{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,27]],"date-time":"2026-04-27T04:32:12Z","timestamp":1777264332438,"version":"3.51.4"},"reference-count":58,"publisher":"Wiley","issue":"4","license":[{"start":{"date-parts":[[2026,4,6]],"date-time":"2026-04-06T00:00:00Z","timestamp":1775433600000},"content-version":"vor","delay-in-days":5,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"},{"start":{"date-parts":[[2026,4,1]],"date-time":"2026-04-01T00:00:00Z","timestamp":1775001600000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["HLA"],"published-print":{"date-parts":[[2026,4]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    Allele\u2010specific variability in HLA class I gene expression levels has been associated with many diseases. Although expression variation at the allelic level has not been well\u2010characterised for the HLA class II genes, differential expression of these genes, as marked by single nucleotide polymorphisms (SNPs), has been implicated in the outcome of several diseases. Here, we measured cell surface expression levels of distinct HLA\u2010DR, HLA\u2010DQ and HLA\u2010DP allotypes in 175 healthy European American donors using locus\u2010specific antibodies. We identified allotype\u2010specific variation in the intrinsic cell surface expression levels. To further characterise genetic associations with differential protein expression levels of the HLA class II molecules, we performed a genome\u2010wide association study (GWAS). This showed that surface expression levels of HLA\u2010DR, HLA\u2010DQ and HLA\u2010DP associated most significantly with rs28383323 (\n                    <jats:italic>p<\/jats:italic>\n                    \u2009=\u20093.9\u2009\u00d7\u200910\n                    <jats:sup>\u221210<\/jats:sup>\n                    ), rs281860696 (\n                    <jats:italic>p<\/jats:italic>\n                    \u2009=\u20092.2\u2009\u00d7\u200910\n                    <jats:sup>\u22125<\/jats:sup>\n                    ) and rs3128928 (\n                    <jats:italic>p<\/jats:italic>\n                    \u2009=\u20091.3\u2009\u00d7\u200910\n                    <jats:sup>\u22128<\/jats:sup>\n                    ), respectively. Two SNPs, rs3128927 and rs9277534, were previously reported to associate with HLA\u2010DP protein expression and hepatitis B virus (HBV) recovery\/persistence. We show that rs3128928, which resides in the\n                    <jats:italic>COL11A2P1<\/jats:italic>\n                    pseudogene, associates most significantly with HLA\u2010DP protein expression, whereas the neighbouring rs3128927 associates most significantly with HBV recovery\/persistence. Thus, protein expression levels of HLA\u2010DR, DQ and DP appear to be controlled by cis\u2010elements that do not mark specific alleles. This finding may help to distinguish disease associations caused by specific HLA allelic effects vs. protein expression levels.\n                  <\/jats:p>","DOI":"10.1111\/tan.70669","type":"journal-article","created":{"date-parts":[[2026,4,6]],"date-time":"2026-04-06T09:16:09Z","timestamp":1775466969000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["<scp>HLA<\/scp>\n                    Class\n                    <scp>II<\/scp>\n                    Protein Expression Regulation Is Strongly Linked to Cis\u2010Acting\n                    <scp>SNPs<\/scp>"],"prefix":"10.1111","volume":"107","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3767-6210","authenticated-orcid":false,"given":"Nicolas","family":"Vince","sequence":"first","affiliation":[{"name":"Basic Science Program, Frederick National Laboratory for Cancer Research in the Laboratory of Integrative Cancer Immunology, National Cancer Institute  Bethesda Maryland USA"},{"name":"Ragon Institute of MGH, MIT and Harvard  Cambridge Massachusetts USA"},{"name":"Nantes Universit\u00e9, INSERM, Center for Research in Transplantation and Translational Immunology  Nantes France"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Veron","family":"Ramsuran","sequence":"additional","affiliation":[{"name":"Basic Science Program, Frederick National Laboratory for Cancer Research in the Laboratory of Integrative Cancer Immunology, National Cancer Institute  Bethesda Maryland USA"},{"name":"Ragon Institute of MGH, MIT and Harvard  Cambridge Massachusetts USA"},{"name":"KwaZulu\u2010Natal Research Innovation and Sequencing Platform (KRISP), School of Laboratory Medicine and Medical Sciences, University of KwaZulu\u2010Natal  Durban South Africa"},{"name":"Centre for the AIDS Programme of Research in South Africa (CAPRISA)  Durban South Africa"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mathias","family":"Viard","sequence":"additional","affiliation":[{"name":"Basic Science Program, Frederick National Laboratory for Cancer Research in the Laboratory of Integrative Cancer Immunology, National Cancer Institute  Bethesda Maryland USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jeremy","family":"Martinson","sequence":"additional","affiliation":[{"name":"University of Pittsburgh Graduate School of Public Health  Pittsburgh Pennsylvania USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Steven","family":"Wolinsky","sequence":"additional","affiliation":[{"name":"Division of Infectious Diseases Northwestern University, The Feinberg School of Medicine  Chicago Illinois USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Shehnaz","family":"Hussain","sequence":"additional","affiliation":[{"name":"Division of Epidemiology University of California Davis, School of Medicine  Davis California USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Eric C.","family":"Seaberg","sequence":"additional","affiliation":[{"name":"Bloomberg School of Public Health, Johns Hopkins University  Baltimore Maryland USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Todd T.","family":"Brown","sequence":"additional","affiliation":[{"name":"Bloomberg School of Public Health, Johns Hopkins University  Baltimore Maryland USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Niamh","family":"Spence","sequence":"additional","affiliation":[{"name":"Basic Science Program, Frederick National Laboratory for Cancer Research in the Laboratory of Integrative Cancer Immunology, National Cancer Institute  Bethesda Maryland USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8851-8319","authenticated-orcid":false,"given":"Chloe L.","family":"Thio","sequence":"additional","affiliation":[{"name":"Johns Hopkins University, School of Medicine  Baltimore Maryland USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"James J.","family":"Goedert","sequence":"additional","affiliation":[{"name":"National Cancer Institute  Washington DC USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"W. 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