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CD4<jats:sup>+<\/jats:sup> T cells are essential to contain infection, but how best to prime them and whether this can reduce viral loads remain unclear. To address these questions, we used ovalbumin (OVA) as a model antigen, delivering it with murine cytomegalovirus (MCMV) to protect mice against OVA\u2010expressing murine herpesvirus\u20104 (MuHV\u20104). Membrane\u2010associated OVA (mOVA) was more effective than soluble OVA, both to prime CD4<jats:sup>+<\/jats:sup> T cells and as an effector target. It was also a better target than an OVA epitope limited to infected cells, suggesting that protective CD4<jats:sup>+<\/jats:sup> T cells recognize infected cell debris rather than infected cells themselves. While MCMV\u2010mOVA protected acutely against MuHV\u20104\u2010mOVA, long\u2010term protection was incomplete, even when OVA\u2010specific CD8<jats:sup>+<\/jats:sup> T cells and B cells were also primed. Thus, even optimized single\u2010target vaccines may poorly reduce long\u2010term \u03b3\u2010herpesvirus infections.<\/jats:p>","DOI":"10.1111\/imcb.12319","type":"journal-article","created":{"date-parts":[[2020,1,30]],"date-time":"2020-01-30T07:13:33Z","timestamp":1580368413000},"page":"332-343","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Membrane association of a model CD4<sup>+<\/sup> T\u2010cell vaccine antigen confers enhanced yet incomplete protection against murid herpesvirus\u20104 infection"],"prefix":"10.1111","volume":"98","author":[{"given":"Joseph","family":"Yunis","sequence":"first","affiliation":[{"name":"School of Chemistry and Molecular Biosciences University of Queensland St Lucia QLD Australia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Alec J","family":"Redwood","sequence":"additional","affiliation":[{"name":"Faculty of Health and Medical Sciences University of Western Australia Perth WA Australia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Gabrielle T","family":"Belz","sequence":"additional","affiliation":[{"name":"Molecular Immunology Walter and Eliza Hall Institute Melbourne VIC Australia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3520-5060","authenticated-orcid":false,"given":"Philip G","family":"Stevenson","sequence":"additional","affiliation":[{"name":"School of Chemistry and Molecular Biosciences University of Queensland St Lucia QLD Australia"},{"name":"Royal Children's Hospital Brisbane QLD Australia"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"311","published-online":{"date-parts":[[2020,2,29]]},"reference":[{"key":"e_1_2_7_2_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.vaccine.2012.09.041"},{"key":"e_1_2_7_3_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.coviro.2013.04.005"},{"key":"e_1_2_7_4_1","doi-asserted-by":"publisher","DOI":"10.1099\/vir.0.013300-0"},{"key":"e_1_2_7_5_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-immunol-072710-081639"},{"key":"e_1_2_7_6_1","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.77.4.2522-2529.2003"},{"key":"e_1_2_7_7_1","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.78.13.6836-6845.2004"},{"key":"e_1_2_7_8_1","doi-asserted-by":"publisher","DOI":"10.1099\/vir.0.023085-0"},{"key":"e_1_2_7_9_1","doi-asserted-by":"publisher","DOI":"10.1186\/1743-422X-9-107"},{"key":"e_1_2_7_10_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.96.16.9281"},{"key":"e_1_2_7_11_1","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.75.17.8283-8288.2001"},{"key":"e_1_2_7_12_1","doi-asserted-by":"publisher","DOI":"10.1099\/0022-1317-80-2-477"},{"key":"e_1_2_7_13_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1006311"},{"key":"e_1_2_7_14_1","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2006-12-063008"},{"key":"e_1_2_7_15_1","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.00787-06"},{"key":"e_1_2_7_16_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.169.4.1199"},{"key":"e_1_2_7_17_1","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.02240-08"},{"key":"e_1_2_7_18_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.169.3.1593"},{"key":"e_1_2_7_19_1","doi-asserted-by":"publisher","DOI":"10.1002\/eji.200324794"},{"key":"e_1_2_7_20_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ejcb.2011.01.007"},{"key":"e_1_2_7_21_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.190.4.567"},{"key":"e_1_2_7_22_1","doi-asserted-by":"publisher","DOI":"10.1034\/j.1600-065X.2002.18510.x"},{"key":"e_1_2_7_23_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0952-7915(98)80227-6"},{"key":"e_1_2_7_24_1","doi-asserted-by":"publisher","DOI":"10.2741\/2096"},{"key":"e_1_2_7_25_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.96.9.5135"},{"key":"e_1_2_7_26_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.virol.2005.05.011"},{"key":"e_1_2_7_27_1","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.66.4.1977-1984.1992"},{"key":"e_1_2_7_28_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1004761"},{"key":"e_1_2_7_29_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1006905"},{"key":"e_1_2_7_30_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.vaccine.2010.03.012"},{"key":"e_1_2_7_31_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0021-9258(19)50633-0"},{"key":"e_1_2_7_32_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.166.10.6099"},{"key":"e_1_2_7_33_1","doi-asserted-by":"publisher","DOI":"10.1002\/eji.200636164"},{"key":"e_1_2_7_34_1","doi-asserted-by":"crossref","first-page":"2067","DOI":"10.4049\/jimmunol.133.4.2067","article-title":"Antigen recognition by H2\u2010restricted T cells. 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