{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,10]],"date-time":"2026-10-10T18:17:20Z","timestamp":1791656240275,"version":"4.3.4"},"reference-count":220,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2019,6,13]],"date-time":"2019-06-13T00:00:00Z","timestamp":1560384000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell. Infect. Microbiol."],"DOI":"10.3389\/fcimb.2019.00207","type":"journal-article","created":{"date-parts":[[2019,6,13]],"date-time":"2019-06-13T06:16:13Z","timestamp":1560406573000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":262,"title":["The PD-1\/PD-L1 Axis and Virus Infections: A Delicate Balance"],"prefix":"10.3389","volume":"9","author":[{"given":"G\u00fcnther","family":"Sch\u00f6nrich","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Martin J.","family":"Raftery","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2019,6,13]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"1593","DOI":"10.1084\/jem.20161760","article-title":"Human memory CD8 T cell effector potential is epigenetically preserved during in vivo homeostasis","volume":"214","author":"Abdelsamed","year":"2017","journal-title":"J. Exp. Med."},{"key":"B2","doi-asserted-by":"crossref","first-page":"765","DOI":"10.1093\/intimm\/8.5.765","article-title":"Expression of the PD-1 antigen on the surface of stimulated mouse T and B lymphocytes","volume":"8","author":"Agata","year":"1996","journal-title":"Int. Immunol"},{"key":"B3","doi-asserted-by":"publisher","first-page":"4749","DOI":"10.1073\/pnas.1718217115","article-title":"Role of PD-1 during effector CD8 T cell differentiation","volume":"115","author":"Ahn","year":"2018","journal-title":"Proc. Natl. Acad. Sci. U.S.A"},{"key":"B4","doi-asserted-by":"publisher","first-page":"4525","DOI":"10.4049\/jimmunol.176.8.4525","article-title":"CD8 T cells specific for lymphocytic choriomeningitis virus require type I IFN receptor for clonal expansion","volume":"176","author":"Aichele","year":"2006","journal-title":"J. Immunol."},{"key":"B5","doi-asserted-by":"publisher","first-page":"e1005224","DOI":"10.1371\/journal.ppat.1005224","article-title":"PD-L1 expression on retrovirus-infected cells mediates immune escape from CD8+ T cell killing","volume":"11","author":"Akhmetzyanova","year":"2015","journal-title":"PLoS Pathog"},{"key":"B6","doi-asserted-by":"publisher","first-page":"362","DOI":"10.1038\/nature24633","article-title":"Origin and differentiation of human memory CD8 T cells after vaccination","volume":"552","author":"Akondy","year":"2017","journal-title":"Nature"},{"key":"B7","doi-asserted-by":"publisher","first-page":"4184","DOI":"10.1128\/JVI.02290-10","article-title":"The role of LAT in increased CD8+ T cell exhaustion in trigeminal ganglia of mice latently infected with herpes simplex virus 1","volume":"85","author":"Allen","year":"2011","journal-title":"J. Virol."},{"key":"B8","doi-asserted-by":"publisher","first-page":"1232","DOI":"10.3389\/fimmu.2017.01232","article-title":"Interferon-lambdas: front-line guardians of immunity and homeostasis in the respiratory tract","volume":"8","author":"Andreakos","year":"2017","journal-title":"Front. Immunol."},{"key":"B9","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1126\/science.1254803","article-title":"Skin-resident memory CD8(+) T cells trigger a state of tissue-wide pathogen alert","volume":"346","author":"Ariotti","year":"2014","journal-title":"Science"},{"key":"B10","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1007\/82_2010_29","article-title":"Varicella-zoster virus T cell tropism and the pathogenesis of skin infection","volume":"342","author":"Arvin","year":"2010","journal-title":"Curr. Top. Microbiol. Immunol."},{"key":"B11","doi-asserted-by":"publisher","first-page":"1052","DOI":"10.1016\/j.immuni.2016.04.022","article-title":"Costimulatory and coinhibitory receptor pathways in infectious disease","volume":"44","author":"Attanasio","year":"2016","journal-title":"Immunity"},{"key":"B12","doi-asserted-by":"publisher","first-page":"682","DOI":"10.1038\/nature04444","article-title":"Restoring function in exhausted CD8 T cells during chronic viral infection","volume":"439","author":"Barber","year":"2006","journal-title":"Nature"},{"key":"B13","doi-asserted-by":"publisher","first-page":"573","DOI":"10.1172\/JCI99035","article-title":"Unravelling the fate of functional PD1+ T cells in chronic viral hepatitis","volume":"128","author":"Barnes","year":"2018","journal-title":"J. Clin. Invest."},{"key":"B14","doi-asserted-by":"publisher","first-page":"551","DOI":"10.1007\/s00281-017-0637-x","article-title":"Molecular pathogenesis of viral hemorrhagic fever","volume":"39","author":"Basler","year":"2017","journal-title":"Semin. Immunopathol."},{"key":"B15","doi-asserted-by":"publisher","first-page":"4836","DOI":"10.4049\/jimmunol.180.7.4836","article-title":"Dendritic cell programming by cytomegalovirus stunts naive T cell responses via the PD-L1\/PD-1 pathway","volume":"180","author":"Benedict","year":"2008","journal-title":"J. Immunol."},{"key":"B16","doi-asserted-by":"publisher","first-page":"358","DOI":"10.1016\/j.immuni.2016.07.008","article-title":"Bioenergetic insufficiencies due to metabolic alterations regulated by the inhibitory receptor PD-1 are an early driver of CD8(+) T cell exhaustion","volume":"45","author":"Bengsch","year":"2016","journal-title":"Immunity"},{"key":"B17","doi-asserted-by":"publisher","first-page":"204","DOI":"10.1126\/science.aal1962","article-title":"Mouse models of acute and chronic hepacivirus infection","volume":"357","author":"Billerbeck","year":"2017","journal-title":"Science"},{"key":"B18","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1007\/978-3-642-56055-2_2","article-title":"Innate immune responses to LCMV infections: natural killer cells and cytokines","volume":"263","author":"Biron","year":"2002","journal-title":"Curr. Top. Microbiol. Immunol."},{"key":"B19","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1038\/ni.1679","article-title":"Coregulation of CD8+ T cell exhaustion by multiple inhibitory receptors during chronic viral infection","volume":"10","author":"Blackburn","year":"2009","journal-title":"Nat. Immunol."},{"key":"B20","doi-asserted-by":"publisher","first-page":"4386","DOI":"10.1128\/JVI.02524-08","article-title":"Impact of epitope escape on PD-1 expression and CD8 T-cell exhaustion during chronic infection","volume":"83","author":"Blattman","year":"2009","journal-title":"J. Virol."},{"key":"B21","doi-asserted-by":"publisher","first-page":"132","DOI":"10.1016\/j.clim.2008.05.009","article-title":"PDL-1 upregulation on monocytes and T cells by HIV via type I interferon: restricted expression of type I interferon receptor by CCR5-expressing leukocytes","volume":"129","author":"Boasso","year":"2008","journal-title":"Clin. Immunol."},{"key":"B22","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1016\/j.canlet.2015.02.027","article-title":"TLR3 triggering regulates PD-L1 (CD274) expression in human neuroblastoma cells","volume":"361","author":"Boes","year":"2015","journal-title":"Cancer Lett"},{"key":"B23","doi-asserted-by":"crossref","first-page":"1977","DOI":"10.1084\/jem.169.6.1977","article-title":"Tumor necrosis factor\/cachectin increases permeability of endothelial cell monolayers by a mechanism involving regulatory G proteins","volume":"169","author":"Brett","year":"1989","journal-title":"J. Exp. Med."},{"key":"B24","doi-asserted-by":"publisher","first-page":"20428","DOI":"10.1073\/pnas.0811139106","article-title":"IL-10 and PD-L1 operate through distinct pathways to suppress T-cell activity during persistent viral infection","volume":"105","author":"Brooks","year":"2008","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B25","doi-asserted-by":"publisher","first-page":"1675","DOI":"10.1172\/JCI26856","article-title":"Reprogramming of antiviral T cells prevents inactivation and restores T cell activity during persistent viral infection","volume":"116","author":"Brooks","year":"","journal-title":"J. Clin. Invest."},{"key":"B26","doi-asserted-by":"publisher","first-page":"1301","DOI":"10.1038\/nm1492","article-title":"Interleukin-10 determines viral clearance or persistence in vivo","volume":"12","author":"Brooks","year":"","journal-title":"Nat. Med."},{"key":"B27","doi-asserted-by":"publisher","first-page":"1257","DOI":"10.4049\/jimmunol.170.3.1257","article-title":"Blockade of programmed death-1 ligands on dendritic cells enhances T cell activation and cytokine production","volume":"170","author":"Brown","year":"2003","journal-title":"J. Immunol."},{"key":"B28","doi-asserted-by":"publisher","first-page":"397","DOI":"10.1016\/j.coi.2010.03.007","article-title":"Role of PD-1 in regulating acute infections","volume":"22","author":"Brown","year":"2010","journal-title":"Curr. Opin. Immunol."},{"key":"B29","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1016\/j.immuni.2007.05.016","article-title":"Programmed death-1 ligand 1 interacts specifically with the B7-1 costimulatory molecule to inhibit T cell responses","volume":"27","author":"Butte","year":"2007","journal-title":"Immunity"},{"key":"B30","doi-asserted-by":"publisher","first-page":"3567","DOI":"10.1016\/j.molimm.2008.05.014","article-title":"Interaction of human PD-L1 and B7-1","volume":"45","author":"Butte","year":"2008","journal-title":"Mol. Immunol."},{"key":"B31","doi-asserted-by":"publisher","first-page":"40","DOI":"10.1126\/science.1259925","article-title":"A neighborhood watch upholds local immune protection","volume":"346","author":"Carbone","year":"2014","journal-title":"Science"},{"key":"B32","doi-asserted-by":"publisher","first-page":"e39757","DOI":"10.1371\/journal.pone.0039757","article-title":"Blocking of PDL-1 interaction enhances primary and secondary CD8 T cell response to herpes simplex virus-1 infection","volume":"7","author":"Channappanavar","year":"2012","journal-title":"PLoS ONE"},{"key":"B33","doi-asserted-by":"publisher","first-page":"921","DOI":"10.1158\/2326-6066.CIR-17-0316","article-title":"PD-L1 binds to B7-1 only in Cis on the same cell surface","volume":"6","author":"Chaudhri","year":"2018","journal-title":"Cancer Immunol. Res."},{"key":"B34","doi-asserted-by":"publisher","first-page":"227","DOI":"10.1038\/nri3405","article-title":"Molecular mechanisms of T cell co-stimulation and co-inhibition","volume":"13","author":"Chen","year":"2013","journal-title":"Nat. Rev. Immunol."},{"key":"B35","doi-asserted-by":"publisher","first-page":"9127","DOI":"10.1128\/JVI.00587-11","article-title":"The herpes simplex virus 1 latency-associated transcript promotes functional exhaustion of virus-specific CD8+ T cells in latently infected trigeminal ganglia: a novel immune evasion mechanism","volume":"85","author":"Chentoufi","year":"2011","journal-title":"J. Virol"},{"key":"B36","doi-asserted-by":"publisher","first-page":"6682","DOI":"10.4049\/jimmunol.0900080","article-title":"PD-1-mediated suppression of IL-2 production induces CD8+ T cell anergy in vivo","volume":"182","author":"Chikuma","year":"2009","journal-title":"J. Immunol"},{"key":"B37","doi-asserted-by":"publisher","first-page":"2372","DOI":"10.1073\/pnas.1018515108","article-title":"Unexpected protective role for Toll-like receptor 3 in the arterial wall","volume":"108","author":"Cole","year":"2011","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B38","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1016\/j.coi.2017.07.016","article-title":"Check point inhibitors as therapies for infectious diseases","volume":"48","author":"Cox","year":"2017","journal-title":"Curr. Opin. Immunol."},{"key":"B39","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1016\/j.immuni.2014.01.005","article-title":"Molecular and transcriptional basis of CD4(+) T cell dysfunction during chronic infection","volume":"40","author":"Crawford","year":"2014","journal-title":"Immunity"},{"key":"B40","doi-asserted-by":"publisher","first-page":"562","DOI":"10.1038\/nm863","article-title":"Blockade of B7-H1 improves myeloid dendritic cell-mediated antitumor immunity","volume":"9","author":"Curiel","year":"2003","journal-title":"Nat. Med."},{"key":"B41","doi-asserted-by":"publisher","first-page":"54","DOI":"10.3389\/fimmu.2019.00054","article-title":"The PD-1\/PD-L1 pathway affects the expansion and function of cytotoxic CD8(+) T cells during an acute retroviral infection","volume":"10","author":"David","year":"2019","journal-title":"Front. Immunol."},{"key":"B42","doi-asserted-by":"publisher","first-page":"252","DOI":"10.1089\/jir.2014.0227","article-title":"Disease-promoting effects of type I interferons in viral, bacterial, and coinfections","volume":"35","author":"Davidson","year":"2015","journal-title":"J. Interferon Cytokine Res."},{"key":"B43","doi-asserted-by":"publisher","first-page":"350","DOI":"10.1038\/nature05115","article-title":"PD-1 expression on HIV-specific T cells is associated with T-cell exhaustion and disease progression","volume":"443","author":"Day","year":"2006","journal-title":"Nature"},{"key":"B44","doi-asserted-by":"publisher","first-page":"1130","DOI":"10.1016\/j.immuni.2012.08.021","article-title":"Network analysis reveals centrally connected genes and pathways involved in CD8+ T cell exhaustion versus memory","volume":"37","author":"Doering","year":"2012","journal-title":"Immunity"},{"key":"B45","doi-asserted-by":"publisher","first-page":"2461","DOI":"10.1084\/jem.20061462","article-title":"Resolution of a chronic viral infection after interleukin-10 receptor blockade","volume":"203","author":"Ejrnaes","year":"2006","journal-title":"J. Exp. Med."},{"key":"B46","doi-asserted-by":"publisher","first-page":"2967","DOI":"10.1172\/JCI62860","article-title":"Viral acute lower respiratory infections impair CD8+ T cells through PD-1","volume":"122","author":"Erickson","year":"2012","journal-title":"J. Clin. Invest."},{"key":"B47","doi-asserted-by":"publisher","first-page":"973","DOI":"10.1016\/j.immuni.2016.04.020","article-title":"CD28 costimulation: from mechanism to therapy","volume":"44","author":"Esensten","year":"2016","journal-title":"Immunity"},{"key":"B48","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1111\/j.1749-6632.2010.05919.x","article-title":"The role of the PD-1 pathway in autoimmunity and peripheral tolerance","volume":"1217","author":"Fife","year":"2011","journal-title":"Ann. N. Y. Acad. Sci"},{"key":"B49","doi-asserted-by":"publisher","first-page":"219","DOI":"10.1111\/j.1600-065X.2010.00923.x","article-title":"The PD-1 pathway in tolerance and autoimmunity","volume":"236","author":"Francisco","year":"2010","journal-title":"Immunol. Rev."},{"key":"B50","doi-asserted-by":"publisher","first-page":"3015","DOI":"10.1084\/jem.20090847","article-title":"PD-L1 regulates the development, maintenance, and function of induced regulatory T cells","volume":"206","author":"Francisco","year":"2009","journal-title":"J. Exp. Med."},{"key":"B51","doi-asserted-by":"publisher","first-page":"2485","DOI":"10.1084\/jem.20121015","article-title":"Programmed death 1 protects from fatal circulatory failure during systemic virus infection of mice","volume":"209","author":"Frebel","year":"2012","journal-title":"J. Exp. Med."},{"key":"B52","doi-asserted-by":"publisher","first-page":"875","DOI":"10.1016\/j.immuni.2017.04.025","article-title":"Interferon-lambda mediates non-redundant front-line antiviral protection against influenza virus infection without compromising host fitness","volume":"46","author":"Galani","year":"2017","journal-title":"Immunity"},{"key":"B53","doi-asserted-by":"publisher","first-page":"872","DOI":"10.1016\/j.bbmt.2009.03.022","article-title":"Increased programmed death-1 molecule expression in cytomegalovirus disease and acute graft-versus-host disease after allogeneic hematopoietic cell transplantation","volume":"15","author":"Gallez-Hawkins","year":"2009","journal-title":"Biol. Blood Marrow Transplant."},{"key":"B54","doi-asserted-by":"crossref","first-page":"1383","DOI":"10.1084\/jem.187.9.1383","article-title":"Induction and exhaustion of lymphocytic choriomeningitis virus-specific cytotoxic T lymphocytes visualized using soluble tetrameric major histocompatibility complex class I-peptide complexes","volume":"187","author":"Gallimore","year":"1998","journal-title":"J. Exp. Med."},{"key":"B55","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1016\/j.jaci.2018.11.048","article-title":"Inhibitory checkpoint receptors control CD8(+) resident memory T cells to prevent skin allergy","volume":"2019","author":"Gamradt","year":"2019","journal-title":"J. Allergy Clin. Immunol."},{"key":"B56","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1128\/JVI.02035-18","article-title":"Contrasting roles of the PD-1 signaling pathway in dendritic cell-mediated induction and regulation of HIV-1-specific effector T cell functions","volume":"93","author":"Garcia-Bates","year":"2019","journal-title":"J. Virol."},{"key":"B57","doi-asserted-by":"publisher","first-page":"1189","DOI":"10.1016\/j.celrep.2017.04.031","article-title":"Interferon receptor signaling pathways regulating PD-L1 and PD-L2 expression","volume":"19","author":"Garcia-Diaz","year":"2017","journal-title":"Cell Rep"},{"key":"B58","doi-asserted-by":"publisher","first-page":"e361","DOI":"10.1016\/S2352-3018(16)30087-X","article-title":"Estimates of global, regional, and national incidence, prevalence, and mortality of HIV, 1980-2015: the Global Burden of Disease Study 2015","volume":"3","year":"2016","journal-title":"Lancet HIV"},{"key":"B59","doi-asserted-by":"publisher","first-page":"524","DOI":"10.1038\/ni.1718","article-title":"Memory T cells in nonlymphoid tissue that provide enhanced local immunity during infection with herpes simplex virus","volume":"10","author":"Gebhardt","year":"2009","journal-title":"Nat. Immunol"},{"key":"B60","doi-asserted-by":"publisher","first-page":"eaau4255","DOI":"10.1126\/sciadv.aau4255","article-title":"Type III interferon signaling restricts enterovirus 71 infection of goblet cells","volume":"5","author":"Good","year":"2019","journal-title":"Sci. Adv."},{"key":"B61","doi-asserted-by":"publisher","first-page":"468","DOI":"10.1016\/j.jhep.2006.07.009","article-title":"Turning on the off switch: regulation of anti-viral T cell responses in the liver by the PD-1\/PD-L1 pathway","volume":"45","author":"Grakoui","year":"2006","journal-title":"J. Hepatol."},{"key":"B62","doi-asserted-by":"publisher","first-page":"1363","DOI":"10.1002\/eji.201343701","article-title":"Japanese encephalitis virus expands regulatory T cells by increasing the expression of PD-L1 on dendritic cells","volume":"44","author":"Gupta","year":"2014","journal-title":"Eur. J. Immunol."},{"key":"B63","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1146\/annurev-med-012017-043208","article-title":"CD8 T cell exhaustion in chronic infection and cancer: opportunities for interventions","volume":"69","author":"Hashimoto","year":"2018","journal-title":"Annu. Rev. Med."},{"key":"B64","doi-asserted-by":"publisher","first-page":"1467","DOI":"10.1038\/ni.3589","article-title":"Programs for the persistence, vigilance and control of human CD8(+) lung-resident memory T cells","volume":"17","author":"Hombrink","year":"2016","journal-title":"Nat. Immunol"},{"key":"B65","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1016\/j.immuni.2013.11.017","article-title":"Tuning of antigen sensitivity by T cell receptor-dependent negative feedback controls T cell effector function in inflamed tissues","volume":"40","author":"Honda","year":"2014","journal-title":"Immunity"},{"key":"B66","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1128\/mBio.00917-17","article-title":"Kaposi's sarcoma-associated herpesvirus increases PD-L1 and proinflammatory cytokine expression in human monocytes","volume":"8","author":"Host","year":"2017","journal-title":"MBio"},{"key":"B67","doi-asserted-by":"publisher","first-page":"224","DOI":"10.1089\/vim.2016.0122","article-title":"Clinical significance of dynamics of programmed death ligand-1 expression on circulating CD14(+) monocytes and CD19(+) B cells with the progression of hepatitis B virus infection","volume":"30","author":"Huang","year":"2017","journal-title":"Viral Immunol"},{"key":"B68","doi-asserted-by":"crossref","first-page":"3887","DOI":"10.1002\/j.1460-2075.1992.tb05481.x","article-title":"Induced expression of PD-1, a novel member of the immunoglobulin gene superfamily, upon programmed cell death","volume":"11","author":"Ishida","year":"1992","journal-title":"EMBO J"},{"key":"B69","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1084\/jem.20022235","article-title":"PD-1 inhibits antiviral immunity at the effector phase in the liver","volume":"198","author":"Iwai","year":"2003","journal-title":"J. Exp. Med."},{"key":"B70","doi-asserted-by":"publisher","first-page":"3711","DOI":"10.4049\/jimmunol.1700417","article-title":"PD-L1\/B7-H1 inhibits viral clearance by macrophages in HSV-1-infected corneas","volume":"200","author":"Jeon","year":"2018","journal-title":"J. Immunol."},{"key":"B71","doi-asserted-by":"publisher","first-page":"6277","DOI":"10.4049\/jimmunol.1300582","article-title":"PD-L1\/B7-H1 regulates the survival but not the function of CD8+ T cells in herpes simplex virus type 1 latently infected trigeminal ganglia","volume":"190","author":"Jeon","year":"2013","journal-title":"J. Immunol."},{"key":"B72","doi-asserted-by":"publisher","first-page":"14733","DOI":"10.1073\/pnas.1009731107","article-title":"Cooperation of Tim-3 and PD-1 in CD8 T-cell exhaustion during chronic viral infection","volume":"107","author":"Jin","year":"2010","journal-title":"Proc. Natl. Acad. Sci. U.S.A"},{"key":"B73","doi-asserted-by":"publisher","first-page":"10527","DOI":"10.1128\/JVI.01546-16","article-title":"Varicella-zoster virus downregulates programmed death ligand 1 and major histocompatibility complex class I in human brain vascular adventitial fibroblasts, perineurial cells, and lung fibroblasts","volume":"90","author":"Jones","year":"2016","journal-title":"J. Virol."},{"key":"B74","doi-asserted-by":"publisher","first-page":"e1007650","DOI":"10.1371\/journal.ppat.1007650","article-title":"Varicella zoster virus productively infects human peripheral blood mononuclear cells to modulate expression of immunoinhibitory proteins and blocking PD-L1 enhances virus-specific CD8+ T cell effector function","volume":"15","author":"Jones","year":"2019","journal-title":"PLoS Pathog."},{"key":"B75","doi-asserted-by":"publisher","first-page":"180","DOI":"10.1016\/j.virol.2014.12.033","article-title":"T cell exhaustion during persistent viral infections","author":"Kahan","year":"2015","journal-title":"Virology"},{"key":"B76","doi-asserted-by":"publisher","first-page":"2","DOI":"10.3390\/v11020156","article-title":"Immune exhaustion: past lessons and new insights from lymphocytic choriomeningitis virus","volume":"11","author":"Kahan","year":"2019","journal-title":"Viruses"},{"key":"B77","doi-asserted-by":"publisher","first-page":"e0205886","DOI":"10.1371\/journal.pone.0205886","article-title":"The immunological function of extracellular vesicles in hepatitis B virus-infected hepatocytes","volume":"13","author":"Kakizaki","year":"2018","journal-title":"PLoS ONE"},{"key":"B78","doi-asserted-by":"publisher","first-page":"1034","DOI":"10.1128\/JVI.01409-09","article-title":"Herpes simplex virus type 1 (HSV-1)-induced apoptosis in human dendritic cells as a result of downregulation of cellular FLICE-inhibitory protein and reduced expression of HSV-1 antiapoptotic latency-associated transcript sequences","volume":"84","author":"Kather","year":"2010","journal-title":"J. Virol."},{"key":"B79","doi-asserted-by":"publisher","first-page":"677","DOI":"10.1146\/annurev.immunol.26.021607.090331","article-title":"PD-1 and its ligands in tolerance and immunity","volume":"26","author":"Keir","year":"2008","journal-title":"Annu. Rev. Immunol."},{"key":"B80","doi-asserted-by":"publisher","first-page":"883","DOI":"10.1084\/jem.20051776","article-title":"Tissue expression of PD-L1 mediates peripheral T cell tolerance","volume":"203","author":"Keir","year":"2006","journal-title":"J. Exp. Med."},{"key":"B81","doi-asserted-by":"publisher","first-page":"593","DOI":"10.1016\/S1074-7613(03)00112-2","article-title":"Herpes simplex virus-specific memory CD8+ T cells are selectively activated and retained in latently infected sensory ganglia","volume":"18","author":"Khanna","year":"2003","journal-title":"Immunity"},{"key":"B82","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1128\/JVI.00207-19","article-title":"By binding CD80 and CD86, the vaccinia virus' M2 protein blocks their interactions with both CD28 and CTLA4 and potentiates CD80's binding to PD-L1","volume":"2019","author":"Kleinpeter","year":"2019","journal-title":"J. Virol"},{"key":"B83","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1126\/science.aao0184","article-title":"Immunology taught by rats","volume":"357","author":"Klenerman","year":"2017","journal-title":"Science"},{"key":"B84","doi-asserted-by":"publisher","first-page":"846","DOI":"10.1172\/JCI21767","article-title":"T cell-mediated vascular dysfunction of human allografts results from IFN-gamma dysregulation of NO synthase","volume":"114","author":"Koh","year":"2004","journal-title":"J. Clin. Invest."},{"key":"B85","doi-asserted-by":"publisher","first-page":"637","DOI":"10.1084\/jem.20050821","article-title":"Type I interferons act directly on CD8 T cells to allow clonal expansion and memory formation in response to viral infection","volume":"202","author":"Kolumam","year":"2005","journal-title":"J. Exp. Med."},{"key":"B86","doi-asserted-by":"publisher","first-page":"2921","DOI":"10.1016\/j.celrep.2017.08.078","article-title":"Human tissue-resident memory T cells are defined by core transcriptional and functional signatures in lymphoid and mucosal sites","volume":"20","author":"Kumar","year":"2017","journal-title":"Cell Rep"},{"key":"B87","doi-asserted-by":"publisher","first-page":"7506","DOI":"10.4049\/jimmunol.180.11.7506","article-title":"Detrimental contribution of the immuno-inhibitor B7-H1 to rabies virus encephalitis","volume":"180","author":"Lafon","year":"2008","journal-title":"J. Immunol."},{"key":"B88","doi-asserted-by":"publisher","first-page":"6667","DOI":"10.1158\/0008-5472.CAN-17-0740","article-title":"IL10 Release upon PD-1 blockade sustains immunosuppression in ovarian cancer","volume":"77","author":"Lamichhane","year":"2017","journal-title":"Cancer Res"},{"key":"B89","doi-asserted-by":"publisher","first-page":"907","DOI":"10.1016\/j.immuni.2019.03.025","article-title":"Shared and distinct functions of type I and type III interferons","volume":"50","author":"Lazear","year":"2019","journal-title":"Immunity"},{"key":"B90","doi-asserted-by":"publisher","first-page":"455","DOI":"10.3389\/fimmu.2013.00455","article-title":"Inhibitory receptor expression depends more dominantly on differentiation and activation than \u201cexhaustion\u201d of human CD8 T cells","volume":"4","author":"Legat","year":"2013","journal-title":"Front. Immunol."},{"key":"B91","doi-asserted-by":"publisher","first-page":"10252","DOI":"10.1128\/JVI.05548-11","article-title":"Longitudinal analysis of the human T cell response during acute hantavirus infection","volume":"85","author":"Lindgren","year":"2011","journal-title":"J. Virol."},{"key":"B92","doi-asserted-by":"publisher","first-page":"1459","DOI":"10.1084\/jem.191.9.1459","article-title":"CD8(+) T cells can block herpes simplex virus type 1 (HSV-1) reactivation from latency in sensory neurons","volume":"191","author":"Liu","year":"2000","journal-title":"J. Exp. Med."},{"key":"B93","doi-asserted-by":"publisher","first-page":"13062","DOI":"10.1128\/JVI.78.23.13062-13071.2004","article-title":"Identification of a mouse cytomegalovirus gene selectively targeting CD86 expression on antigen-presenting cells","volume":"78","author":"Loewendorf","year":"2004","journal-title":"J. Virol."},{"key":"B94","doi-asserted-by":"publisher","first-page":"5336","DOI":"10.1073\/pnas.0931259100","article-title":"PD-L1 and PD-L2 are differentially regulated by Th1 and Th2 cells","volume":"100","author":"Loke","year":"2003","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B95","doi-asserted-by":"publisher","first-page":"2173","DOI":"10.4049\/jimmunol.1102719","article-title":"Maintenance of T cell function in the face of chronic antigen stimulation and repeated reactivation for a latent virus infection","volume":"188","author":"Mackay","year":"2012","journal-title":"J. Immunol."},{"key":"B96","doi-asserted-by":"publisher","first-page":"2714","DOI":"10.4049\/jimmunol.178.5.2714","article-title":"PD-1:PD-L1 interactions contribute to the functional suppression of virus-specific CD8+ T lymphocytes in the liver","volume":"178","author":"Maier","year":"2007","journal-title":"J. Immunol."},{"key":"B97","doi-asserted-by":"publisher","first-page":"1129","DOI":"10.1016\/j.immuni.2017.11.021","article-title":"Transcription factor IRF4 promotes CD8(+) T cell exhaustion and limits the development of memory-like T cells during chronic infection","volume":"47","author":"Man","year":"2017","journal-title":"Immunity"},{"key":"B98","doi-asserted-by":"publisher","first-page":"553","DOI":"10.1084\/jem.20090858","article-title":"Dynamic T cell migration program provides resident memory within intestinal epithelium","volume":"207","author":"Masopust","year":"2010","journal-title":"J. Exp. Med."},{"key":"B99","doi-asserted-by":"crossref","first-page":"8056","DOI":"10.1128\/JVI.68.12.8056-8063.1994","article-title":"CD4+ T cells are required to sustain CD8+ cytotoxic T-cell responses during chronic viral infection","volume":"68","author":"Matloubian","year":"1994","journal-title":"J. Virol."},{"key":"B100","doi-asserted-by":"crossref","first-page":"1043","DOI":"10.1084\/jem.172.4.1043","article-title":"Genetic basis of viral persistence: single amino acid change in the viral glycoprotein affects ability of lymphocytic choriomeningitis virus to persist in adult mice","volume":"172","author":"Matloubian","year":"1990","journal-title":"J. Exp. Med."},{"key":"B101","doi-asserted-by":"publisher","first-page":"4719","DOI":"10.1073\/pnas.1502619112","article-title":"Human Ebola virus infection results in substantial immune activation","volume":"112","author":"McElroy","year":"2015","journal-title":"Proc. Natl. Acad. Sci. U.S.A"},{"key":"B102","doi-asserted-by":"publisher","first-page":"642","DOI":"10.1164\/rccm.201504-0782OC","article-title":"Dysregulation of antiviral function of CD8(+) T cells in the chronic obstructive pulmonary disease lung. Role of the PD-1-PD-L1 Axis","volume":"193","author":"McKendry","year":"2016","journal-title":"Am. J. Respir. Crit. Care Med."},{"key":"B103","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1016\/j.coi.2016.09.005","article-title":"T cell exhaustion and immune-mediated disease-the potential for therapeutic exhaustion","volume":"43","author":"McKinney","year":"2016","journal-title":"Curr. Opin. Immunol."},{"key":"B104","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1038\/s41590-018-0045-y","article-title":"Metabolic exhaustion in infection, cancer and autoimmunity","volume":"19","author":"McKinney","year":"2018","journal-title":"Nat. Immunol."},{"key":"B105","doi-asserted-by":"publisher","first-page":"12916","DOI":"10.1128\/JVI.02423-13","article-title":"Local blockade of epithelial PDL-1 in the airways enhances T cell function and viral clearance during influenza virus infection","volume":"87","author":"McNally","year":"2013","journal-title":"J. Virol."},{"key":"B106","doi-asserted-by":"publisher","first-page":"655","DOI":"10.1097\/QAD.0b013e3282f4de23","article-title":"Upregulation of PD-L1 on monocytes and dendritic cells by HIV-1 derived TLR ligands","volume":"22","author":"Meier","year":"2008","journal-title":"AIDS"},{"key":"B107","doi-asserted-by":"publisher","first-page":"1372","DOI":"10.3389\/fimmu.2017.01372","article-title":"Transcriptome analysis of circulating immune cell subsets highlight the role of monocytes in zaire ebola virus makona pathogenesis","volume":"8","author":"Menicucci","year":"2017","journal-title":"Front. Immunol."},{"key":"B108","doi-asserted-by":"publisher","first-page":"758","DOI":"10.1038\/362758a0","article-title":"Virus persistence in acutely infected immunocompetent mice by exhaustion of antiviral cytotoxic effector T cells","volume":"362","author":"Moskophidis","year":"1993","journal-title":"Nature"},{"key":"B109","doi-asserted-by":"publisher","first-page":"2508","DOI":"10.1172\/JCI40040","article-title":"PD-L1 has distinct functions in hematopoietic and nonhematopoietic cells in regulating T cell responses during chronic infection in mice","volume":"120","author":"Mueller","year":"2010","journal-title":"J. Clin. Invest."},{"key":"B110","doi-asserted-by":"publisher","first-page":"520","DOI":"10.1016\/j.jhep.2006.05.007","article-title":"PD-L1 is induced in hepatocytes by viral infection and by interferon-alpha and -gamma and mediates T cell apoptosis","volume":"45","author":"Muhlbauer","year":"2006","journal-title":"J. Hepatol."},{"key":"B111","doi-asserted-by":"publisher","first-page":"3488","DOI":"10.1002\/(SICI)1521-4141(199811)28:11<3488::AID-IMMU3488>3.0.CO;2-Y","article-title":"Paralysis of B7 co-stimulation through the effect of viral IL-10 on T cells as a mechanism of local tolerance induction","volume":"28","author":"Muller","year":"1998","journal-title":"Eur. J. Immunol."},{"key":"B112","doi-asserted-by":"publisher","first-page":"941","DOI":"10.1002\/eji.200324509","article-title":"Frontline: Induction of apoptosis and modulation of c-FLIPL and p53 in immature dendritic cells infected with herpes simplex virus","volume":"34","author":"Muller","year":"2004","journal-title":"Eur. J. Immunol."},{"key":"B113","doi-asserted-by":"publisher","first-page":"18","DOI":"10.1172\/jci.insight.122940","article-title":"Combination anti-PD-1 and antiretroviral therapy provides therapeutic benefit against SIV","volume":"3","author":"Mylvaganam","year":"2018","journal-title":"JCI Insight"},{"key":"B114","doi-asserted-by":"publisher","first-page":"652","DOI":"10.1016\/j.chom.2013.05.014","article-title":"Networking at the level of host immunity: immune cell interactions during persistent viral infections","volume":"13","author":"Ng","year":"2013","journal-title":"Cell Host Microbe"},{"key":"B115","doi-asserted-by":"publisher","first-page":"1028","DOI":"10.1189\/jlb.0208105","article-title":"Viral replication and paracrine effects result in distinct, functional responses of dendritic cells following infection with dengue 2 virus","volume":"84","author":"Nightingale","year":"2008","journal-title":"J. Leukoc. Biol."},{"key":"B116","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1016\/S1074-7613(00)80089-8","article-title":"Development of lupus-like autoimmune diseases by disruption of the PD-1 gene encoding an ITIM motif-carrying immunoreceptor","volume":"11","author":"Nishimura","year":"1999","journal-title":"Immunity"},{"key":"B117","doi-asserted-by":"publisher","first-page":"319","DOI":"10.1126\/science.291.5502.319","article-title":"Autoimmune dilated cardiomyopathy in PD-1 receptor-deficient mice","volume":"291","author":"Nishimura","year":"2001","journal-title":"Science"},{"key":"B118","doi-asserted-by":"publisher","first-page":"1125","DOI":"10.1084\/jem.20142237","article-title":"Genetic absence of PD-1 promotes accumulation of terminally differentiated exhausted CD8+ T cells","volume":"212","author":"Odorizzi","year":"2015","journal-title":"J. Exp. Med."},{"key":"B119","doi-asserted-by":"publisher","first-page":"360","DOI":"10.1016\/j.virusres.2011.07.011","article-title":"Type-III interferon, not type-I, is the predominant interferon induced by respiratory viruses in nasal epithelial cells","volume":"160","author":"Okabayashi","year":"2011","journal-title":"Virus Res"},{"key":"B120","doi-asserted-by":"publisher","first-page":"1212","DOI":"10.1038\/ni.2762","article-title":"A rheostat for immune responses: the unique properties of PD-1 and their advantages for clinical application","volume":"14","author":"Okazaki","year":"2013","journal-title":"Nat. Immunol"},{"key":"B121","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1016\/j.it.2006.02.001","article-title":"The PD-1-PD-L pathway in immunological tolerance","volume":"27","author":"Okazaki","year":"2006","journal-title":"Trends Immunol"},{"key":"B122","doi-asserted-by":"publisher","first-page":"337","DOI":"10.1038\/d41586-017-08280-8","article-title":"The origins of memory T cells","volume":"552","author":"Omilusik","year":"2017","journal-title":"Nature"},{"key":"B123","doi-asserted-by":"publisher","first-page":"e1005776","DOI":"10.1371\/journal.ppat.1005776","article-title":"Viral manipulation of host inhibitory receptor signaling for immune evasion","volume":"12","author":"Ong","year":"2016","journal-title":"PLoS Pathog"},{"key":"B124","doi-asserted-by":"publisher","first-page":"2212","DOI":"10.1016\/j.vaccine.2011.12.116","article-title":"Global epidemiology of hepatitis B virus infection: new estimates of age-specific HBsAg seroprevalence and endemicity","volume":"30","author":"Ott","year":"2012","journal-title":"Vaccine"},{"key":"B125","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1099\/vir.0.058966-0","article-title":"IL-10 encoded by viruses: a remarkable example of independent acquisition of a cellular gene by viruses and its subsequent evolution in the viral genome","volume":"95","author":"Ouyang","year":"2014","journal-title":"J. Gen. Virol."},{"key":"B126","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1146\/annurev-immunol-031210-101312","article-title":"Regulation and functions of the IL-10 family of cytokines in inflammation and disease","volume":"29","author":"Ouyang","year":"2011","journal-title":"Annu. Rev. Immunol."},{"key":"B127","doi-asserted-by":"publisher","first-page":"411","DOI":"10.1146\/annurev-pathol-020712-164041","article-title":"Pathogenesis of the viral hemorrhagic fevers","volume":"8","author":"Paessler","year":"2013","journal-title":"Annu. Rev. Pathol."},{"key":"B128","doi-asserted-by":"publisher","first-page":"1220","DOI":"10.1126\/science.1229620","article-title":"Progenitor and terminal subsets of CD8+ T cells cooperate to contain chronic viral infection","volume":"338","author":"Paley","year":"2012","journal-title":"Science"},{"key":"B129","doi-asserted-by":"publisher","first-page":"1567","DOI":"10.1084\/jem.20162115","article-title":"IL-2(high) tissue-resident T cells in the human liver: sentinels for hepatotropic infection","volume":"214","author":"Pallett","year":"2017","journal-title":"J. Exp. Med."},{"key":"B130","doi-asserted-by":"publisher","first-page":"1160","DOI":"10.1126\/science.aaf2807","article-title":"Epigenetic stability of exhausted T cells limits durability of reinvigoration by PD-1 blockade","volume":"354","author":"Pauken","year":"2016","journal-title":"Science"},{"key":"B131","doi-asserted-by":"publisher","first-page":"6107","DOI":"10.1172\/JCI96107","article-title":"PD-1+CD8+ T cells are clonally expanding effectors in human chronic inflammation","volume":"2018","author":"Petrelli","year":"2018","journal-title":"J. Clin. Invest"},{"key":"B132","doi-asserted-by":"publisher","first-page":"2281","DOI":"10.1084\/jem.20061496","article-title":"PD-1 is a regulator of virus-specific CD8+ T cell survival in HIV infection","volume":"203","author":"Petrovas","year":"2006","journal-title":"J. Exp. Med."},{"key":"B133","doi-asserted-by":"publisher","first-page":"6672","DOI":"10.1128\/JVI.00825-14","article-title":"HIV-1 Tat protein induces PD-L1 (B7-H1) expression on dendritic cells through tumor necrosis factor alpha- and toll-like receptor 4-mediated mechanisms","volume":"88","author":"Planes","year":"2014","journal-title":"J. Virol."},{"key":"B134","doi-asserted-by":"publisher","first-page":"280","DOI":"10.1038\/ni1165","article-title":"Resting dendritic cells induce peripheral CD8+ T cell tolerance through PD-1 and CTLA-4","volume":"6","author":"Probst","year":"2005","journal-title":"Nat. Immunol."},{"key":"B135","doi-asserted-by":"publisher","first-page":"3634","DOI":"10.4049\/jimmunol.0900974","article-title":"TLR3-stimulated dendritic cells up-regulate B7-H1 expression and influence the magnitude of CD8 T cell responses to tumor vaccination","volume":"183","author":"Pulko","year":"2009","journal-title":"J. Immunol"},{"key":"B136","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1023\/A:1008184330127","article-title":"Herpesvirus homologues of cellular genes","volume":"21","author":"Raftery","year":"2000","journal-title":"Virus Genes"},{"key":"B137","doi-asserted-by":"publisher","first-page":"2560","DOI":"10.3389\/fimmu.2018.02560","article-title":"Hantavirus-driven PD-L1\/PD-L2 upregulation: an imperfect viral immune evasion mechanism","volume":"9","author":"Raftery","year":"2018","journal-title":"Front. Immunol."},{"key":"B138","doi-asserted-by":"crossref","first-page":"1103","DOI":"10.1084\/jem.190.8.1103","article-title":"Herpes simplex virus type 1 infection of activated cytotoxic T cells: induction of fratricide as a mechanism of viral immune evasion","volume":"190","author":"Raftery","year":"1999","journal-title":"J. Exp. Med."},{"key":"B139","doi-asserted-by":"publisher","first-page":"997","DOI":"10.1016\/S1074-7613(01)00239-4","article-title":"Targeting the function of mature dendritic cells by human cytomegalovirus: a multilayered viral defense strategy","volume":"15","author":"Raftery","year":"2001","journal-title":"Immunity"},{"key":"B140","doi-asserted-by":"publisher","first-page":"3383","DOI":"10.4049\/jimmunol.173.5.3383","article-title":"Shaping phenotype, function, and survival of dendritic cells by cytomegalovirus-encoded IL-10","volume":"173","author":"Raftery","year":"2004","journal-title":"J. Immunol."},{"key":"B141","doi-asserted-by":"publisher","first-page":"507","DOI":"10.1189\/jlb.0610327","article-title":"Expression of PD-L1 and PD-L2 on human macrophages is up-regulated by HIV-1 and differentially modulated by IL-10","volume":"89","author":"Rodriguez-Garcia","year":"2011","journal-title":"J. Leukoc. Biol."},{"key":"B142","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1016\/j.celrep.2018.03.026","article-title":"Shp-2 is dispensable for establishing T cell exhaustion and for PD-1 signaling in vivo","volume":"23","author":"Rota","year":"2018","journal-title":"Cell Rep"},{"key":"B143","doi-asserted-by":"publisher","first-page":"100","DOI":"10.1038\/nature17949","article-title":"Unique human immune signature of Ebola virus disease in Guinea","volume":"533","author":"Ruibal","year":"2016","journal-title":"Nature"},{"key":"B144","doi-asserted-by":"publisher","first-page":"1636","DOI":"10.1128\/JVI.02851-13","article-title":"Highly pathological influenza A virus infection is associated with augmented expression of PD-1 by functionally compromised virus-specific CD8+ T cells","volume":"88","author":"Rutigliano","year":"2014","journal-title":"J. Virol."},{"key":"B145","doi-asserted-by":"publisher","first-page":"2569","DOI":"10.3389\/fimmu.2018.02569","article-title":"T-cell exhaustion in chronic infections: reversing the state of exhaustion and reinvigorating optimal protective immune responses","volume":"9","author":"Saeidi","year":"2018","journal-title":"Front. Immunol."},{"key":"B146","doi-asserted-by":"publisher","first-page":"708","DOI":"10.1038\/44385","article-title":"Two subsets of memory T lymphocytes with distinct homing potentials and effector functions","volume":"401","author":"Sallusto","year":"1999","journal-title":"Nature"},{"key":"B147","doi-asserted-by":"publisher","first-page":"313","DOI":"10.1016\/j.cell.2018.09.035","article-title":"A paradigm shift in cancer immunotherapy: from enhancement to normalization","volume":"175","author":"Sanmamed","year":"2018","journal-title":"Cell"},{"key":"B148","doi-asserted-by":"publisher","first-page":"98","DOI":"10.1126\/science.1254536","article-title":"Resident memory CD8 T cells trigger protective innate and adaptive immune responses","volume":"346","author":"Schenkel","year":"2014","journal-title":"Science"},{"key":"B149","doi-asserted-by":"publisher","first-page":"509","DOI":"10.1038\/ni.2568","article-title":"Sensing and alarm function of resident memory CD8(+) T cells","volume":"14","author":"Schenkel","year":"2013","journal-title":"Nat. Immunol"},{"key":"B150","doi-asserted-by":"publisher","first-page":"762","DOI":"10.1007\/s11262-017-1460-0","article-title":"Herpesviral capture of immunomodulatory host genes","volume":"53","author":"Schonrich","year":"2017","journal-title":"Virus Genes"},{"key":"B151","doi-asserted-by":"publisher","first-page":"163","DOI":"10.1111\/j.1600-065X.2008.00694.x","article-title":"Hantavirus-induced immunity in rodent reservoirs and humans","volume":"225","author":"Schonrich","year":"2008","journal-title":"Immunol. Rev."},{"key":"B152","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1016\/j.coi.2016.02.007","article-title":"Recent advances in viral evasion of the MHC Class I processing pathway","volume":"40","author":"Schuren","year":"2016","journal-title":"Curr. Opin. Immunol."},{"key":"B153","doi-asserted-by":"publisher","first-page":"1243","DOI":"10.1016\/j.celrep.2016.06.078","article-title":"Distinct metabolic requirements of exhausted and functional virus-specific CD8 T cells in the same host","volume":"16","author":"Schurich","year":"2016","journal-title":"Cell Rep"},{"key":"B154","doi-asserted-by":"publisher","first-page":"1546","DOI":"10.1016\/S0140-6736(15)61412-X","article-title":"Estimations of worldwide prevalence of chronic hepatitis B virus infection: a systematic review of data published between 1965 and 2013","volume":"386","author":"Schweitzer","year":"2015","journal-title":"Lancet"},{"key":"B155","doi-asserted-by":"publisher","first-page":"3637","DOI":"10.4049\/jimmunol.170.7.3637","article-title":"B7-H1 (programmed death-1 ligand) on dendritic cells is involved in the induction and maintenance of T cell anergy","volume":"170","author":"Selenko-Gebauer","year":"2003","journal-title":"J. Immunol."},{"key":"B156","doi-asserted-by":"publisher","first-page":"1165","DOI":"10.1126\/science.aae0491","article-title":"The epigenetic landscape of T cell exhaustion","volume":"354","author":"Sen","year":"2016","journal-title":"Science"},{"key":"B157","doi-asserted-by":"publisher","first-page":"e2446","DOI":"10.1038\/cddis.2016.350","article-title":"CD169(+) macrophages regulate PD-L1 expression via type I interferon and thereby prevent severe immunopathology after LCMV infection","volume":"7","author":"Shaabani","year":"2016","journal-title":"Cell Death Dis"},{"key":"B158","doi-asserted-by":"publisher","first-page":"153","DOI":"10.1038\/nri.2017.108","article-title":"The diverse functions of the PD1 inhibitory pathway","volume":"18","author":"Sharpe","year":"2018","journal-title":"Nat. Rev. Immunol"},{"key":"B159","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1038\/ni1443","article-title":"The function of programmed cell death 1 and its ligands in regulating autoimmunity and infection","volume":"8","author":"Sharpe","year":"2007","journal-title":"Nat. Immunol."},{"key":"B160","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1016\/j.coviro.2017.11.001","article-title":"Formation and function of tissue-resident memory T cells during viral infection","volume":"28","author":"Shin","year":"2018","journal-title":"Curr. Opin. Virol."},{"key":"B161","doi-asserted-by":"publisher","first-page":"941","DOI":"10.1084\/jem.20061937","article-title":"Viral antigen and extensive division maintain virus-specific CD8 T cells during chronic infection","volume":"204","author":"Shin","year":"2007","journal-title":"J. Exp. Med."},{"key":"B162","doi-asserted-by":"publisher","first-page":"953","DOI":"10.1038\/icb.2017.62","article-title":"Maintenance of PD-1 on brain-resident memory CD8 T cells is antigen independent","volume":"95","author":"Shwetank","year":"2017","journal-title":"Immunol. Cell Biol."},{"key":"B163","doi-asserted-by":"crossref","first-page":"1835","DOI":"10.1126\/science.284.5421.1835","article-title":"The nature of the principal type 1 interferon-producing cells in human blood","volume":"284","author":"Siegal","year":"1999","journal-title":"Science"},{"key":"B164","doi-asserted-by":"publisher","first-page":"4593","DOI":"10.1038\/s41467-018-07053-9","article-title":"Tissue-resident memory T cells populate the human brain","volume":"9","author":"Smolders","year":"2018","journal-title":"Nat. Commun."},{"key":"B165","doi-asserted-by":"publisher","first-page":"768","DOI":"10.1038\/nri3740","article-title":"T cell differentiation in chronic infection and cancer: functional adaptation or exhaustion?","volume":"14","author":"Speiser","year":"2014","journal-title":"Nat. Rev. Immunol."},{"key":"B166","doi-asserted-by":"publisher","first-page":"1081","DOI":"10.1016\/S0140-6736(16)30579-7","article-title":"The global burden of viral hepatitis from 1990 to 2013: findings from the Global Burden of Disease Study 2013","volume":"388","author":"Stanaway","year":"2016","journal-title":"Lancet"},{"key":"B167","doi-asserted-by":"publisher","first-page":"404","DOI":"10.1086\/499275","article-title":"Expression of programmed death-1 ligand (PD-L) 1, PD-L2, B7-H3, and inducible costimulator ligand on human respiratory tract epithelial cells and regulation by respiratory syncytial virus and type 1 and 2 cytokines","volume":"193","author":"Stanciu","year":"2006","journal-title":"J. Infect. Dis."},{"key":"B168","doi-asserted-by":"publisher","first-page":"e0121527","DOI":"10.1371\/journal.pone.0121527","article-title":"Viral infection of human lung macrophages increases PDL1 expression via IFNbeta","volume":"10","author":"Staples","year":"2015","journal-title":"PLoS ONE"},{"key":"B169","doi-asserted-by":"publisher","first-page":"802","DOI":"10.1016\/j.immuni.2014.10.013","article-title":"The transcription factor FoxO1 sustains expression of the inhibitory receptor PD-1 and survival of antiviral CD8(+) T cells during chronic infection","volume":"41","author":"Staron","year":"2014","journal-title":"Immunity"},{"key":"B170","doi-asserted-by":"publisher","first-page":"2142","DOI":"10.1016\/j.celrep.2018.04.038","article-title":"Long-term persistence of exhausted CD8 T cells in chronic infection is regulated by microRNA-155","volume":"23","author":"Stelekati","year":"2018","journal-title":"Cell Rep"},{"key":"B171","doi-asserted-by":"publisher","first-page":"7062","DOI":"10.1126\/science.aav7062","article-title":"Restriction of PD-1 function by cis-PD-L1\/CD80 interactions is required for optimal T cell responses","volume":"2019","author":"Sugiura","year":"2019","journal-title":"Science"},{"key":"B172","doi-asserted-by":"publisher","first-page":"434","DOI":"10.1016\/j.immuni.2018.03.014","article-title":"Regulation and function of the PD-L1 checkpoint","volume":"48","author":"Sun","year":"2018","journal-title":"Immunity"},{"key":"B173","doi-asserted-by":"publisher","first-page":"1635","DOI":"10.1158\/0008-5472.CAN-14-3016","article-title":"IL10 and PD-1 cooperate to limit the activity of tumor-specific CD8+ T cells","volume":"75","author":"Sun","year":"2015","journal-title":"Cancer Res"},{"key":"B174","doi-asserted-by":"publisher","first-page":"848","DOI":"10.1016\/j.immuni.2008.11.002","article-title":"Homeostasis of naive and memory T cells","volume":"29","author":"Surh","year":"2008","journal-title":"Immunity"},{"key":"B175","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1126\/sciimmunol.aas9673","article-title":"Location, location, location: tissue resident memory T cells in mice and humans","volume":"4","author":"Szabo","year":"2019","journal-title":"Sci. Immunol."},{"key":"B176","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1126\/science.1235214","article-title":"Persistent LCMV infection is controlled by blockade of type I interferon signaling","volume":"340","author":"Teijaro","year":"2013","journal-title":"Science"},{"key":"B177","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1093\/infdis\/jiq020","article-title":"RSV-induced bronchial epithelial cell PD-L1 expression inhibits CD8+ T cell nonspecific antiviral activity","volume":"203","author":"Telcian","year":"2011","journal-title":"J. Infect. Dis."},{"key":"B178","doi-asserted-by":"publisher","first-page":"2772","DOI":"10.4049\/jimmunol.1003208","article-title":"IFN-alpha directly promotes programmed cell death-1 transcription and limits the duration of T cell-mediated immunity","volume":"186","author":"Terawaki","year":"2011","journal-title":"J. Immunol"},{"key":"B179","doi-asserted-by":"publisher","first-page":"2734","DOI":"10.3389\/fimmu.2018.02734","article-title":"Signatures of T and B cell development, functional responses and PD-1 upregulation after HCMV latent infections and reactivations in Nod.Rag.gamma mice humanized with cord blood CD34(+) cells","volume":"9","author":"Theobald","year":"2018","journal-title":"Front. Immunol."},{"key":"B180","doi-asserted-by":"publisher","first-page":"1198","DOI":"10.1038\/nm1482","article-title":"Upregulation of PD-1 expression on HIV-specific CD8+ T cells leads to reversible immune dysfunction","volume":"12","author":"Trautmann","year":"2006","journal-title":"Nat. Med."},{"key":"B181","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1016\/j.bbrc.2005.02.161","article-title":"Expression of B7-H1 and B7-DC on the airway epithelium is enhanced by double-stranded RNA","volume":"330","author":"Tsuda","year":"2005","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"B182","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1053\/j.gastro.2009.09.009","article-title":"Hepatitis C virus core protein subverts the antiviral activities of human Kupffer cells","volume":"138","author":"Tu","year":"2010","journal-title":"Gastroenterology"},{"key":"B183","doi-asserted-by":"publisher","first-page":"1104","DOI":"10.1126\/science.aal3204","article-title":"Can T cells be too exhausted to fight back?","volume":"354","author":"Turner","year":"2016","journal-title":"Science"},{"key":"B184","doi-asserted-by":"publisher","first-page":"1819","DOI":"10.1084\/jem.20150598","article-title":"High antigen levels induce an exhausted phenotype in a chronic infectiwithout impairing T cell expansion and survival","volume":"213","author":"Utzschneider","year":"2016","journal-title":"J. Exp. Med"},{"key":"B185","doi-asserted-by":"publisher","first-page":"603","DOI":"10.1038\/ni.2606","article-title":"T cells maintain an exhausted phenotype after antigen withdrawal and population reexpansion","volume":"14","author":"Utzschneider","year":"2013","journal-title":"Nat. Immunol."},{"key":"B186","doi-asserted-by":"publisher","first-page":"989673","DOI":"10.1155\/2013\/989673","article-title":"PD-L1 expression induced by the 2009 pandemic influenza A(H1N1) virus impairs the human T cell response","volume":"2013","author":"Valero-Pacheco","year":"2013","journal-title":"Clin. Dev. Immunol."},{"key":"B187","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1111\/imr.12085","article-title":"Role of regulatory T cells during virus infection","volume":"255","author":"Veiga-Parga","year":"2013","journal-title":"Immunol. Rev."},{"key":"B188","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1186\/s12977-015-0144-x","article-title":"Role of PD-1 co-inhibitory pathway in HIV infection and potential therapeutic options","volume":"12","author":"Velu","year":"2015","journal-title":"Retrovirology"},{"key":"B189","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1038\/nature07662","article-title":"Enhancing SIV-specific immunity in vivo by PD-1 blockade","volume":"458","author":"Velu","year":"2009","journal-title":"Nature"},{"key":"B190","doi-asserted-by":"publisher","first-page":"3496","DOI":"10.1073\/pnas.0610847104","article-title":"Selective retention of herpes simplex virus-specific T cells in latently infected human trigeminal ganglia","volume":"104","author":"Verjans","year":"2007","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B191","doi-asserted-by":"publisher","first-page":"30","DOI":"10.1016\/j.cell.2009.06.036","article-title":"Redefining chronic viral infection","volume":"138","author":"Virgin","year":"2009","journal-title":"Cell"},{"key":"B192","doi-asserted-by":"publisher","first-page":"802","DOI":"10.1038\/ni.3212","article-title":"Guarding the frontiers: the biology of type III interferons","volume":"16","author":"Wack","year":"2015","journal-title":"Nat. Immunol."},{"key":"B193","doi-asserted-by":"publisher","first-page":"198","DOI":"10.1126\/science.1151869","article-title":"Dendritic cell-induced memory T cell activation in nonlymphoid tissues","volume":"319","author":"Wakim","year":"2008","journal-title":"Science"},{"key":"B194","doi-asserted-by":"publisher","first-page":"11823","DOI":"10.1073\/pnas.0505497102","article-title":"Establishment of NOD-Pdcd1-\/- mice as an efficient animal model of type I diabetes","volume":"102","author":"Wang","year":"2005","journal-title":"Proc. Natl. Acad. Sci. U.S.A"},{"key":"B195","doi-asserted-by":"publisher","first-page":"3226","DOI":"10.1002\/eji.200838285","article-title":"B7-H1 up-regulation impairs myeloid DC and correlates with disease progression in chronic HIV-1 infection","volume":"38","author":"Wang","year":"2008","journal-title":"Eur. J. Immunol."},{"key":"B196","doi-asserted-by":"publisher","first-page":"E2480","DOI":"10.1073\/pnas.1305394110","article-title":"Strength of PD-1 signaling differentially affects T-cell effector functions","volume":"110","author":"Wei","year":"2013","journal-title":"Proc. Natl. Acad. Sci. U.S.A"},{"key":"B197","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1002\/9780471729259.mc15a01s8","article-title":"Lymphocytic choriomeningitis virus (LCMV): propagation, quantitation, and storage","author":"Welsh","year":"2008","journal-title":"Curr. Protoc. Microbiol."},{"key":"B198","doi-asserted-by":"publisher","first-page":"5535","DOI":"10.1128\/JVI.78.11.5535-5545.2004","article-title":"Memory CD8 T-cell differentiation during viral infection","volume":"78","author":"Wherry","year":"2004","journal-title":"J. Virol."},{"key":"B199","doi-asserted-by":"publisher","first-page":"16004","DOI":"10.1073\/pnas.0407192101","article-title":"Antigen-independent memory CD8 T cells do not develop during chronic viral infection","volume":"101","author":"Wherry","year":"2004","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B200","doi-asserted-by":"publisher","first-page":"4911","DOI":"10.1128\/jvi.77.8.4911-4927.2003","article-title":"Viral persistence alters CD8 T-cell immunodominance and tissue distribution and results in distinct stages of functional impairment","volume":"77","author":"Wherry","year":"2003","journal-title":"J. Virol."},{"key":"B201","doi-asserted-by":"publisher","first-page":"670","DOI":"10.1016\/j.immuni.2007.09.006","article-title":"Molecular signature of CD8+ T cell exhaustion during chronic viral infection","volume":"27","author":"Wherry","year":"2007","journal-title":"Immunity"},{"key":"B202","doi-asserted-by":"publisher","first-page":"486","DOI":"10.1038\/nri3862","article-title":"Molecular and cellular insights into T cell exhaustion","volume":"15","author":"Wherry","year":"2015","journal-title":"Nat. Rev. Immunol."},{"key":"B203","doi-asserted-by":"publisher","first-page":"15050","DOI":"10.1038\/ncomms15050","article-title":"TCF1(+) hepatitis C virus-specific CD8(+) T cells are maintained after cessation of chronic antigen stimulation","volume":"8","author":"Wieland","year":"2017","journal-title":"Nat. Commun."},{"key":"B204","doi-asserted-by":"publisher","first-page":"202","DOI":"10.1126\/science.1235208","article-title":"Blockade of chronic type I interferon signaling to control persistent LCMV infection","volume":"340","author":"Wilson","year":"2013","journal-title":"Science"},{"key":"B205","doi-asserted-by":"publisher","first-page":"1452","DOI":"10.1016\/j.cell.2018.11.006","article-title":"Putting the immunologic brakes on cancer","volume":"175","author":"Wolchok","year":"2018","journal-title":"Cell"},{"key":"B206","doi-asserted-by":"publisher","first-page":"2093","DOI":"10.3389\/fimmu.2018.02093","article-title":"CD8(+) resident memory T cells and viral infection","volume":"9","author":"Wu","year":"2018","journal-title":"Front. Immunol."},{"key":"B207","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1038\/nri.2017.112","article-title":"Immune checkpoint blockade in infectious diseases","volume":"18","author":"Wykes","year":"2018","journal-title":"Nat. Rev. Immunol."},{"key":"B208","doi-asserted-by":"publisher","first-page":"943","DOI":"10.1084\/jem.20130790","article-title":"RGMb is a novel binding partner for PD-L2 and its engagement with PD-L2 promotes respiratory tolerance","volume":"211","author":"Xiao","year":"2014","journal-title":"J. Exp. Med."},{"key":"B209","doi-asserted-by":"publisher","first-page":"7340","DOI":"10.4049\/jimmunol.1001642","article-title":"Increased B7-H1 expression on dendritic cells correlates with programmed death 1 expression on T cells in simian immunodeficiency virus-infected macaques and may contribute to T cell dysfunction and disease progression","volume":"185","author":"Xu","year":"2010","journal-title":"J. Immunol."},{"key":"B210","doi-asserted-by":"publisher","first-page":"5538","DOI":"10.4049\/jimmunol.169.10.5538","article-title":"Expression of programmed death 1 ligands by murine T cells and AP","volume":"169","author":"Yamazaki","year":"2002","journal-title":"J. Immunol."},{"key":"B211","doi-asserted-by":"publisher","first-page":"404","DOI":"10.1038\/nature25144","article-title":"Effector CD8 T cells dedifferentiate into long-lived memory cells","volume":"552","author":"Youngblood","year":"2017","journal-title":"Nature"},{"key":"B212","doi-asserted-by":"publisher","first-page":"400","DOI":"10.1016\/j.immuni.2011.06.015","article-title":"Chronic virus infection enforces demethylation of the locus that encodes PD-1 in antigen-specific CD8(+) T cells","volume":"35","author":"Youngblood","year":"2011","journal-title":"Immunity"},{"key":"B213","doi-asserted-by":"crossref","first-page":"2205","DOI":"10.1084\/jem.188.12.2205","article-title":"Viral immune evasion due to persistence of activated T cells without effector function","volume":"188","author":"Zajac","year":"1998","journal-title":"J. Exp. Med."},{"key":"B214","doi-asserted-by":"publisher","first-page":"1159","DOI":"10.1038\/ni1519","article-title":"Immunopathology of highly virulent pathogens: insights from Ebola virus","volume":"8","author":"Zampieri","year":"2007","journal-title":"Nat. Immunol."},{"key":"B215","doi-asserted-by":"publisher","first-page":"1661","DOI":"10.3389\/fimmu.2017.01661","article-title":"Interferon (IFN)-lambda takes the helm: immunomodulatory roles of type III IFNs","volume":"8","author":"Zanoni","year":"2017","journal-title":"Front. Immunol."},{"key":"B216","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1016\/j.coviro.2016.01.002","article-title":"Immune-surveillance through exhausted effector T-cells","volume":"16","author":"Zehn","year":"2016","journal-title":"Curr. Opin. Virol."},{"key":"B217","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1007\/978-3-319-15774-0_10","article-title":"Immune memory and exhaustion: clinically relevant lessons from the LCMV model","volume":"850","author":"Zehn","year":"2015","journal-title":"Adv. Exp. Med. Biol."},{"key":"B218","doi-asserted-by":"publisher","first-page":"e0170516","DOI":"10.1371\/journal.pone.0170516","article-title":"Hepatitis C virus induces MDSCs-like monocytes through TLR2\/PI3K\/AKT\/STAT3 signaling","volume":"12","author":"Zhai","year":"2017","journal-title":"PLoS ONE"},{"key":"B219","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1016\/j.immuni.2011.07.010","article-title":"CD8(+) T cells: foot soldiers of the immune system","volume":"35","author":"Zhang","year":"2011","journal-title":"Immunity"},{"key":"B220","doi-asserted-by":"publisher","first-page":"757","DOI":"10.1084\/jem.20121416","article-title":"PD-1 promotes immune exhaustion by inducing antiviral T cell motility paralysis","volume":"210","author":"Zinselmeyer","year":"2013","journal-title":"J. Exp. Med"}],"container-title":["Frontiers in Cellular and Infection Microbiology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/article\/10.3389\/fcimb.2019.00207\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,1,5]],"date-time":"2021-01-05T18:29:35Z","timestamp":1609871375000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/article\/10.3389\/fcimb.2019.00207\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,6,13]]},"references-count":220,"alternative-id":["10.3389\/fcimb.2019.00207"],"URL":"https:\/\/doi.org\/10.3389\/fcimb.2019.00207","relation":{},"ISSN":["2235-2988"],"issn-type":[{"value":"2235-2988","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,6,13]]},"article-number":"207"}}