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    Raymond M Welsh PhD

    TitleProfessor
    InstitutionUniversity of Massachusetts Medical School
    DepartmentPathology
    AddressUniversity of Massachusetts Medical School
    55 Lake Avenue North
    Worcester MA 01655
    Phone508-856-5819
      Other Positions
      InstitutionUMMS - School of Medicine
      DepartmentMicrobiology and Physiological Systems

      InstitutionUMMS - Graduate School of Biomedical Sciences
      DepartmentImmunology and Virology

      InstitutionUMMS - Graduate School of Biomedical Sciences
      DepartmentMD/PhD Program

      InstitutionUMMS - Graduate School of Biomedical Sciences
      DepartmentMolecular Genetics and Microbiology

      InstitutionUMMS - Programs, Centers and Institutes
      DepartmentCenter for AIDS Research

        Overview 
        Narrative

        Academic Background

        PhD, 1972, University of Massachusetts, Amherst

        Dr. Ray Welsh

        Viral Immunology and pathogenesis

        The focus of our research is on analyzing basic mechanisms of viral immunology and immunopathology. To do this, we study mouse model infections with a variety of viruses, including lymphocytic choriomeningitis virus, murine cytomegalovirus (MCMV), vaccinia virus, Pichinde virus, mouse hepatitis virus, herpes simplex virus, polyomavirus, Sindbis virus, and others. Our laboratory has demonstrated that natural killer cells become activated during viral infections and contribute to natural resistance to viruses, though they sometimes can act antagonistically by interfering with anti-viral T cell responses. We identified a receptor on NK cells that enables them to control MCMV infection. We also examine the dynamics of CD8 T cell activation, death by apoptosis, and memory in viral infections. A major area of research defined by our lab is on heterologous immunity, whereby the T and B cell immune response elicited by one virus may alter the pathogenesis of disease initiated by a second unrelated virus. This is sometimes mediated by T cells cross-reactive between unrelated viruses. We have also shown that virus-specific T cells often cross-react with allogeneic tissue, and we are studying the role of those cross-reactive T cells in allograft rejection.



        Bibliographic 
        selected publications
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        1. Mishra R, Polic B, Welsh RM, Szomolanyi-Tsuda E. Inflammatory cytokine-mediated evasion of virus-induced tumors from NK cell control. J Immunol. 2013 Jul 15; 191(2):961-70.
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        2. Rodriguez IA, Welsh RM. Possible role of a cell surface carbohydrate in evolution of resistance to viral infections in old world primates. J Virol. 2013 Aug; 87(15):8317-26.
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        3. Priyadharshini B, Thornley TB, Daniels KA, Cuthbert A, Welsh RM, Greiner DL, Brehm MA. Alloreactive CD8 T cells rescued from apoptosis during co-stimulation blockade by Toll-like receptor stimulation remain susceptible to Fas-induced cell death. Immunology. 2013 Apr; 138(4):322-32.
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        4. Welsh RM, Waggoner SN. NK cells controlling virus-specific T cells: Rheostats for acute vs. persistent infections. Virology. 2013 Jan 5; 435(1):37-45.
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        5. Chen AT, Cornberg M, Gras S, Guillonneau C, Rossjohn J, Trees A, Emonet S, de la Torre JC, Welsh RM, Selin LK. Loss of anti-viral immunity by infection with a virus encoding a cross-reactive pathogenic epitope. PLoS Pathog. 2012 Apr; 8(4):e1002633.
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        6. Seedhom MO, Mathurin KS, Kim SK, Welsh RM. Increased protection from vaccinia virus infection in mice genetically prone to lymphoproliferative disorders. J Virol. 2012 Jun; 86(11):6010-22.
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        7. Welsh RM, Bahl K, Marshall HD, Urban SL. Type 1 interferons and antiviral CD8 T-cell responses. PLoS Pathog. 2012 Jan; 8(1):e1002352.
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        8. Kincaid EZ, Che JW, York I, Escobar H, Reyes-Vargas E, Delgado JC, Welsh RM, Karow ML, Murphy AJ, Valenzuela DM, Yancopoulos GD, Rock KL. Mice completely lacking immunoproteasomes show major changes in antigen presentation. Nat Immunol. 2011; 13(2):129-35.
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        9. Waggoner SN, Cornberg M, Selin LK, Welsh RM. Natural killer cells act as rheostats modulating antiviral T cells. Nature. 2012 Jan 19; 481(7381):394-8.
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        10. Marshall HD, Urban SL, Welsh RM. Virus-induced transient immune suppression and the inhibition of T cell proliferation by type I interferon. J Virol. 2011 Jun; 85(12):5929-39.
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        11. Calcagno C, Puzone R, Pearson YE, Cheng Y, Ghersi D, Selin LK, Welsh RM, Celada F. Computer simulations of heterologous immunity: Highlights of an interdisciplinary cooperation. Autoimmunity. 2011 Jun; 44(4):304-14.
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        12. Priyadharshini B, Welsh RM, Greiner DL, Gerstein RM, Brehm MA. Maturation-Dependent Licensing of Naive T Cells for Rapid TNF Production. PLoS One. 2010; 5(11):e15038.
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        13. Clute SC, Naumov YN, Watkin LB, Aslan N, Sullivan JL, Thorley-Lawson DA, Luzuriaga K, Welsh RM, Puzone R, Celada F, Selin LK. Broad cross-reactive TCR repertoires recognizing dissimilar epstein-barr and influenza A virus epitopes. J Immunol. 2010 Dec 1; 185(11):6753-64.
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        14. Shen ZT, Brehm MA, Daniels KA, Sigalov AB, Selin LK, Welsh RM, Stern LJ. Bi-specific MHC heterodimers for characterization of cross-reactive T cells. J Biol Chem. 2010 Oct 22; 285(43):33144-53.
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        15. Brehm MA, Daniels KA, Priyadharshini B, Thornley TB, Greiner DL, Rossini AA, Welsh RM. Allografts stimulate cross-reactive virus-specific memory CD8 T cells with private specificity. Am J Transplant. 2010 Aug; 10(8):1738-48.
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        16. Marshall HD, Prince AL, Berg LJ, Welsh RM. IFN-alpha beta and self-MHC divert CD8 T cells into a distinct differentiation pathway characterized by rapid acquisition of effector functions. J Immunol. 2010 Aug 1; 185(3):1419-28.
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        17. Mishra R, Chen AT, Welsh RM, Szomolanyi-Tsuda E. NK cells and gammadelta T cells mediate resistance to polyomavirus-induced tumors. PLoS Pathog. 2010 May; 6(5):e1000924.
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        18. Waggoner SN, Taniguchi RT, Mathew PA, Kumar V, Welsh RM. Absence of mouse 2B4 promotes NK cell-mediated killing of activated CD8+ T cells, leading to prolonged viral persistence and altered pathogenesis. J Clin Invest. 2010 Jun 1; 120(6):1925-38.
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        19. Welsh RM, Che JW, Brehm MA, Selin LK. Heterologous immunity between viruses. Immunol Rev. 2010 May; 235(1):244-66.
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        20. Nie S, Lin SJ, Kim SK, Welsh RM, Selin LK. Pathological features of heterologous immunity are regulated by the private specificities of the immune repertoire. Am J Pathol. 2010 May; 176(5):2107-12.
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        21. Bahl K, Hüebner A, Davis RJ, Welsh RM. Analysis of apoptosis of memory T cells and dendritic cells during the early stages of viral infection or exposure to toll-like receptor agonists. J Virol. 2010 May; 84(10):4866-77.
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        22. Cornberg M, Clute SC, Watkin LB, Saccoccio FM, Kim SK, Naumov YN, Brehm MA, Aslan N, Welsh RM, Selin LK. CD8 T cell cross-reactivity networks mediate heterologous immunity in human EBV and murine vaccinia virus infections. J Immunol. 2010 Mar 15; 184(6):2825-38.
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        23. Seedhom MO, Jellison ER, Daniels KA, Welsh RM. High frequencies of virus-specific CD8+ T-cell precursors. J Virol. 2009 Dec; 83(24):12907-16.
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        24. Welsh RM. Blimp hovers over T cell immunity. Immunity. 2009 Aug 21; 31(2):178-80.
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        25. Guay HM, Mishra R, Garcea RL, Welsh RM, Szomolanyi-Tsuda E. Generation of protective T cell-independent antiviral antibody responses in SCID mice reconstituted with follicular or marginal zone B cells. J Immunol. 2009 Jul 1; 183(1):518-23.
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        26. Welsh RM, Selin LK. Attrition of memory CD8 T cells. Nature. 2009 Jun 4; 459(7247):E3-4; discussion E4.
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        27. Miller DM, Thornley TB, Pearson T, Kruger AJ, Yamazaki M, Shultz LD, Welsh RM, Brehm MA, Rossini AA, Greiner DL. TLR agonists prevent the establishment of allogeneic hematopoietic chimerism in mice treated with costimulation blockade. J Immunol. 2009 May 1; 182(9):5547-59.
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        28. Mathurin KS, Martens GW, Kornfeld H, Welsh RM. CD4 T-cell-mediated heterologous immunity between mycobacteria and poxviruses. J Virol. 2009 Apr; 83(8):3528-39.
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        29. Naumov YN, Naumova EN, Yassai MB, Kota K, Welsh RM, Selin LK. Multiple glycines in TCR alpha-chains determine clonally diverse nature of human T cell memory to influenza A virus. J Immunol. 2008 Nov 15; 181(10):7407-19.
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        30. Lin SJ, Chen AT, Welsh RM. Immune system derived from homeostatic proliferation generates normal CD8 T-cell memory but altered repertoires and diminished heterologous immune responses. Blood. 2008 Aug 1; 112(3):680-9.
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        31. Welsh RM, Seedhom MO. Lymphocytic choriomeningitis virus (LCMV): propagation, quantitation, and storage. Curr Protoc Microbiol. 2008 Feb; Chapter 15:Unit 15A.1.
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        32. Thornley TB, Phillips NE, Beaudette-Zlatanova BC, Markees TG, Bahl K, Brehm MA, Shultz LD, Kurt-Jones EA, Mordes JP, Welsh RM, Rossini AA, Greiner DL. Type 1 IFN mediates cross-talk between innate and adaptive immunity that abrogates transplantation tolerance. J Immunol. 2007 Nov 15; 179(10):6620-9.
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        33. Lin SJ, Peacock CD, Bahl K, Welsh RM. Programmed death-1 (PD-1) defines a transient and dysfunctional oligoclonal T cell population in acute homeostatic proliferation. J Exp Med. 2007 Oct 1; 204(10):2321-33.
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        34. Abdel-Motal UM, Guay HM, Wigglesworth K, Welsh RM, Galili U. Immunogenicity of influenza virus vaccine is increased by anti-gal-mediated targeting to antigen-presenting cells. J Virol. 2007 Sep; 81(17):9131-41.
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        35. Welsh RM, Fujinami RS. Pathogenic epitopes, heterologous immunity and vaccine design. Nat Rev Microbiol. 2007 Jul; 5(7):555-63.
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        36. Guay HM, Andreyeva TA, Garcea RL, Welsh RM, Szomolanyi-Tsuda E. MyD88 is required for the formation of long-term humoral immunity to virus infection. J Immunol. 2007 Apr 15; 178(8):5124-31.
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        37. Jellison ER, Guay HM, Szomolanyi-Tsuda E, Welsh RM. Dynamics and magnitude of virus-induced polyclonal B cell activation mediated by BCR-independent presentation of viral antigen. Eur J Immunol. 2007 Jan; 37(1):119-28.
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        38. Brehm MA, Mangada J, Markees TG, Pearson T, Daniels KA, Thornley TB, Welsh RM, Rossini AA, Greiner DL. Rapid quantification of naive alloreactive T cells by TNF-alpha production and correlation with allograft rejection in mice. Blood. 2007 Jan 15; 109(2):819-26.
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        39. Selin LK, Brehm MA, Naumov YN, Cornberg M, Kim SK, Clute SC, Welsh RM. Memory of mice and men: CD8+ T-cell cross-reactivity and heterologous immunity. Immunol Rev. 2006 Jun; 211:164-81.
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        40. Szomolanyi-Tsuda E, Liang X, Welsh RM, Kurt-Jones EA, Finberg RW. Role for TLR2 in NK cell-mediated control of murine cytomegalovirus in vivo. J Virol. 2006 May; 80(9):4286-91.
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        41. Cornberg M, Chen AT, Wilkinson LA, Brehm MA, Kim SK, Calcagno C, Ghersi D, Puzone R, Celada F, Welsh RM, Selin LK. Narrowed TCR repertoire and viral escape as a consequence of heterologous immunity. J Clin Invest. 2006 May; 116(5):1443-56.
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        42. Welsh RM. Private specificities of heterologous immunity. Curr Opin Immunol. 2006 Jun; 18(3):331-7.
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        43. Bahl K, Kim SK, Calcagno C, Ghersi D, Puzone R, Celada F, Selin LK, Welsh RM. IFN-induced attrition of CD8 T cells in the presence or absence of cognate antigen during the early stages of viral infections. J Immunol. 2006 Apr 1; 176(7):4284-95.
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        44. Thornley TB, Brehm MA, Markees TG, Shultz LD, Mordes JP, Welsh RM, Rossini AA, Greiner DL. TLR agonists abrogate costimulation blockade-induced prolongation of skin allografts. J Immunol. 2006 Feb 1; 176(3):1561-70.
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        45. Atherly LO, Brehm MA, Welsh RM, Berg LJ. Tec kinases Itk and Rlk are required for CD8+ T cell responses to virus infection independent of their role in CD4+ T cell help. J Immunol. 2006 Feb 1; 176(3):1571-81.
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        46. Welsh RM, Kim SK, Cornberg M, Clute SC, Selin LK, Naumov YN. The privacy of T cell memory to viruses. Curr Top Microbiol Immunol. 2006; 311:117-53.
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        47. Clute SC, Watkin LB, Cornberg M, Naumov YN, Sullivan JL, Luzuriaga K, Welsh RM, Selin LK. Cross-reactive influenza virus-specific CD8+ T cells contribute to lymphoproliferation in Epstein-Barr virus-associated infectious mononucleosis. J Clin Invest. 2005 Dec; 115(12):3602-12.
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        48. Brehm MA, Daniels KA, Welsh RM. Rapid production of TNF-alpha following TCR engagement of naive CD8 T cells. J Immunol. 2005 Oct 15; 175(8):5043-9.
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        49. Khachemoune A, Welsh R, Ehrsam E. An elderly woman with a non-healing ulcer. Marjolin's ulcer. Am Fam Physician. 2005 Jun 1; 71(11):2163-5.
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        50. Brehm MA, Daniels KA, Ortaldo JR, Welsh RM. Rapid conversion of effector mechanisms from NK to T cells during virus-induced lysis of allogeneic implants in vivo. J Immunol. 2005 Jun 1; 174(11):6663-71.
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        51. Kim SK, Cornberg M, Wang XZ, Chen HD, Selin LK, Welsh RM. Private specificities of CD8 T cell responses control patterns of heterologous immunity. J Exp Med. 2005 Feb 21; 201(4):523-33.
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        52. Yang H, Kim SK, Kim M, Reche PA, Morehead TJ, Damon IK, Welsh RM, Reinherz EL. Antiviral chemotherapy facilitates control of poxvirus infections through inhibition of cellular signal transduction. J Clin Invest. 2005 Feb; 115(2):379-87.
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        53. Jellison ER, Kim SK, Welsh RM. Cutting edge: MHC class II-restricted killing in vivo during viral infection. J Immunol. 2005 Jan 15; 174(2):614-8.
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        54. Wang XZ, Brehm MA, Welsh RM. Preapoptotic phenotype of viral epitope-specific CD8 T cells precludes memory development and is an intrinsic property of the epitope. J Immunol. 2004 Oct 15; 173(8):5138-47.
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        55. Selin LK, Cornberg M, Brehm MA, Kim SK, Calcagno C, Ghersi D, Puzone R, Celada F, Welsh RM. CD8 memory T cells: cross-reactivity and heterologous immunity. Semin Immunol. 2004 Oct; 16(5):335-47.
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        56. Szomolanyi-Tsuda E, Welsh RM. Judging a virus by its cover. J Clin Invest. 2004 Oct; 114(7):895-7.
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        57. Gavanescu I, Pihan G, Halilovic E, Szomolanyi-Tsuda E, Welsh RM, Doxsey S. Mycoplasma infection induces a scleroderma-like centrosome autoantibody response in mice. Clin Exp Immunol. 2004 Aug; 137(2):288-97.
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        58. Peacock CD, Welsh RM. Origin and fate of lymphocytic choriomeningitis virus-specific CD8+ T cells coexpressing the inhibitory NK cell receptor Ly49G2. J Immunol. 2004 Jul 1; 173(1):478-84.
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        59. Welsh RM, Bahl K, Wang XZ. Apoptosis and loss of virus-specific CD8+ T-cell memory. Curr Opin Immunol. 2004 Jun; 16(3):271-6.
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        60. Brehm MA, Selin LK, Welsh RM. CD8 T cell responses to viral infections in sequence. Cell Microbiol. 2004 May; 6(5):411-21.
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        61. Kim M, Yang H, Kim SK, Reche PA, Tirabassi RS, Hussey RE, Chishti Y, Rheinwald JG, Morehead TJ, Zech T, Damon IK, Welsh RM, Reinherz EL. Biochemical and functional analysis of smallpox growth factor (SPGF) and anti-SPGF monoclonal antibodies. J Biol Chem. 2004 Jun 11; 279(24):25838-48.
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        62. Kim SK, Welsh RM. Comprehensive early and lasting loss of memory CD8 T cells and functional memory during acute and persistent viral infections. J Immunol. 2004 Mar 1; 172(5):3139-50.
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        63. Welsh RM, Selin LK, Szomolanyi-Tsuda E. Immunological memory to viral infections. Annu Rev Immunol. 2004; 22:711-43.
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        64. Selin LK, Welsh RM. Plasticity of T cell memory responses to viruses. Immunity. 2004 Jan; 20(1):5-16.
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        65. Chen HD, Fraire AE, Joris I, Welsh RM, Selin LK. Specific history of heterologous virus infections determines anti-viral immunity and immunopathology in the lung. Am J Pathol. 2003 Oct; 163(4):1341-55.
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        66. Peacock CD, Kim SK, Welsh RM. Attrition of virus-specific memory CD8+ T cells during reconstitution of lymphopenic environments. J Immunol. 2003 Jul 15; 171(2):655-63.
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        67. Welsh RM, Rothman AL. Dengue immune response: low affinity, high febrility. Nat Med. 2003 Jul; 9(7):820-2.
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        68. Wang XZ, Stepp SE, Brehm MA, Chen HD, Selin LK, Welsh RM. Virus-specific CD8 T cells in peripheral tissues are more resistant to apoptosis than those in lymphoid organs. Immunity. 2003 May; 18(5):631-42.
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        69. Brehm MA, Markees TG, Daniels KA, Greiner DL, Rossini AA, Welsh RM. Direct visualization of cross-reactive effector and memory allo-specific CD8 T cells generated in response to viral infections. J Immunol. 2003 Apr 15; 170(8):4077-86.
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        70. Zipris D, Hillebrands JL, Welsh RM, Rozing J, Xie JX, Mordes JP, Greiner DL, Rossini AA. Infections that induce autoimmune diabetes in BBDR rats modulate CD4+CD25+ T cell populations. J Immunol. 2003 Apr 1; 170(7):3592-602.
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        71. Welsh RM, Stepp SE, Szomolanyi-Tsuda E. B cell memory: sapping the T cell. Nat Med. 2003 Feb; 9(2):164-6.
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        72. Peacock CD, Xu W, Stepp SE, Welsh RM. Dynamics of Ly49 expressing cytotoxic lymphocyte subsets in response to virus infection. Microbes Infect. 2002 Dec; 4(15):1481-90.
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        73. Kim SK, Brehm MA, Welsh RM, Selin LK. Dynamics of memory T cell proliferation under conditions of heterologous immunity and bystander stimulation. J Immunol. 2002 Jul 1; 169(1):90-8.
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        74. Forman D, Welsh RM, Markees TG, Woda BA, Mordes JP, Rossini AA, Greiner DL. Viral abrogation of stem cell transplantation tolerance causes graft rejection and host death by different mechanisms. J Immunol. 2002 Jun 15; 168(12):6047-56.
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        75. Brehm MA, Pinto AK, Daniels KA, Schneck JP, Welsh RM, Selin LK. T cell immunodominance and maintenance of memory regulated by unexpectedly cross-reactive pathogens. Nat Immunol. 2002 Jul; 3(7):627-34.
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        76. Welsh RM, Selin LK. No one is naive: the significance of heterologous T-cell immunity. Nat Rev Immunol. 2002 Jun; 2(6):417-26.
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        77. Welsh RM, Stepp SE, Szomolanyi-Tsuda E, Peacock CD. Tumor viral escape from inhibited T cells. Nat Immunol. 2002 Feb; 3(2):112-4.
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        78. McNally JM, Welsh RM. Bystander T cell activation and attrition. Curr Top Microbiol Immunol. 2002; 263:29-41.
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        79. Chen HD, Fraire AE, Joris I, Brehm MA, Welsh RM, Selin LK. Memory CD8+ T cells in heterologous antiviral immunity and immunopathology in the lung. Nat Immunol. 2001 Nov; 2(11):1067-76.
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        80. Daniels KA, Devora G, Lai WC, O'Donnell CL, Bennett M, Welsh RM. Murine cytomegalovirus is regulated by a discrete subset of natural killer cells reactive with monoclonal antibody to Ly49H. J Exp Med. 2001 Jul 2; 194(1):29-44.
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        81. McNally JM, Zarozinski CC, Lin MY, Brehm MA, Chen HD, Welsh RM. Attrition of bystander CD8 T cells during virus-induced T-cell and interferon responses. J Virol. 2001 Jul; 75(13):5965-76.
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        82. Selin LK, Santolucito PA, Pinto AK, Szomolanyi-Tsuda E, Welsh RM. Innate immunity to viruses: control of vaccinia virus infection by gamma delta T cells. J Immunol. 2001 Jun 1; 166(11):6784-94.
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        83. Welsh RM. Assessing CD8 T cell number and dysfunction in the presence of antigen. J Exp Med. 2001 Mar 5; 193(5):F19-22.
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        84. Szomolanyi-Tsuda E, Brien JD, Dorgan JE, Garcea RL, Woodland RT, Welsh RM. Antiviral T-cell-independent type 2 antibody responses induced in vivo in the absence of T and NK cells. Virology. 2001 Feb 15; 280(2):160-8.
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        85. Varga SM, Selin LK, Welsh RM. Independent regulation of lymphocytic choriomeningitis virus-specific T cell memory pools: relative stability of CD4 memory under conditions of CD8 memory T cell loss. J Immunol. 2001 Feb 1; 166(3):1554-61.
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        86. Turgeon NA, Iwakoshi NN, Phillips NE, Meyers WC, Welsh RM, Greiner DL, Mordes JP, Rossini AA. Viral infection abrogates CD8(+) T-cell deletion induced by costimulation blockade. J Surg Res. 2000 Sep; 93(1):63-9.
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        87. Peacock CD, Lin MY, Ortaldo JR, Welsh RM. The virus-specific and allospecific cytotoxic T-lymphocyte response to lymphocytic choriomeningitis virus is modified in a subpopulation of CD8(+) T cells coexpressing the inhibitory major histocompatibility complex class I receptor Ly49G2. J Virol. 2000 Aug; 74(15):7032-8.
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        88. Lin MY, Selin LK, Welsh RM. Evolution of the CD8 T-cell repertoire during infections. Microbes Infect. 2000 Jul; 2(9):1025-39.
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        89. Szomolanyi-Tsuda E, Brien JD, Dorgan JE, Welsh RM, Garcea RL. The role of CD40-CD154 interaction in antiviral T cell-independent IgG responses. J Immunol. 2000 Jun 1; 164(11):5877-82.
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        90. Varga SM, Welsh RM. High frequency of virus-specific interleukin-2-producing CD4(+) T cells and Th1 dominance during lymphocytic choriomeningitis virus infection. J Virol. 2000 May; 74(9):4429-32.
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        91. Welsh RM, McNally JM, Brehm MA, Selin LK. Consequences of cross-reactive and bystander CTL responses during viral infections. Virology. 2000 Apr 25; 270(1):4-8.
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        92. Zarozinski CC, McNally JM, Lohman BL, Daniels KA, Welsh RM. Bystander sensitization to activation-induced cell death as a mechanism of virus-induced immune suppression. J Virol. 2000 Apr; 74(8):3650-8.
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        93. Welsh RM, Markees TG, Woda BA, Daniels KA, Brehm MA, Mordes JP, Greiner DL, Rossini AA. Virus-induced abrogation of transplantation tolerance induced by donor-specific transfusion and anti-CD154 antibody. J Virol. 2000 Mar; 74(5):2210-8.
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        94. Selin LK, Lin MY, Kraemer KA, Pardoll DM, Schneck JP, Varga SM, Santolucito PA, Pinto AK, Welsh RM. Attrition of T cell memory: selective loss of LCMV epitope-specific memory CD8 T cells following infections with heterologous viruses. Immunity. 1999 Dec; 11(6):733-42.
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        95. Welsh RM, McNally JM. Immune deficiency, immune silencing, and clonal exhaustion of T cell responses during viral infections. Curr Opin Microbiol. 1999 Aug; 2(4):382-7.
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        96. Tay CH, Yu LY, Kumar V, Mason L, Ortaldo JR, Welsh RM. The role of LY49 NK cell subsets in the regulation of murine cytomegalovirus infections. J Immunol. 1999 Jan 15; 162(2):718-26.
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        97. Lin MY, Welsh RM. Stability and diversity of T cell receptor repertoire usage during lymphocytic choriomeningitis virus infection of mice. J Exp Med. 1998 Dec 7; 188(11):1993-2005.
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        98. Selin LK, Varga SM, Wong IC, Welsh RM. Protective heterologous antiviral immunity and enhanced immunopathogenesis mediated by memory T cell populations. J Exp Med. 1998 Nov 2; 188(9):1705-15.
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        99. Lohman BL, Welsh RM. Apoptotic regulation of T cells and absence of immune deficiency in virus-infected gamma interferon receptor knockout mice. J Virol. 1998 Oct; 72(10):7815-21.
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        100. Varga SM, Welsh RM. Detection of a high frequency of virus-specific CD4+ T cells during acute infection with lymphocytic choriomeningitis virus. J Immunol. 1998 Oct 1; 161(7):3215-8.
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        101. Szomolanyi-Tsuda E, Le QP, Garcea RL, Welsh RM. T-Cell-independent immunoglobulin G responses in vivo are elicited by live-virus infection but not by immunization with viral proteins or virus-like particles. J Virol. 1998 Aug; 72(8):6665-70.
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        102. Szomolanyi-Tsuda E, Welsh RM. T-cell-independent antiviral antibody responses. Curr Opin Immunol. 1998 Aug; 10(4):431-5.
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        103. Varga SM, Welsh RM. Stability of virus-specific CD4+ T cell frequencies from acute infection into long term memory. J Immunol. 1998 Jul 1; 161(1):367-74.
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        104. Welsh RM, O'Donnell CL, Reed DJ, Rother RP. Evaluation of the Galalpha1-3Gal epitope as a host modification factor eliciting natural humoral immunity to enveloped viruses. J Virol. 1998 Jun; 72(6):4650-6.
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        105. Tay CH, Szomolanyi-Tsuda E, Welsh RM. Control of infections by NK cells. Curr Top Microbiol Immunol. 1998; 230:193-220.
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        106. Welsh RM, Lin MY, Lohman BL, Varga SM, Zarozinski CC, Selin LK. Alpha beta and gamma delta T-cell networks and their roles in natural resistance to viral infections. Immunol Rev. 1997 Oct; 159:79-93.
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        107. Selin LK, Welsh RM. Cytolytically active memory CTL present in lymphocytic choriomeningitis virus-immune mice after clearance of virus infection. J Immunol. 1997 Jun 1; 158(11):5366-73.
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        108. Zarozinski CC, Welsh RM. Minimal bystander activation of CD8 T cells during the virus-induced polyclonal T cell response. J Exp Med. 1997 May 5; 185(9):1629-39.
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        109. Tay CH, Welsh RM. Distinct organ-dependent mechanisms for the control of murine cytomegalovirus infection by natural killer cells. J Virol. 1997 Jan; 71(1):267-75.
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        110. Lohman BL, Razvi ES, Welsh RM. T-lymphocyte downregulation after acute viral infection is not dependent on CD95 (Fas) receptor-ligand interactions. J Virol. 1996 Nov; 70(11):8199-203.
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        111. Selin LK, Vergilis K, Welsh RM, Nahill SR. Reduction of otherwise remarkably stable virus-specific cytotoxic T lymphocyte memory by heterologous viral infections. J Exp Med. 1996 Jun 1; 183(6):2489-99.
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        112. Szomolanyi-Tsuda E, Welsh RM. T cell-independent antibody-mediated clearance of polyoma virus in T cell-deficient mice. J Exp Med. 1996 Feb 1; 183(2):403-11.
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        113. Varga SM, Welsh RM. The CD45RB-associated epitope defined by monoclonal antibody CZ-1 is an activation and memory marker for mouse CD4 T cells. Cell Immunol. 1996 Jan 10; 167(1):56-62.
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        114. Welsh RM, Selin LK, Razvi ES. Role of apoptosis in the regulation of virus-induced T cell responses, immune suppression, and memory. J Cell Biochem. 1995 Oct; 59(2):135-42.
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        115. Brutkiewicz RR, Welsh RM. Major histocompatibility complex class I antigens and the control of viral infections by natural killer cells. J Virol. 1995 Jul; 69(7):3967-71.
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        116. Razvi ES, Jiang Z, Woda BA, Welsh RM. Lymphocyte apoptosis during the silencing of the immune response to acute viral infections in normal, lpr, and Bcl-2-transgenic mice. Am J Pathol. 1995 Jul; 147(1):79-91.
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        117. Zarozinski CC, Fynan EF, Selin LK, Robinson HL, Welsh RM. Protective CTL-dependent immunity and enhanced immunopathology in mice immunized by particle bombardment with DNA encoding an internal virion protein. J Immunol. 1995 Apr 15; 154(8):4010-7.
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        118. Tay CH, Welsh RM, Brutkiewicz RR. NK cell response to viral infections in beta 2-microglobulin-deficient mice. J Immunol. 1995 Jan 15; 154(2):780-9.
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        119. Razvi ES, Welsh RM, McFarland HI. In vivo state of antiviral CTL precursors. Characterization of a cycling cell population containing CTL precursors in immune mice. J Immunol. 1995 Jan 15; 154(2):620-32.
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        120. Razvi ES, Welsh RM. Apoptosis in viral infections. Adv Virus Res. 1995; 45:1-60.
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        121. Welsh RM, O'Donnell CL, Shultz LD. Antiviral activity of NK 1.1+ natural killer cells in C57BL/6 scid mice infected with murine cytomegalovirus. Nat Immun. 1994 Sep-Oct; 13(5):239-45.
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        122. Selin LK, Welsh RM. Specificity and editing by apoptosis of virus-induced cytotoxic T lymphocytes. Curr Opin Immunol. 1994 Aug; 6(4):553-9.
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        123. Selin LK, Nahill SR, Welsh RM. Cross-reactivities in memory cytotoxic T lymphocyte recognition of heterologous viruses. J Exp Med. 1994 Jun 1; 179(6):1933-43.
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        124. Szomolanyi-Tsuda E, Dundon PL, Joris I, Shultz LD, Woda BA, Welsh RM. Acute, lethal, natural killer cell-resistant myeloproliferative disease induced by polyomavirus in severe combined immunodeficient mice. Am J Pathol. 1994 Feb; 144(2):359-71.
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        125. Nishioka WK, Welsh RM. Susceptibility to cytotoxic T lymphocyte-induced apoptosis is a function of the proliferative status of the target. J Exp Med. 1994 Feb 1; 179(2):769-74.
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        126. Brown DW, Welsh RM, Like AA. Infection of peripancreatic lymph nodes but not islets precedes Kilham rat virus-induced diabetes in BB/Wor rats. J Virol. 1993 Oct; 67(10):5873-8.
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        127. Razvi ES, Welsh RM. Programmed cell death of T lymphocytes during acute viral infection: a mechanism for virus-induced immune deficiency. J Virol. 1993 Oct; 67(10):5754-65.
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        128. Brutkiewicz RR, O'Donnell CL, Maciaszek JW, Welsh RM, Vargas-Cortes M. The monoclonal antibody CZ-1 identifies a mouse CD45-associated epitope expressed on interleukin-2-responsive cells. Eur J Immunol. 1993 Oct; 23(10):2427-33.
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        129. Nishioka WK, Welsh RM. B cells induce apoptosis via a novel mechanism in fibroblasts infected with mouse hepatitis virus. Nat Immun. 1993 May-Jun; 12(3):113-27.
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        130. Nahill SR, Welsh RM. High frequency of cross-reactive cytotoxic T lymphocytes elicited during the virus-induced polyclonal cytotoxic T lymphocyte response. J Exp Med. 1993 Feb 1; 177(2):317-27.
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        131. Nahill SR, Welsh RM. Polyclonality of the cytotoxic T lymphocyte response to virus infection. Proc Soc Exp Biol Med. 1992 Sep; 200(4):453-7.
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        132. McFarland HI, Nahill SR, Maciaszek JW, Welsh RM. CD11b (Mac-1): a marker for CD8+ cytotoxic T cell activation and memory in virus infection. J Immunol. 1992 Aug 15; 149(4):1326-33.
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        133. Brutkiewicz RR, Klaus SJ, Welsh RM. Window of vulnerability of vaccinia virus-infected cells to natural killer (NK) cell-mediated cytolysis correlates with enhanced NK cell triggering and is concomitant with a decrease in H-2 class I antigen expression. Nat Immun. 1992 Jul-Aug; 11(4):203-14.
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        134. Vargas-Cortes M, O'Donnell CL, Maciaszek JW, Welsh RM. Generation of "natural killer cell-escape" variants of Pichinde virus during acute and persistent infections. J Virol. 1992 Apr; 66(4):2532-5.
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        135. Vargas-Cortes M, O'Donnell CL, Appel MC, Maciaszek JW, Yurkunas KS, Welsh RM. A lymphocyte differentiation and activation antigen, CZ-1, that distinguishes between CD8+ and unstimulated CD4+ T lymphocytes. Eur J Immunol. 1992 Apr; 22(4):1043-7.
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        136. Nishioka WK, Welsh RM. Inhibition of cytotoxic T lymphocyte-induced target cell DNA fragmentation, but not lysis, by inhibitors of DNA topoisomerases I and II. J Exp Med. 1992 Jan 1; 175(1):23-7.
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        137. Guberski DL, Thomas VA, Shek WR, Like AA, Handler ES, Rossini AA, Wallace JE, Welsh RM. Induction of type I diabetes by Kilham's rat virus in diabetes-resistant BB/Wor rats. Science. 1991 Nov 15; 254(5034):1010-3.
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        138. Brubaker JO, Chong KT, Welsh RM. Lymphokine-activated killer cells are rejected in vivo by activated natural killer cells. J Immunol. 1991 Aug 15; 147(4):1439-44.
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        139. Welsh RM, Brubaker JO, Vargas-Cortes M, O'Donnell CL. Natural killer (NK) cell response to virus infections in mice with severe combined immunodeficiency. The stimulation of NK cells and the NK cell-dependent control of virus infections occur independently of T and B cell function. J Exp Med. 1991 May 1; 173(5):1053-63.
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        140. Welsh RM, Nishioka WK, Antia R, Dundon PL. Mechanism of killing by virus-induced cytotoxic T lymphocytes elicited in vivo. J Virol. 1990 Aug; 64(8):3726-33.
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        141. Welsh RM, Dundon PL, Eynon EE, Brubaker JO, Koo GC, O'Donnell CL. Demonstration of the antiviral role of natural killer cells in vivo with a natural killer cell-specific monoclonal antibody (NK 1.1). Nat Immun Cell Growth Regul. 1990; 9(2):112-20.
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        142. Natuk RJ, Bukowski JF, Brubaker JO, Welsh RM. Antiviral effect of lymphokine-activated killer cells: chemotaxis and homing to sites of virus infection. J Virol. 1989 Nov; 63(11):4969-71.
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        143. Yang HY, Dundon PL, Nahill SR, Welsh RM. Virus-induced polyclonal cytotoxic T lymphocyte stimulation. J Immunol. 1989 Mar 1; 142(5):1710-8.
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        144. Bukowski JF, Yang H, Welsh RM. Antiviral effect of lymphokine-activated killer cells: characterization of effector cells mediating prophylaxis. J Virol. 1988 Oct; 62(10):3642-8.
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        145. McIntyre KW, Natuk RJ, Biron CA, Kase K, Greenberger J, Welsh RM. Blastogenesis of large granular lymphocytes in nonlymphoid organs. J Leukoc Biol. 1988 Jun; 43(6):492-501.
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        146. Natuk RJ, Welsh RM. Chemotactic effect of human recombinant interleukin 2 on mouse activated large granular lymphocytes. J Immunol. 1987 Oct 15; 139(8):2737-43.
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        147. Welsh RM, Doe WF. Cytotoxic cells induced during lymphocytic choriomeningitis virus infection of mice: natural killer cell activity in cultured spleen leukocytes concomitant with T-cell-dependent immune interferon production. Infect Immun. 1980 Nov; 30(2):473-83.
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        148. Kiessling RW, Welsh RM. Killing of normal cells by activated mouse natural killer cells: evidence for two patterns of genetic regulation of lysis. Int J Cancer. 1980 May 15; 25(5):611-5.
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        149. Zinkernagel RM, Althage A, Callahan G, Welsh RM. On the immunocompetence of H-2 incompatible irradiation bone marrow chimeras. J Immunol. 1980 May; 124(5):2356-65.
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        150. Altman A, Cardenas JM, Welsh RM, Katz DH. The biological effects of allogeneic effect factor (AEF) on T lymphocytes. III.--Interferon does not contribute to the biological activities displayed by AEF on both T and B lymphocytes. Ann Immunol (Paris). 1980 May-Jun; 131C(3):335-47.
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        151. Welsh RM, Hallenbeck LA. Effect of virus infections on target cell susceptibility to natural killer cell-mediated lysis. J Immunol. 1980 May; 124(5):2491-7.
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        152. Welsh RM, Kiessling RW. Natural killer cell response to lymphocytic choriomeningitis virus in beige mice. Scand J Immunol. 1980; 11(4):363-7.
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        153. Welsh RM, Buchmeier MJ. Protein analysis of defective interfering lymphocytic choriomeningitis virus and persistently infected cells. Virology. 1979 Jul 30; 96(2):503-15.
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        154. Welsh RM, Zinkernagel RM, Hallenbeck LA. Cytotoxic cells induced during lymphocytic choriomeningitis virus infection of mice. II. "Specificities" of the natural killer cells. J Immunol. 1979 Feb; 122(2):475-81.
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        155. Welsh RM. Mouse natural killer cells: induction specificity, and function. J Immunol. 1978 Nov; 121(5):1631-5.
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        156. Welsh RM. Cytotoxic cells induced during lymphocytic choriomeningitis virus infection of mice. I. Characterization of natural killer cell induction. J Exp Med. 1978 Jul 1; 148(1):163-81.
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        157. Welsh RM, Zinkernagel RM. Heterospecific cytotoxic cell activity induced during the first three days of acute lymphocytic choriomeningitis virus infection in mice. Nature. 1977 Aug 18; 268(5621):646-8.
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        158. Oldstone MB, Holmstoen J, Welsh RM. Alterations of acetylcholine enzymes in neuroblastoma cells persistently infected with lymphocytic choriomeningitis virus. J Cell Physiol. 1977 Jun; 91(3):459-72.
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        159. Welsh RM, Jensen FC, Cooper NR, Oldstone MB. Inactivation of lysis of oncornaviruses by human serum. Virology. 1976 Oct 15; 74(2):432-40.
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        160. Cooper NR, Jensen FC, Welsh RM, Oldstone MB. Lysis of RNA tumor viruses by human serum: direct antibody-independent triggering of the classical complement pathway. J Exp Med. 1976 Oct 1; 144(4):970-84.
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        161. Welsh RM, Lampert PW, Burner PA, Oldstone MB. Antibody-complement interactions with purified lymphocytic choriomeningitis virus. Virology. 1976 Aug; 73(1):59-71.
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        162. Oldstone MB, Perrin LH, Welsh RM. Potential pathogenic mechanisms of injury in amyotrophic lateral sclerosis. UCLA Forum Med Sci. 1976; (19):251-62.
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        163. Welsh RM, Trowbridge RS, Kowalski JB, O'Connell CM, Pfau CJ. Amantadine hydrochloride inhibition of early and late stages of lymphocytic choriomeningitis virus-cell interactions. Virology. 1971 Sep; 45(3):679-86.
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