0021-9258.

McClain MS, Shaffer CL, Israel DA, Peek RM, Cover TL. Genome sequence analysis of Helicobacter pylori strains associated with gastric ulceration and gastric cancer. BMC Genomics. 2009; 10: 3. PMID: 19123947, PMCID: PMC2627912, PII: 1471-2164-10-3, DOI: 10.1186/1471-2164-10-3, ISSN: 1471-2164.

Loh JT, Torres VJ, Algood HM, McClain MS, Cover TL. Helicobacter pylori HopQ outer membrane protein attenuates bacterial adherence to gastric epithelial cells. FEMS Microbiol. Lett. 2008 Dec; 289(1): 53-8. PMID: 19065710, PMCID: PMC2651568, ISSN: 0378-1097.

Wang F, Xia P, Wu F, Wang D, Wang W, Ward T, Liu Y, Aikhionbare F, Guo Z, Powell M, Liu B, Bi F, Shaw A, Zhu Z, Elmoselhi A, Fan D, Cover TL, Ding X, Yao X. Helicobacter pylori VacA disrupts apical membrane-cytoskeletal interactions in gastric parietal cells. J. Biol. Chem [print-electronic]. 2008 Sep 9/26/2008; 283(39): 26714-25. PMID: 18625712, PMCID: PMC2546541, PII: M800527200, DOI: 10.1074/jbc.M800527200, ISSN: 0021-9258.

Ding SZ, Olekhnovich IN, Cover TL, Peek RM, Smith MF, Goldberg JB. Helicobacter pylori and mitogen-activated protein kinases mediate activator protein-1 (AP-1) subcomponent protein expression and DNA-binding activity in gastric epithelial cells. FEMS Immunol. Med. Microbiol [print-electronic]. 2008 Aug; 53(3): 385-94. PMID: 18625013, PMCID: PMC2553690, PII: FIM439, DOI: 10.1111/j.1574-695X.2008.00439.x, ISSN: 0928-8244.

Ivie SE, McClain MS, Torres VJ, Algood HM, Lacy DB, Yang R, Blanke SR, Cover TL. Helicobacter pylori VacA subdomain required for intracellular toxin activity and assembly of functional oligomeric complexes. Infect. Immun [print-electronic]. 2008 Jul; 76(7): 2843-51. PMID: 18443094, PMCID: PMC2446698, PII: IAI.01664-07, DOI: 10.1128/IAI.01664-07, ISSN: 1098-5522.

Cover TL. Beta2 integrin mediates entry of a bacterial toxin into T lymphocytes. Cell Host Microbe. 2008 Jan 1/17/2008; 3(1): 5-6. PMID: 18191787, PII: S1931-3128(07)00312-5, DOI: 10.1016/j.chom.2007.12.004, ISSN: 1934-6069.

Algood HM, Gallo-Romero J, Wilson KT, Peek RM, Cover TL. Host response to Helicobacter pylori infection before initiation of the adaptive immune response. FEMS Immunol. Med. Microbiol [print-electronic]. 2007 Dec; 51(3): 577-86. PMID: 17919297, PII: FIM338, DOI: 10.1111/j.1574-695X.2007.00338.x, ISSN: 0928-8244.

Torres VJ, VanCompernolle SE, Sundrud MS, Unutmaz D, Cover TL. Helicobacter pylori vacuolating cytotoxin inhibits activation-induced proliferation of human T and B lymphocyte subsets. J. Immunol. 2007 Oct 10/15/2007; 179(8): 5433-40. PMID: 17911630, PII: 179/8/5433, ISSN: 0022-1767.

Gangwer KA, Mushrush DJ, Stauff DL, Spiller B, McClain MS, Cover TL, Lacy DB. Crystal structure of the Helicobacter pylori vacuolating toxin p55 domain. Proc. Natl. Acad. Sci. U.S.A [print-electronic]. 2007 Oct 10/9/2007; 104(41): 16293-8. PMID: 17911250, PMCID: PMC2042200, PII: 0707447104, DOI: 10.1073/pnas.0707447104, ISSN: 0027-8424.

Chaturvedi R, Asim M, Lewis ND, Algood HM, Cover TL, Kim PY, Wilson KT. L-arginine availability regulates inducible nitric oxide synthase-dependent host defense against Helicobacter pylori. Infect. Immun [print-electronic]. 2007 Sep; 75(9): 4305-15. PMID: 17562760, PMCID: PMC1951193, PII: IAI.00578-07, DOI: 10.1128/IAI.00578-07, ISSN: 0019-9567.

Loh JT, Torres VJ, Cover TL. Regulation of Helicobacter pylori cagA expression in response to salt. Cancer Res. 2007 May 5/15/2007; 67(10): 4709-15. PMID: 17510398, PII: 67/10/4709, DOI: 10.1158/0008-5472.CAN-06-4746, ISSN: 0008-5472.

Ghose C, Perez-Perez GI, Torres VJ, Crosatti M, Nomura A, Peek RM, Cover TL, Francois F, Blaser MJ. Serological assays for identification of human gastric colonization by Helicobacter pylori strains expressing VacA m1 or m2. Clin. Vaccine Immunol [print-electronic]. 2007 Apr; 14(4): 442-50. PMID: 17267587, PMCID: PMC1865612, PII: CVI.00434-06, DOI: 10.1128/CVI.00434-06, ISSN: 1556-6811.

McClain MS, Cover TL. Functional analysis of neutralizing antibodies against Clostridium perfringens epsilon-toxin. Infect. Immun [print-electronic]. 2007 Apr; 75(4): 1785-93. PMID: 17261609, PMCID: PMC1865726, PII: IAI.01643-06, DOI: 10.1128/IAI.01643-06, ISSN: 0019-9567.

Yan F, Cao H, Cover TL, Whitehead R, Washington MK, Polk DB. Soluble proteins produced by probiotic bacteria regulate intestinal epithelial cell survival and growth. Gastroenterology [print-electronic]. 2007 Feb; 132(2): 562-75. PMID: 17258729, PMCID: PMC3036990, PII: S0016-5085(06)02477-2, DOI: 10.1053/j.gastro.2006.11.022, ISSN: 0016-5085.

Algood HM, Torres VJ, Unutmaz D, Cover TL. Resistance of primary murine CD4+ T cells to Helicobacter pylori vacuolating cytotoxin. Infect. Immun [print-electronic]. 2007 Jan; 75(1): 334-41. PMID: 17074854, PMCID: PMC1828377, PII: IAI.01063-06, DOI: 10.1128/IAI.01063-06, ISSN: 0019-9567.

Oswald-Richter K, Torres VJ, Sundrud MS, VanCompernolle SE, Cover TL, Unutmaz D. Helicobacter pylori VacA toxin inhibits human immunodeficiency virus infection of primary human T cells. J. Virol [print-electronic]. 2006 Dec; 80(23): 11767-75. PMID: 17005643, PMCID: PMC1642621, PII: JVI.00213-06, DOI: 10.1128/JVI.00213-06, ISSN: 0022-538X.

Cover TL. Role of Helicobacter pylori outer membrane proteins in gastroduodenal disease [editorial]. J. Infect. Dis [print-electronic]. 2006 Nov 11/15/2006; 194(10): 1343-5. PMID: 17054062, PII: JID37089, DOI: 10.1086/508432, ISSN: 0022-1899.

McClain MS, Czajkowsky DM, Torres VJ, Szabo G, Shao Z, Cover TL. Random mutagenesis of Helicobacter pylori vacA to identify amino acids essential for vacuolating cytotoxic activity. Infect. Immun [print-electronic]. 2006 Nov; 74(11): 6188-95. PMID: 16954403, PMCID: PMC1695532, PII: IAI.00915-06, DOI: 10.1128/IAI.00915-06, ISSN: 0019-9567.

Algood HM, Cover TL. Helicobacter pylori persistence: an overview of interactions between H. pylori and host immune defenses. Clin. Microbiol. Rev. 2006 Oct; 19(4): 597-613. PMID: 17041136, PMCID: PMC1592695, PII: 19/4/597, DOI: 10.1128/CMR.00006-06, ISSN: 0893-8512.

Lee WK, Ogura K, Loh JT, Cover TL, Berg DE. Quantitative effect of luxS gene inactivation on the fitness of Helicobacter pylori. Appl. Environ. Microbiol [print-electronic]. 2006 Oct; 72(10): 6615-22. PMID: 16936059, PMCID: PMC1610275, PII: AEM.01291-06, DOI: 10.1128/AEM.01291-06, ISSN: 0099-2240.

Busler VJ, Torres VJ, McClain MS, Tirado O, Friedman DB, Cover TL. Protein-protein interactions among Helicobacter pylori cag proteins. J. Bacteriol. 2006 Jul; 188(13): 4787-800. PMID: 16788188, PMCID: PMC1482994, PII: 188/13/4787, DOI: 10.1128/JB.00066-06, ISSN: 0021-9193.

Loh JT, Cover TL. Requirement of histidine kinases HP0165 and HP1364 for acid resistance in Helicobacter pylori. Infect. Immun. 2006 May; 74(5): 3052-9. PMID: 16622250, PMCID: PMC1459715, PII: 74/5/3052, DOI: 10.1128/IAI.74.5.3052-3059.2006, ISSN: 0019-9567.

Hennig EE, Allen JM, Cover TL. Multiple chromosomal loci for the babA gene in Helicobacter pylori. Infect. Immun. 2006 May; 74(5): 3046-51. PMID: 16622249, PMCID: PMC1459736, PII: 74/5/3046, DOI: 10.1128/IAI.74.5.3046-3051.2006, ISSN: 0019-9567.

Torres VJ, McClain MS, Cover TL. Mapping of a domain required for protein-protein interactions and inhibitory activity of a Helicobacter pylori dominant-negative VacA mutant protein. Infect. Immun. 2006 Apr; 74(4): 2093-101. PMID: 16552038, PMCID: PMC1418911, PII: 74/4/2093, DOI: 10.1128/IAI.74.4.2093-2101.2006, ISSN: 0019-9567.

Johansson M, Lehto M, Tanhuanpää K, Cover TL, Olkkonen VM. The oxysterol-binding protein homologue ORP1L interacts with Rab7 and alters functional properties of late endocytic compartments. Mol. Biol. Cell [print-electronic]. 2005 Dec; 16(12): 5480-92. PMID: 16176980, PMCID: PMC1289395, PII: E05-03-0189, DOI: 10.1091/mbc.E05-03-0189, ISSN: 1059-1524.

Czajkowsky DM, Iwamoto H, Szabo G, Cover TL, Shao Z. Mimicry of a host anion channel by a Helicobacter pylori pore-forming toxin. Biophys. J [print-electronic]. 2005 Nov; 89(5): 3093-101. PMID: 16100263, PMCID: PMC1366806, PII: S0006-3495(05)72952-0, DOI: 10.1529/biophysj.105.066746, ISSN: 0006-3495.

Cao P, Lee KJ, Blaser MJ, Cover TL. Analysis of hopQ alleles in East Asian and Western strains of Helicobacter pylori. FEMS Microbiol. Lett. 2005 Oct 10/1/2005; 251(1): 37-43. PMID: 16102915, PII: S0378-1097(05)00484-2, DOI: 10.1016/j.femsle.2005.07.023, ISSN: 0378-1097.

El-Bez C, Adrian M, Dubochet J, Cover TL. High resolution structural analysis of Helicobacter pylori VacA toxin oligomers by cryo-negative staining electron microscopy. J. Struct. Biol. 2005 Sep; 151(3): 215-28. PMID: 16125415, PII: S1047-8477(05)00145-0, DOI: 10.1016/j.jsb.2005.07.001, ISSN: 1047-8477.

Torres VJ, Ivie SE, McClain MS, Cover TL. Functional properties of the p33 and p55 domains of the Helicobacter pylori vacuolating cytotoxin. J. Biol. Chem [print-electronic]. 2005 Jun 6/3/2005; 280(22): 21107-14. PMID: 15817461, PII: M501042200, DOI: 10.1074/jbc.M501042200, ISSN: 0021-9258.

Terry CE, McGinnis LM, Madigan KC, Cao P, Cover TL, Liechti GW, Peek RM, Forsyth MH. Genomic Comparison of cag pathogenicity island (PAI)-positive and -negative Helicobacter pylori strains: identification of novel markers for cag PAI-positive strains. Infect. Immun. 2005 Jun; 73(6): 3794-8. PMID: 15908415, PMCID: PMC1111835, PII: 73/6/3794, DOI: 10.1128/IAI.73.6.3794-3798.2005, ISSN: 0019-9567.

Cover TL, Blanke SR. Helicobacter pylori VacA, a paradigm for toxin multifunctionality. Nat. Rev. Microbiol. 2005 Apr; 3(4): 320-32. PMID: 15759043, PII: nrmicro1095, DOI: 10.1038/nrmicro1095, ISSN: 1740-1526.

Loh JT, Forsyth MH, Cover TL. Growth phase regulation of flaA expression in Helicobacter pylori is luxS dependent. Infect. Immun. 2004 Sep; 72(9): 5506-10. PMID: 15322055, PMCID: PMC517482, PII: 72/9/5506, DOI: 10.1128/IAI.72.9.5506-5510.2004, ISSN: 0019-9567.

Geisse NA, Cover TL, Henderson RM, Edwardson JM. Targeting of Helicobacter pylori vacuolating toxin to lipid raft membrane domains analysed by atomic force microscopy. Biochem. J. 2004 Aug 8/1/2004; 381(Pt 3): 911-7. PMID: 15128269, PMCID: PMC1133903, PII: BJ20031719, DOI: 10.1042/BJ20031719, ISSN: 1470-8728.

Hennig EE, Mernaugh R, Edl J, Cao P, Cover TL. Heterogeneity among Helicobacter pylori strains in expression of the outer membrane protein BabA. Infect. Immun. 2004 Jun; 72(6): 3429-35. PMID: 15155649, PMCID: PMC415674, PII: 72/6/3429, DOI: 10.1128/IAI.72.6.3429-3435.2004, ISSN: 0019-9567.

Sundrud MS, Torres VJ, Unutmaz D, Cover TL. 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. U.S.A [print-electronic]. 2004 May 5/18/2004; 101(20): 7727-32. PMID: 15128946, PMCID: PMC419674, PII: 0401528101, DOI: 10.1073/pnas.0401528101, ISSN: 0027-8424.

Li Y, Wandinger-Ness A, Goldenring JR, Cover TL. Clustering and redistribution of late endocytic compartments in response to Helicobacter pylori vacuolating toxin. Mol. Biol. Cell [print-electronic]. 2004 Apr; 15(4): 1946-59. PMID: 14742715, PMCID: PMC379289, PII: E03-08-0618, DOI: 10.1091/mbc.E03-08-0618, ISSN: 1059-1524.

Torres VJ, McClain MS, Cover TL. Interactions between p-33 and p-55 domains of the Helicobacter pylori vacuolating cytotoxin (VacA). J. Biol. Chem [print-electronic]. 2004 Jan 1/16/2004; 279(3): 2324-31. PMID: 14593124, PII: M310159200, DOI: 10.1074/jbc.M310159200, ISSN: 0021-9258.

Fonteneau JF, Kavanagh DG, Lirvall M, Sanders C, Cover TL, Bhardwaj N, Larsson M. Characterization of the MHC class I cross-presentation pathway for cell-associated antigens by human dendritic cells. Blood [print-electronic]. 2003 Dec 12/15/2003; 102(13): 4448-55. PMID: 12933572, PII: 2003-06-1801, DOI: 10.1182/blood-2003-06-1801, ISSN: 0006-4971.

Willhite DC, Cover TL, Blanke SR. Cellular vacuolation and mitochondrial cytochrome c release are independent outcomes of Helicobacter pylori vacuolating cytotoxin activity that are each dependent on membrane channel formation. J. Biol. Chem [print-electronic]. 2003 Nov 11/28/2003; 278(48): 48204-9. PMID: 13129933, PII: M304131200, DOI: 10.1074/jbc.M304131200, ISSN: 0021-9258.

Marlink KL, Bacon KD, Sheppard BC, Ashktorab H, Smoot DT, Cover TL, Deveney CW, Rutten MJ. Effects of Helicobacter pylori on intracellular Ca2+ signaling in normal human gastric mucous epithelial cells. Am. J. Physiol. Gastrointest. Liver Physiol [print-electronic]. 2003 Jul; 285(1): G163-76. PMID: 12606301, PII: 00257.2002, DOI: 10.1152/ajpgi.00257.2002, ISSN: 0193-1857.

McGowan CC, Necheva AS, Forsyth MH, Cover TL, Blaser MJ. Promoter analysis of Helicobacter pylori genes with enhanced expression at low pH. Mol. Microbiol. 2003 Jun; 48(5): 1225-39. PMID: 12787351, PII: 3500, ISSN: 0950-382X.

McClain MS, Iwamoto H, Cao P, Vinion-Dubiel AD, Li Y, Szabo G, Shao Z, Cover TL. Essential role of a GXXXG motif for membrane channel formation by Helicobacter pylori vacuolating toxin. J. Biol. Chem [print-electronic]. 2003 Apr 4/4/2003; 278(14): 12101-8. PMID: 12562777, PII: M212595200, DOI: 10.1074/jbc.M212595200, ISSN: 0021-9258.

McClain MS, Cover TL. Expression of Helicobacter pylori vacuolating toxin in Escherichia coli. Infect. Immun. 2003 Apr; 71(4): 2266-71. PMID: 12654854, PMCID: PMC152094, ISSN: 0019-9567.

Cover TL, Krishna US, Israel DA, Peek RM. Induction of gastric epithelial cell apoptosis by Helicobacter pylori vacuolating cytotoxin. Cancer Res. 2003 Mar 3/1/2003; 63(5): 951-7. PMID: 12615708, ISSN: 0008-5472.

Nabekura T, Morishima S, Cover TL, Mori S, Kannan H, Komune S, Okada Y. Recovery from lactacidosis-induced glial cell swelling with the aid of exogenous anion channels. Glia. 2003 Feb; 41(3): 247-59. PMID: 12528180, DOI: 10.1002/glia.10190, ISSN: 0894-1491.

Cao P, Cover TL. Two different families of hopQ alleles in Helicobacter pylori. J. Clin. Microbiol. 2002 Dec; 40(12): 4504-11. PMID: 12454143, PMCID: PMC154617, ISSN: 0095-1137.

Ikonomov OC, Sbrissa D, Yoshimori T, Cover TL, Shisheva A. PIKfyve Kinase and SKD1 AAA ATPase define distinct endocytic compartments. Only PIKfyve expression inhibits the cell-vacoulating activity of Helicobacter pylori VacA toxin. J. Biol. Chem [print-electronic]. 2002 Nov 11/29/2002; 277(48): 46785-90. PMID: 12213828, PII: M208068200, DOI: 10.1074/jbc.M208068200, ISSN: 0021-9258.

Drake WP, Pei Z, Pride DT, Collins RD, Cover TL, Blaser MJ. Molecular analysis of sarcoidosis tissues for mycobacterium species DNA. Emerging Infect. Dis. 2002 Nov; 8(11): 1334-41. PMID: 12453366, PMCID: PMC2738555, DOI: 10.3201/eid0811.020318, ISSN: 1080-6040.

Graham JE, Peek RM, Krishna U, Cover TL. Global analysis of Helicobacter pylori gene expression in human gastric mucosa. Gastroenterology. 2002 Nov; 123(5): 1637-48. PMID: 12404238, PMCID: PMC1361305, PII: S0016508502002974, ISSN: 0016-5085.

Schraw W, Li Y, McClain MS, van der Goot FG, Cover TL. Association of Helicobacter pylori vacuolating toxin (VacA) with lipid rafts. J. Biol. Chem [print-electronic]. 2002 Sep 9/13/2002; 277(37): 34642-50. PMID: 12121984, PII: M203466200, DOI: 10.1074/jbc.M203466200, ISSN: 0021-9258.

Forsyth MH, Cao P, Garcia PP, Hall JD, Cover TL. Genome-wide transcriptional profiling in a histidine kinase mutant of Helicobacter pylori identifies members of a regulon. J. Bacteriol. 2002 Aug; 184(16): 4630-5. PMID: 12142435, PMCID: PMC135264, ISSN: 0021-9193.

Adrian M, Cover TL, Dubochet J, Heuser JE. Multiple oligomeric states of the Helicobacter pylori vacuolating toxin demonstrated by cryo-electron microscopy. J. Mol. Biol. 2002 Apr 4/19/2002; 318(1): 121-33. PMID: 12054773, PII: S0022-2836(02)00047-5, DOI: 10.1016/S0022-2836(02)00047-5, ISSN: 0022-2836.

McClain MS, Cao P, Iwamoto H, Vinion-Dubiel AD, Szabo G, Shao Z, Cover TL. A 12-amino-acid segment, present in type s2 but not type s1 Helicobacter pylori VacA proteins, abolishes cytotoxin activity and alters membrane channel formation. J. Bacteriol. 2001 Nov; 183(22): 6499-508. PMID: 11673417, PMCID: PMC95478, DOI: 10.1128/JB.183.22.6499-6508.2001, ISSN: 0021-9193.

Papini E, Zoratti M, Cover TL. In search of the Helicobacter pylori VacA mechanism of action. Toxicon. 2001 Nov; 39(11): 1757-67. PMID: 11595638, PII: S0041010101001623, ISSN: 0041-0101.

Vinion-Dubiel AD, McClain MS, Cao P, Mernaugh RL, Cover TL. Antigenic diversity among Helicobacter pylori vacuolating toxins. Infect. Immun. 2001 Jul; 69(7): 4329-36. PMID: 11401970, PMCID: PMC98503, DOI: 10.1128/IAI.69.7.4329-4336.2001, ISSN: 0019-9567.

Tham KT, Peek RM, Atherton JC, Cover TL, Perez-Perez GI, Shyr Y, Blaser MJ. Helicobacter pylori genotypes, host factors, and gastric mucosal histopathology in peptic ulcer disease. Hum. Pathol. 2001 Mar; 32(3): 264-73. PMID: 11274634, PII: S0046-8177(01)25759-1, DOI: 10.1053/hupa.2001.21136, ISSN: 0046-8177.

McClain MS, Cao P, Cover TL. Amino-terminal hydrophobic region of Helicobacter pylori vacuolating cytotoxin (VacA) mediates transmembrane protein dimerization. Infect. Immun. 2001 Feb; 69(2): 1181-4. PMID: 11160018, PMCID: PMC98002, DOI: 10.1128/IAI.69.2.1181-1184.2001, ISSN: 0019-9567.

Nguyen VQ, Caprioli RM, Cover TL. Carboxy-terminal proteolytic processing of Helicobacter pylori vacuolating toxin. Infect. Immun. 2001 Jan; 69(1): 543-6. PMID: 11119549, PMCID: PMC97915, DOI: 10.1128/IAI.69.1.543-546.2001, ISSN: 0019-9567.

McClain MS, Schraw W, Ricci V, Boquet P, Cover TL. Acid activation of Helicobacter pylori vacuolating cytotoxin (VacA) results in toxin internalization by eukaryotic cells. Mol. Microbiol. 2000 Jul; 37(2): 433-42. PMID: 10931337, PII: mmi2013, ISSN: 0950-382X.

Forsyth MH, Cover TL. Intercellular communication in Helicobacter pylori: luxS is essential for the production of an extracellular signaling molecule. Infect. Immun. 2000 Jun; 68(6): 3193-9. PMID: 10816463, PMCID: PMC97560, ISSN: 0019-9567.

Hotchin NA, Cover TL, Akhtar N. Cell vacuolation induced by the VacA cytotoxin of Helicobacter pylori is regulated by the Rac1 GTPase. J. Biol. Chem. 2000 May 5/12/2000; 275(19): 14009-12. PMID: 10747859, PII: C000153200, ISSN: 0021-9258.

Vinion-Dubiel AD, McClain MS, Czajkowsky DM, Iwamoto H, Ye D, Cao P, Schraw W, Szabo G, Blanke SR, Shao Z, Cover TL. A dominant negative mutant of Helicobacter pylori vacuolating toxin (VacA) inhibits VacA-induced cell vacuolation. J. Biol. Chem. 1999 Dec 12/31/1999; 274(53): 37736-42. PMID: 10608833, ISSN: 0021-9258.

Peek RM, Blaser MJ, Mays DJ, Forsyth MH, Cover TL, Song SY, Krishna U, Pietenpol JA. Helicobacter pylori strain-specific genotypes and modulation of the gastric epithelial cell cycle. Cancer Res. 1999 Dec 12/15/1999; 59(24): 6124-31. PMID: 10626802, ISSN: 0008-5472.

Schraw W, McClain MS, Cover TL. Kinetics and mechanisms of extracellular protein release by Helicobacter pylori. Infect. Immun. 1999 Oct; 67(10): 5247-52. PMID: 10496902, PMCID: PMC96877, ISSN: 0019-9567.

Atherton JC, Sharp PM, Cover TL, Gonzalez-Valencia G, Peek RM, Thompson SA, Hawkey CJ, Blaser MJ. Vacuolating cytotoxin (vacA) alleles of Helicobacter pylori comprise two geographically widespread types, m1 and m2, and have evolved through limited recombination. Curr. Microbiol. 1999 Oct; 39(4): 211-8. PMID: 10486057, PMCID: PMC2762107, PII: 3022, ISSN: 0343-8651.

Atherton JC, Cover TL, Twells RJ, Morales MR, Hawkey CJ, Blaser MJ. Simple and accurate PCR-based system for typing vacuolating cytotoxin alleles of Helicobacter pylori. J. Clin. Microbiol. 1999 Sep; 37(9): 2979-82. PMID: 10449485, PMCID: PMC85427, ISSN: 0095-1137.

Pai R, Cover TL, Tarnawski AS. Helicobacter pylori vacuolating cytotoxin (VacA) disorganizes the cytoskeletal architecture of gastric epithelial cells. Biochem. Biophys. Res. Commun. 1999 Aug 8/19/1999; 262(1): 245-50. PMID: 10448099, PII: S0006-291X(99)91194-4, DOI: 10.1006/bbrc.1999.1194, ISSN: 0006-291X.

Perez-Perez GI, Peek RM, Atherton JC, Blaser MJ, Cover TL. Detection of anti-VacA antibody responses in serum and gastric juice samples using type s1/m1 and s2/m2 Helicobacter pylori VacA antigens. Clin. Diagn. Lab. Immunol. 1999 Jul; 6(4): 489-93. PMID: 10391848, PMCID: PMC95713, ISSN: 1071-412X.

Cover TL, Blaser MJ. Helicobacter pylori factors associated with disease [editorial]. Gastroenterology. 1999 Jul; 117(1): 257-61. PMID: 10381935, PII: S0016508599000797, ISSN: 0016-5085.

Fiocca R, Necchi V, Sommi P, Ricci V, Telford J, Cover TL, Solcia E. Release of Helicobacter pylori vacuolating cytotoxin by both a specific secretion pathway and budding of outer membrane vesicles. Uptake of released toxin and vesicles by gastric epithelium. J. Pathol. 1999 Jun; 188(2): 220-6. PMID: 10398168, PII: 10.1002/(SICI)1096-9896(199906)188:2<220::AID-PATH307>3.0.CO;2-C, DOI: 10.1002/(SICI)1096-9896(199906)188:2<220::AID-PATH307>3.0.CO;2-C, ISSN: 0022-3417.

Iwamoto H, Czajkowsky DM, Cover TL, Szabo G, Shao Z. VacA from Helicobacter pylori: a hexameric chloride channel. FEBS Lett. 1999 Apr 4/30/1999; 450(1-2): 101-4. PMID: 10350065, PII: S0014-5793(99)00474-3, ISSN: 0014-5793.

Forsyth MH, Cover TL. Mutational analysis of the vacA promoter provides insight into gene transcription in Helicobacter pylori. J. Bacteriol. 1999 Apr; 181(7): 2261-6. PMID: 10094707, PMCID: PMC93642, ISSN: 0021-9193.

Czajkowsky DM, Iwamoto H, Cover TL, Shao Z. The vacuolating toxin from Helicobacter pylori forms hexameric pores in lipid bilayers at low pH. Proc. Natl. Acad. Sci. U.S.A. 1999 Mar 3/2/1999; 96(5): 2001-6. PMID: 10051584, PMCID: PMC26726, ISSN: 0027-8424.

McGowan CC, Necheva A, Thompson SA, Cover TL, Blaser MJ. Acid-induced expression of an LPS-associated gene in Helicobacter pylori. Mol. Microbiol. 1998 Oct; 30(1): 19-31. PMID: 9786182, ISSN: 0950-382X.

Pagliaccia C, de Bernard M, Lupetti P, Ji X, Burroni D, Cover TL, Papini E, Rappuoli R, Telford JL, Reyrat JM. The m2 form of the Helicobacter pylori cytotoxin has cell type-specific vacuolating activity. Proc. Natl. Acad. Sci. U.S.A. 1998 Aug 8/18/1998; 95(17): 10212-7. PMID: 9707626, PMCID: PMC21487, ISSN: 0027-8424.

Forsyth MH, Atherton JC, Blaser MJ, Cover TL. Heterogeneity in levels of vacuolating cytotoxin gene (vacA) transcription among Helicobacter pylori strains. Infect. Immun. 1998 Jul; 66(7): 3088-94. PMID: 9632570, PMCID: PMC108317, ISSN: 0019-9567.

Pai R, Wyle FA, Cover TL, Itani RM, Domek MJ, Tarnawski AS. Helicobacter pylori culture supernatant interferes with epidermal growth factor-activated signal transduction in human gastric KATO III cells. Am. J. Pathol. 1998 Jun; 152(6): 1617-24. PMID: 9626065, PMCID: PMC1858446, ISSN: 0002-9440.

Cao P, McClain MS, Forsyth MH, Cover TL. Extracellular release of antigenic proteins by Helicobacter pylori. Infect. Immun. 1998 Jun; 66(6): 2984-6. PMID: 9596777, PMCID: PMC108299, ISSN: 0019-9567.

Romano M, Ricci V, Di Popolo A, Sommi P, Del Vecchio Blanco C, Bruni CB, Ventura U, Cover TL, Blaser MJ, Coffey RJ, Zarrilli R. Helicobacter pylori upregulates expression of epidermal growth factor-related peptides, but inhibits their proliferative effect in MKN 28 gastric mucosal cells. J. Clin. Invest. 1998 Apr 4/15/1998; 101(8): 1604-13. PMID: 9541490, PMCID: PMC508741, DOI: 10.1172/JCI1174, ISSN: 0021-9738.

Cover TL. An intracellular target for Helicobacter pylori vacuolating toxin. Trends Microbiol. 1998 Apr; 6(4): 127-8; discussion 128. PMID: 9587185, PII: S0966-842X(98)01231-1, ISSN: 0966-842X.

Cover TL. Commentary: Helicobacter pylori transmission, host factors, and bacterial factors. Gastroenterology. 1997 Dec; 113(6 Suppl): S29-30. PMID: 9394756, ISSN: 0016-5085.

Cover TL, Hanson PI, Heuser JE. Acid-induced dissociation of VacA, the Helicobacter pylori vacuolating cytotoxin, reveals its pattern of assembly. J. Cell Biol. 1997 Aug 8/25/1997; 138(4): 759-69. PMID: 9265644, PMCID: PMC2138037, ISSN: 0021-9525.

Eaton KA, Cover TL, Tummuru MK, Blaser MJ, Krakowka S. Role of vacuolating cytotoxin in gastritis due to Helicobacter pylori in gnotobiotic piglets. Infect. Immun. 1997 Aug; 65(8): 3462-4. PMID: 9234813, PMCID: PMC175490, ISSN: 0019-9567.

McGowan CC, Cover TL, Blaser MJ. Analysis of F1F0-ATPase from Helicobacter pylori. Infect. Immun. 1997 Jul; 65(7): 2640-7. PMID: 9199431, PMCID: PMC175373, ISSN: 0019-9567.

Cover TL. Genetic diversity in Helicobacter pylori. Helicobacter. 1997 Jun; 2(2): 108-9. PMID: 9432328, ISSN: 1083-4389.

Cao P, Cover TL. High-level genetic diversity in the vapD chromosomal region of Helicobacter pylori. J. Bacteriol. 1997 May; 179(9): 2852-6. PMID: 9139899, PMCID: PMC179045, ISSN: 0021-9193.

Atherton JC, Peek RM, Tham KT, Cover TL, Blaser MJ. Clinical and pathological importance of heterogeneity in vacA, the vacuolating cytotoxin gene of Helicobacter pylori. Gastroenterology. 1997 Jan; 112(1): 92-9. PMID: 8978347, PII: S0016508597000243, ISSN: 0016-5085.

Smoot DT, Resau JH, Earlington MH, Simpson M, Cover TL. Effects of Helicobacter pylori vacuolating cytotoxin on primary cultures of human gastric epithelial cells. Gut. 1996 Dec; 39(6): 795-9. PMID: 9038659, PMCID: PMC1383449, ISSN: 0017-5749.

Harris PR, Cover TL, Crowe DR, Orenstein JM, Graham MF, Blaser MJ, Smith PD. Helicobacter pylori cytotoxin induces vacuolation of primary human mucosal epithelial cells. Infect. Immun. 1996 Nov; 64(11): 4867-71. PMID: 8890255, PMCID: PMC174461, ISSN: 0019-9567.

Garner JA, Cover TL. Binding and internalization of the Helicobacter pylori vacuolating cytotoxin by epithelial cells. Infect. Immun. 1996 Oct; 64(10): 4197-203. PMID: 8926088, PMCID: PMC174356, ISSN: 0019-9567.

Atherton JC, Tham KT, Peek RM, Cover TL, Blaser MJ. Density of Helicobacter pylori infection in vivo as assessed by quantitative culture and histology. J. Infect. Dis. 1996 Sep; 174(3): 552-6. PMID: 8769613, ISSN: 0022-1899.

Ricci V, Ciacci C, Zarrilli R, Sommi P, Tummuru MK, Del Vecchio Blanco C, Bruni CB, Cover TL, Blaser MJ, Romano M. Effect of Helicobacter pylori on gastric epithelial cell migration and proliferation in vitro: role of VacA and CagA. Infect. Immun. 1996 Jul; 64(7): 2829-33. PMID: 8698518, PMCID: PMC174149, ISSN: 0019-9567.

Cover TL. The vacuolating cytotoxin of Helicobacter pylori. Mol. Microbiol. 1996 Apr; 20(2): 241-6. PMID: 8733223, ISSN: 0950-382X.

McGowan CC, Cover TL, Blaser MJ. Helicobacter pylori and gastric acid: biological and therapeutic implications. Gastroenterology. 1996 Mar; 110(3): 926-38. PMID: 8608904, PII: S0016508596001394, ISSN: 0016-5085.

Cover TL, Blaser MJ. Helicobacter pylori infection, a paradigm for chronic mucosal inflammation: pathogenesis and implications for eradication and prevention. Adv Intern Med. 1996; 41: 85-117. PMID: 8903587, ISSN: 0065-2822.

Ghiara P, Marchetti M, Blaser MJ, Tummuru MK, Cover TL, Segal ED, Tompkins LS, Rappuoli R. Role of the Helicobacter pylori virulence factors vacuolating cytotoxin, CagA, and urease in a mouse model of disease. Infect. Immun. 1995 Oct; 63(10): 4154-60. PMID: 7558333, PMCID: PMC173584, ISSN: 0019-9567.

Atherton JC, Cao P, Peek RM, Tummuru MK, Blaser MJ, Cover TL. Mosaicism in vacuolating cytotoxin alleles of Helicobacter pylori. Association of specific vacA types with cytotoxin production and peptic ulceration. J. Biol. Chem. 1995 Jul 7/28/1995; 270(30): 17771-7. PMID: 7629077, ISSN: 0021-9258.

Garner JA, Cover TL. Analysis of genetic diversity in cytotoxin-producing and non-cytotoxin-producing Helicobacter pylori strains. J. Infect. Dis. 1995 Jul; 172(1): 290-3. PMID: 7797934, ISSN: 0022-1899.

Cover TL, Glupczynski Y, Lage AP, Burette A, Tummuru MK, Perez-Perez GI, Blaser MJ. Serologic detection of infection with cagA+ Helicobacter pylori strains. J. Clin. Microbiol. 1995 Jun; 33(6): 1496-500. PMID: 7650174, PMCID: PMC228203, ISSN: 0095-1137.

Blaser MJ, Perez-Perez GI, Kleanthous H, Cover TL, Peek RM, Chyou PH, Stemmermann GN, Nomura A. Infection with Helicobacter pylori strains possessing cagA is associated with an increased risk of developing adenocarcinoma of the stomach. Cancer Res. 1995 May 5/15/1995; 55(10): 2111-5. PMID: 7743510, ISSN: 0008-5472.

Peek RM, Miller GG, Tham KT, Pérez-Pérez GI, Cover TL, Atherton JC, Dunn GD, Blaser MJ. Detection of Helicobacter pylori gene expression in human gastric mucosa. J. Clin. Microbiol. 1995 Jan; 33(1): 28-32. PMID: 7699060, PMCID: PMC227873, ISSN: 0095-1137.

McGowan CC, Cover TL, Blaser MJ. The proton pump inhibitor omeprazole inhibits acid survival of Helicobacter pylori by a urease-independent mechanism. Gastroenterology. 1994 Nov; 107(5): 1573-8. PMID: 7926527, PII: S0016508594003501, ISSN: 0016-5085.

McGowan CC, Cover TL, Blaser MJ. The proton pump inhibitor omeprazole inhibits acid survival of Helicobacter pylori by a urease-independent mechanism. Gastroenterology. 1994 Sep; 107(3): 738-43. PMID: 8076759, ISSN: 0016-5085.

Tummuru MK, Cover TL, Blaser MJ. Mutation of the cytotoxin-associated cagA gene does not affect the vacuolating cytotoxin activity of Helicobacter pylori. Infect. Immun. 1994 Jun; 62(6): 2609-13. PMID: 8188385, PMCID: PMC186552, ISSN: 0019-9567.

Pérez-Pérez GI, Brown WR, Cover TL, Dunn BE, Cao P, Blaser MJ. Correlation between serological and mucosal inflammatory responses to Helicobacter pylori. Clin. Diagn. Lab. Immunol. 1994 May; 1(3): 325-9. PMID: 7496970, PMCID: PMC368256, ISSN: 1071-412X.

Cover TL, Tummuru MK, Cao P, Thompson SA, Blaser MJ. Divergence of genetic sequences for the vacuolating cytotoxin among Helicobacter pylori strains. J. Biol. Chem. 1994 Apr 4/8/1994; 269(14): 10566-73. PMID: 8144644, ISSN: 0021-9258.

Cover TL, Cao P, Lind CD, Tham KT, Blaser MJ. Correlation between vacuolating cytotoxin production by Helicobacter pylori isolates in vitro and in vivo. Infect. Immun. 1993 Dec; 61(12): 5008-12. PMID: 8225576, PMCID: PMC281276, ISSN: 0019-9567.

Blaser MJ, Kobayashi K, Cover TL, Cao P, Feurer ID, Pérez-Pérez GI. Helicobacter pylori infection in Japanese patients with adenocarcinoma of the stomach. Int. J. Cancer. 1993 Nov 11/11/1993; 55(5): 799-802. PMID: 8244577, ISSN: 0020-7136.

Tummuru MK, Cover TL, Blaser MJ. Cloning and expression of a high-molecular-mass major antigen of Helicobacter pylori: evidence of linkage to cytotoxin production. Infect. Immun. 1993 May; 61(5): 1799-809. PMID: 8478069, PMCID: PMC280768, ISSN: 0019-9567.

Cover TL, Reddy LY, Blaser MJ. Effects of ATPase inhibitors on the response of HeLa cells to Helicobacter pylori vacuolating toxin. Infect. Immun. 1993 Apr; 61(4): 1427-31. PMID: 8454346, PMCID: PMC281381, ISSN: 0019-9567.

Cover TL, Vaughn SG, Cao P, Blaser MJ. Potentiation of Helicobacter pylori vacuolating toxin activity by nicotine and other weak bases. J. Infect. Dis. 1992 Nov; 166(5): 1073-8. PMID: 1402018, ISSN: 0022-1899.

Cover TL, Cao P, Murthy UK, Sipple MS, Blaser MJ. Serum neutralizing antibody response to the vacuolating cytotoxin of Helicobacter pylori. J. Clin. Invest. 1992 Sep; 90(3): 913-8. PMID: 1522241, PMCID: PMC329946, DOI: 10.1172/JCI115967, ISSN: 0021-9738.

Pérez-Pérez GI, Olivares AZ, Cover TL, Blaser MJ. Characteristics of Helicobacter pylori variants selected for urease deficiency. Infect. Immun. 1992 Sep; 60(9): 3658-63. PMID: 1500174, PMCID: PMC257374, ISSN: 0019-9567.

Cover TL, Halter SA, Blaser MJ. Characterization of HeLa cell vacuoles induced by Helicobacter pylori broth culture supernatant. Hum. Pathol. 1992 Sep; 23(9): 1004-10. PMID: 1381332, PII: 0046-8177(92)90261-Z, ISSN: 0046-8177.

Cover TL, Blaser MJ. Purification and characterization of the vacuolating toxin from Helicobacter pylori. J. Biol. Chem. 1992 May 5/25/1992; 267(15): 10570-5. PMID: 1587837, ISSN: 0021-9258.

Cover TL, Blaser MJ. Helicobacter pylori and gastroduodenal disease. Annu. Rev. Med. 1992; 43: 135-45. PMID: 1580578, DOI: 10.1146/annurev.me.43.020192.001031, ISSN: 0066-4219.

Cover TL, Puryear W, Perez-Perez GI, Blaser MJ. Effect of urease on HeLa cell vacuolation induced by Helicobacter pylori cytotoxin. Infect. Immun. 1991 Apr; 59(4): 1264-70. PMID: 2004808, PMCID: PMC257837, ISSN: 0019-9567.

Cover TL, Perez-Perez GI, Blaser MJ. Evaluation of cytotoxic activity in fecal filtrates from patients with Campylobacter jejuni or Campylobacter coli enteritis. FEMS Microbiol. Lett. 1990 Aug; 58(3): 301-4. PMID: 2227365, ISSN: 0378-1097.

Cover TL, Dooley CP, Blaser MJ. Characterization of and human serologic response to proteins in Helicobacter pylori broth culture supernatants with vacuolizing cytotoxin activity. Infect. Immun. 1990 Mar; 58(3): 603-10. PMID: 2307514, PMCID: PMC258508, ISSN: 0019-9567.

Cover TL, Aber RC. Yersinia enterocolitica. N. Engl. J. Med. 1989 Jul 7/6/1989; 321(1): 16-24. PMID: 2659991, DOI: 10.1056/NEJM198907063210104, ISSN: 0028-4793.

Cover TL, Blaser MJ. The pathobiology of Campylobacter infections in humans. Annu. Rev. Med. 1989; 40: 269-85. PMID: 2658752, DOI: 10.1146/annurev.me.40.020189.001413, ISSN: 0066-4219.

Available Postdoctoral Position Details
Posted: 4/12/2015
Postdoctoral position available to study the molecular pathogenesis of Helicobacter pylori infection and mechanisms by which H. pylori infection leads to gastric cancer. Areas of interest in the laboratory include a broad range of topics, including investigation of two H. pylori toxins (VacA and CagA), investigation of a type IV secretion system encoded by the H. pylori cag pathogenicity island, regulation of H. pylori gene expression, and comparative genome sequence analysis of H. pylori strains isolated from patients with different disease states. Please send CV to Dr. Timothy L. Cover, Division of Infectious Diseases, A2200 Medical Center North, Vanderbilt University School of Medicine, Nashville, TN 37232. Email: timothy.L.cover@vanderbilt.edu.