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Raymond M Welsh PhD

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

    InstitutionUMMS - School of Medicine
    DepartmentPathology
    DivisionAcademic Pathology

    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


    Collapse Biography 
    Collapse education and training
    University of Massachusetts Amherst, Amherst, MA, United StatesBSMicrobiology
    Rensselaer Polytechnic Institute, Troy, NY, United StatesPH DMicrobiology & Virology

    Collapse Overview 
    Collapse overview

    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.




    Collapse Bibliographic 
    Collapse selected publications
    Publications listed below are automatically derived from MEDLINE/PubMed and other sources, which might result in incorrect or missing publications. Faculty can login to make corrections and additions.
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    1. Jangalwe S, Kapoor VN, Xu J, Girnius N, Kennedy NJ, Edwards YJK, Welsh RM, Davis RJ, Brehm MA. Cutting Edge: Early Attrition of Memory T Cells during Inflammation and Costimulation Blockade Is Regulated Concurrently by Proapoptotic Proteins Fas and Bim. J Immunol. 2019 Jan 04. PMID: 30610162.
      View in: PubMed
    2. Hatfield SD, Daniels KA, O'Donnell CL, Waggoner SN, Welsh RM. Weak vaccinia virus-induced NK cell regulation of CD4 T cells is associated with reduced NK cell differentiation and cytolytic activity. Virology. 2018 06; 519:131-144. PMID: 29715623.
      View in: PubMed
    3. Aslan N, Watkin LB, Gil A, Mishra R, Clark FG, Welsh RM, Ghersi D, Luzuriaga K, Selin LK. Severity of Acute Infectious Mononucleosis Correlates with Cross-Reactive Influenza CD8 T-Cell Receptor Repertoires. MBio. 2017 Dec 05; 8(6). PMID: 29208744.
      View in: PubMed
    4. Shin HM, Kapoor VN, Kim G, Li P, Kim HR, Suresh M, Kaech SM, Wherry EJ, Selin LK, Leonard WJ, Welsh RM, Berg LJ. Transient expression of ZBTB32 in anti-viral CD8+ T cells limits the magnitude of the effector response and the generation of memory. PLoS Pathog. 2017 Aug; 13(8):e1006544. PMID: 28827827.
      View in: PubMed
    5. Wells AC, Daniels KA, Angelou CC, Fagerberg E, Burnside AS, Markstein M, Alfandari D, Welsh RM, Pobezinskaya EL, Pobezinsky LA. Modulation of let-7 miRNAs controls the differentiation of effector CD8 T cells. Elife. 2017 07 24; 6. PMID: 28737488.
      View in: PubMed
    6. Che JW, Daniels KA, Selin LK, Welsh RM. Heterologous Immunity and Persistent Murine Cytomegalovirus Infection. J Virol. 2017 Jan 15; 91(2). PMID: 27807227.
      View in: PubMed
    7. Urban SL, Berg LJ, Welsh RM. Type 1 interferon licenses naïve CD8 T cells to mediate anti-viral cytotoxicity. Virology. 2016 Jun; 493:52-9. PMID: 26999026.
      View in: PubMed
    8. Nayar R, Schutten E, Jangalwe S, Durost PA, Kenney LL, Conley JM, Daniels K, Brehm MA, Welsh RM, Berg LJ. IRF4 Regulates the Ratio of T-Bet to Eomesodermin in CD8+ T Cells Responding to Persistent LCMV Infection. PLoS One. 2015; 10(12):e0144826. PMID: 26714260.
      View in: PubMed
    9. Moriwaki K, Farias Luz N, Balaji S, De Rosa MJ, O'Donnell CL, Gough PJ, Bertin J, Welsh RM, Chan FK. The Mitochondrial Phosphatase PGAM5 Is Dispensable for Necroptosis but Promotes Inflammasome Activation in Macrophages. J Immunol. 2016 Jan 01; 196(1):407-15. PMID: 26582950.
      View in: PubMed
    10. Che JW, Selin LK, Welsh RM. Evaluation of non-reciprocal heterologous immunity between unrelated viruses. Virology. 2015 Aug; 482:89-97. PMID: 25838115.
      View in: PubMed
    11. Rydyznski C, Daniels KA, Karmele EP, Brooks TR, Mahl SE, Moran MT, Li C, Sutiwisesak R, Welsh RM, Waggoner SN. Generation of cellular immune memory and B-cell immunity is impaired by natural killer cells. Nat Commun. 2015 Feb 27; 6:6375. PMID: 25721802.
      View in: PubMed
    12. Che JW, Kraft AR, Selin LK, Welsh RM. Regulatory T cells resist virus infection-induced apoptosis. J Virol. 2015 Feb; 89(4):2112-20. PMID: 25473049.
      View in: PubMed
    13. Urban SL, Welsh RM. Out-of-sequence signal 3 as a mechanism for virus-induced immune suppression of CD8 T cell responses. PLoS Pathog. 2014 Sep; 10(9):e1004357. PMID: 25255454.
      View in: PubMed
    14. Daniels KA, Hatfield SD, Welsh RM, Brehm MA. MHC basis of T cell-dependent heterologous immunity to arenaviruses. Virology. 2014 Sep; 464-465:213-7. PMID: 25094042.
      View in: PubMed
    15. Kapoor VN, Shin HM, Cho OH, Berg LJ, Kang J, Welsh RM. Regulation of tissue-dependent differences in CD8+ T cell apoptosis during viral infection. J Virol. 2014 Sep 1; 88(17):9490-503. PMID: 24942579.
      View in: PubMed
    16. Nayar R, Schutten E, Bautista B, Daniels K, Prince AL, Enos M, Brehm MA, Swain SL, Welsh RM, Berg LJ. Graded levels of IRF4 regulate CD8+ T cell differentiation and expansion, but not attrition, in response to acute virus infection. J Immunol. 2014 Jun 15; 192(12):5881-93. PMID: 24835398.
      View in: PubMed
    17. Mishra R, Welsh R, Szomolanyi-Tsuda E. NK cells and virus-related cancers. Crit Rev Oncog. 2014; 19(1-2):107-19. PMID: 24941377.
      View in: PubMed
    18. Cornberg M, Kenney LL, Chen AT, Waggoner SN, Kim SK, Dienes HP, Welsh RM, Selin LK. Clonal exhaustion as a mechanism to protect against severe immunopathology and death from an overwhelming CD8 T cell response. Front Immunol. 2013; 4:475. PMID: 24391647.
      View in: PubMed
    19. Waggoner SN, Daniels KA, Welsh RM. Therapeutic depletion of natural killer cells controls persistent infection. J Virol. 2014 Feb; 88(4):1953-60. PMID: 24284324.
      View in: PubMed
    20. Raval FM, Mishra R, Garcea RL, Welsh RM, Szomolanyi-Tsuda E. Long-lasting T cell-independent IgG responses require MyD88-mediated pathways and are maintained by high levels of virus persistence. MBio. 2013 Nov 05; 4(6):e00812-13. PMID: 24194540.
      View in: PubMed
    21. Shen ZT, Nguyen TT, Daniels KA, Welsh RM, Stern LJ. Disparate epitopes mediating protective heterologous immunity to unrelated viruses share peptide-MHC structural features recognized by cross-reactive T cells. J Immunol. 2013 Nov 15; 191(10):5139-52. PMID: 24127554.
      View in: PubMed
    22. Shin HM, Kapoor VN, Guan T, Kaech SM, Welsh RM, Berg LJ. Epigenetic modifications induced by Blimp-1 Regulate CD8? T cell memory progression during acute virus infection. Immunity. 2013 Oct 17; 39(4):661-75. PMID: 24120360.
      View in: PubMed
    23. 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. PMID: 23772039.
      View in: PubMed
    24. 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. PMID: 23740988.
      View in: PubMed
    25. 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. PMID: 23190301.
      View in: PubMed
    26. 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. PMID: 23217614.
      View in: PubMed
    27. 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; 8(4):e1002633. PMID: 22536152.
      View in: PubMed
    28. 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. PMID: 22438562.
      View in: PubMed
    29. Welsh RM, Bahl K, Marshall HD, Urban SL. Type 1 interferons and antiviral CD8 T-cell responses. PLoS Pathog. 2012 Jan; 8(1):e1002352. PMID: 22241987.
      View in: PubMed
    30. 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 Dec 25; 13(2):129-35. PMID: 22197977.
      View in: PubMed
    31. Waggoner SN, Cornberg M, Selin LK, Welsh RM. Natural killer cells act as rheostats modulating antiviral T cells. Nature. 2011 Nov 20; 481(7381):394-8. PMID: 22101430.
      View in: PubMed
    32. 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. PMID: 21471240.
      View in: PubMed
    33. 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. PMID: 21271821.
      View in: PubMed
    34. Priyadharshini B, Welsh RM, Greiner DL, Gerstein RM, Brehm MA. Maturation-dependent licensing of naive T cells for rapid TNF production. PLoS One. 2010 Nov 24; 5(11):e15038. PMID: 21124839.
      View in: PubMed
    35. 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. PMID: 21048112.
      View in: PubMed
    36. 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. PMID: 20729210.
      View in: PubMed
    37. 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. PMID: 20592282.
      View in: PubMed
    38. 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 27; 6(5):e1000924. PMID: 20523894.
      View in: PubMed
    39. 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; 120(6):1925-38. PMID: 20440077.
      View in: PubMed
    40. Welsh RM, Che JW, Brehm MA, Selin LK. Heterologous immunity between viruses. Immunol Rev. 2010 May; 235(1):244-66. PMID: 20536568.
      View in: PubMed
    41. 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. PMID: 20348239.
      View in: PubMed
    42. 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. PMID: 20200235.
      View in: PubMed
    43. 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. PMID: 20164414.
      View in: PubMed
    44. 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. PMID: 19812167.
      View in: PubMed
    45. Welsh RM. Blimp hovers over T cell immunity. Immunity. 2009 Aug 21; 31(2):178-80. PMID: 19699168.
      View in: PubMed
    46. 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. PMID: 19542462.
      View in: PubMed
    47. Welsh RM, Selin LK. Attrition of memory CD8 T cells. Nature. 2009 Jun 4; 459(7247):E3-4; discussion E4. PMID: 19494857.
      View in: PubMed
    48. 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. PMID: 19380803.
      View in: PubMed
    49. 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. PMID: 19193795.
      View in: PubMed
    50. 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. PMID: 18981164.
      View in: PubMed
    51. 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. PMID: 18509089.
      View in: PubMed
    52. Welsh RM, Seedhom MO. Lymphocytic choriomeningitis virus (LCMV): propagation, quantitation, and storage. Curr Protoc Microbiol. 2008 Feb; Chapter 15:Unit 15A.1. PMID: 18770534.
      View in: PubMed
    53. 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. PMID: 17982052.
      View in: PubMed
    54. 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. PMID: 17785507.
      View in: PubMed
    55. 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. PMID: 17609270.
      View in: PubMed
    56. Welsh RM, Fujinami RS. Pathogenic epitopes, heterologous immunity and vaccine design. Nat Rev Microbiol. 2007 Jul; 5(7):555-63. PMID: 17558423.
      View in: PubMed
    57. 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. PMID: 17404295.
      View in: PubMed
    58. 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. PMID: 17163452.
      View in: PubMed
    59. 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. PMID: 16973964.
      View in: PubMed
    60. 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. PMID: 16824126.
      View in: PubMed
    61. 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. PMID: 16611887.
      View in: PubMed
    62. 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. PMID: 16614754.
      View in: PubMed
    63. Welsh RM. Private specificities of heterologous immunity. Curr Opin Immunol. 2006 Jun; 18(3):331-7. PMID: 16597500.
      View in: PubMed
    64. 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. PMID: 16547266.
      View in: PubMed
    65. 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. PMID: 16424185.
      View in: PubMed
    66. 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. PMID: 16424186.
      View in: PubMed
    67. 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. PMID: 17048707.
      View in: PubMed
    68. 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. PMID: 16308574.
      View in: PubMed
    69. 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. PMID: 16210607.
      View in: PubMed
    70. 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. PMID: 15952447.
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    71. 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. PMID: 15905505.
      View in: PubMed
    72. 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. PMID: 15710651.
      View in: PubMed
    73. 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. PMID: 15690085.
      View in: PubMed
    74. 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. PMID: 15634878.
      View in: PubMed
    75. 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. PMID: 15470058.
      View in: PubMed
    76. 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. PMID: 15528078.
      View in: PubMed
    77. Szomolanyi-Tsuda E, Welsh RM. Judging a virus by its cover. J Clin Invest. 2004 Oct; 114(7):895-7. PMID: 15467827.
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    78. 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. PMID: 15210808.
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    79. 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. PMID: 15134774.
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    80. Brehm MA, Selin LK, Welsh RM. CD8 T cell responses to viral infections in sequence. Cell Microbiol. 2004 May; 6(5):411-21. PMID: 15056212.
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    81. 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. PMID: 15070899.
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    82. 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. PMID: 14978120.
      View in: PubMed
    83. Selin LK, Welsh RM. Plasticity of T cell memory responses to viruses. Immunity. 2004 Jan; 20(1):5-16. PMID: 14738760.
      View in: PubMed
    84. Welsh RM, Selin LK, Szomolanyi-Tsuda E. Immunological memory to viral infections. Annu Rev Immunol. 2004; 22:711-43. PMID: 15032594.
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    85. 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. PMID: 14507643.
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    86. 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. PMID: 12847230.
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    87. Welsh RM, Rothman AL. Dengue immune response: low affinity, high febrility. Nat Med. 2003 Jul; 9(7):820-2. PMID: 12835692.
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    88. 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. PMID: 12753740.
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    89. 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. PMID: 12682237.
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    90. 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. PMID: 12646622.
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    91. Welsh RM, Stepp SE, Szomolanyi-Tsuda E. B cell memory: sapping the T cell. Nat Med. 2003 Feb; 9(2):164-6. PMID: 12563333.
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    92. 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. PMID: 12505520.
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    93. 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. PMID: 12077233.
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    94. 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. PMID: 12055213.
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    95. 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. PMID: 12055626.
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    96. Welsh RM, Selin LK. No one is naive: the significance of heterologous T-cell immunity. Nat Rev Immunol. 2002 Jun; 2(6):417-26. PMID: 12093008.
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    97. Welsh RM, Stepp SE, Szomolanyi-Tsuda E, Peacock CD. Tumor viral escape from inhibited T cells. Nat Immunol. 2002 Feb; 3(2):112-4. PMID: 11812984.
      View in: PubMed
    98. Welsh RM. Assessing CD8 T cell number and dysfunction in the presence of antigen. J Exp Med. 2001 Mar 5; 193(5):F19-22. PMID: 11238597.
      View in: PubMed
    99. 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. PMID: 10772974.
      View in: PubMed
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