{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,27]],"date-time":"2026-08-27T01:19:09Z","timestamp":1787793549375,"version":"build-2784847793"},"reference-count":41,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T00:00:00Z","timestamp":1787702400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["24 K11484"],"award-info":[{"award-number":["24 K11484"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Cellular Immunology"],"published-print":{"date-parts":[[2026,9]]},"DOI":"10.1016\/j.cellimm.2026.105144","type":"journal-article","created":{"date-parts":[[2026,8,8]],"date-time":"2026-08-08T15:01:34Z","timestamp":1786201294000},"page":"105144","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"special_numbering":"C","title":["HVEM expressed in CD4 T cells promotes self-tolerance to an epidermal antigen through Treg expansion and modulation of B cell responses"],"prefix":"10.1016","volume":"427","author":[{"given":"Yasushi","family":"Hara","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Akikazu","family":"Murakami","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryo","family":"Goitsuka","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Naoko","family":"Nakano","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.cellimm.2026.105144_bb0005","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1126\/science.280.5361.243","article-title":"Homeostasis and self-tolerance in the immune system: turning lymphocytes off","volume":"280","author":"Van Parijs","year":"1998","journal-title":"Science"},{"key":"10.1016\/j.cellimm.2026.105144_bb0010","doi-asserted-by":"crossref","first-page":"455","DOI":"10.1016\/S0092-8674(00)80856-9","article-title":"Regulatory T cells: key controllers of immunologic self-tolerance","volume":"101","author":"Sakaguchi","year":"2000","journal-title":"Cell"},{"key":"10.1016\/j.cellimm.2026.105144_bb0015","first-page":"844","article-title":"Specialized role of migratory dendritic cells in peripheral tolerance induction","volume":"123","author":"Idoyaga","year":"2013","journal-title":"J. Clin. Invest."},{"key":"10.1016\/j.cellimm.2026.105144_bb0020","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1016\/j.it.2016.08.012","article-title":"Induced regulatory T cells: their development, stability, and applications","volume":"37","author":"Kanamori","year":"2016","journal-title":"Trends Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0025","doi-asserted-by":"crossref","first-page":"896","DOI":"10.1016\/j.immuni.2015.10.011","article-title":"CD4+ T cell tolerance to tissue-restricted self antigens is mediated by antigen-specific regulatory T cells rather than deletion","volume":"43","author":"Legoux","year":"2015","journal-title":"Immunity"},{"key":"10.1016\/j.cellimm.2026.105144_bb0030","doi-asserted-by":"crossref","first-page":"624","DOI":"10.1038\/s41586-020-2611-3","article-title":"cDC1 prime and are licensed by CD4(+) T cells to induce anti-tumour immunity","volume":"584","author":"Ferris","year":"2020","journal-title":"Nature"},{"key":"10.1016\/j.cellimm.2026.105144_bb0035","doi-asserted-by":"crossref","first-page":"eadc9584","DOI":"10.1126\/sciimmunol.adc9584","article-title":"CCR7 acts as both a sensor and a sink for CCL19 to coordinate collective leukocyte migration","volume":"8","author":"Alanko","year":"2023","journal-title":"Sci Immunol"},{"key":"10.1016\/j.cellimm.2026.105144_bb0040","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1084\/jem.20091964","article-title":"CD207+ CD103+ dermal dendritic cells cross-present keratinocyte-derived antigens irrespective of the presence of Langerhans cells","volume":"207","author":"Henri","year":"2010","journal-title":"J. Exp. Med."},{"key":"10.1016\/j.cellimm.2026.105144_bb0045","doi-asserted-by":"crossref","first-page":"252","DOI":"10.1016\/j.jaut.2008.04.017","article-title":"Role of Th1 and Th17 cells in organ-specific autoimmunity","volume":"31","author":"Dardalhon","year":"2008","journal-title":"J. Autoimmun."},{"key":"10.1016\/j.cellimm.2026.105144_bb0050","doi-asserted-by":"crossref","first-page":"304","DOI":"10.1038\/ni.3331","article-title":"CD4(+) T cell anergy prevents autoimmunity and generates regulatory T cell precursors","volume":"17","author":"Kalekar","year":"2016","journal-title":"Nat. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0055","doi-asserted-by":"crossref","first-page":"719","DOI":"10.1016\/S0092-8674(02)00767-5","article-title":"Transcriptional mechanisms underlying lymphocyte tolerance","volume":"109","author":"Macian","year":"2002","journal-title":"Cell"},{"key":"10.1016\/j.cellimm.2026.105144_bb0060","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1111\/j.1600-065X.2011.01064.x","article-title":"The signaling networks of the herpesvirus entry mediator (TNFRSF14) in immune regulation","volume":"244","author":"Steinberg","year":"2011","journal-title":"Immunol. Rev."},{"key":"10.1016\/j.cellimm.2026.105144_bb0065","doi-asserted-by":"crossref","first-page":"4105","DOI":"10.4049\/jimmunol.164.8.4105","article-title":"LIGHT, a TNF-like molecule, costimulates T cell proliferation and is required for dendritic cell-mediated allogeneic T cell response","volume":"164","author":"Tamada","year":"2000","journal-title":"J. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0070","doi-asserted-by":"crossref","first-page":"13471","DOI":"10.1074\/jbc.272.21.13471","article-title":"ATAR, a novel tumor necrosis factor receptor family member, signals through TRAF2 and TRAF5","volume":"272","author":"Hsu","year":"1997","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.cellimm.2026.105144_bb0075","doi-asserted-by":"crossref","first-page":"1482","DOI":"10.4049\/jimmunol.2200748","article-title":"Lack of herpes virus entry mediator signals in Thymocytes impairs conventional CD8 T cell selection and promotes memory-like CD8 T cell development","volume":"210","author":"Kugii","year":"2023","journal-title":"J. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0080","doi-asserted-by":"crossref","first-page":"671","DOI":"10.1038\/nri1917","article-title":"Balancing co-stimulation and inhibition with BTLA and HVEM","volume":"6","author":"Murphy","year":"2006","journal-title":"Nat. Rev. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0085","doi-asserted-by":"crossref","DOI":"10.1016\/j.immuni.2019.05.022","article-title":"The HVEM-BTLA axis restrains T cell help to germinal center B cells and functions as a cell-extrinsic suppressor in lymphomagenesis","volume":"51","author":"Mintz","year":"2019","journal-title":"Immunity"},{"key":"10.1016\/j.cellimm.2026.105144_bb0090","first-page":"296","article-title":"Targeted somatic mutagenesis in mouse epidermis","volume":"54","author":"Indra","year":"2000","journal-title":"Horm. Res."},{"key":"10.1016\/j.cellimm.2026.105144_bb0095","doi-asserted-by":"crossref","first-page":"633","DOI":"10.1038\/35036595","article-title":"Skin abnormalities generated by temporally controlled RXRalpha mutations in mouse epidermis","volume":"407","author":"Li","year":"2000","journal-title":"Nature"},{"key":"10.1016\/j.cellimm.2026.105144_bb0100","doi-asserted-by":"crossref","first-page":"1539","DOI":"10.1084\/jem.179.5.1539","article-title":"Resting and anergic B cells are defective in CD28-dependent costimulation of naive CD4+ T cells","volume":"179","author":"Ho","year":"1994","journal-title":"J. Exp. Med."},{"key":"10.1016\/j.cellimm.2026.105144_bb0105","doi-asserted-by":"crossref","first-page":"786","DOI":"10.1038\/ni.1745","article-title":"Immune complex relay by subcapsular sinus macrophages and noncognate B cells drives antibody affinity maturation","volume":"10","author":"Phan","year":"2009","journal-title":"Nat. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0110","doi-asserted-by":"crossref","first-page":"1565","DOI":"10.4049\/jimmunol.180.3.1565","article-title":"Th2 lymphoproliferative disorder of LatY136F mutant mice unfolds independently of TCR-MHC engagement and is insensitive to the action of Foxp3+ regulatory T cells","volume":"180","author":"Wang","year":"2008","journal-title":"J. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0115","doi-asserted-by":"crossref","first-page":"765","DOI":"10.1038\/353765a0","article-title":"Elimination from peripheral lymphoid tissues of self-reactive B lymphocytes recognizing membrane-bound antigens","volume":"353","author":"Hartley","year":"1991","journal-title":"Nature"},{"key":"10.1016\/j.cellimm.2026.105144_bb0120","doi-asserted-by":"crossref","DOI":"10.1016\/j.chom.2018.07.008","article-title":"LIGHT-HVEM signaling in innate lymphoid cell subsets protects against enteric bacterial infection","volume":"24","author":"Seo","year":"2018","journal-title":"Cell Host Microbe"},{"key":"10.1016\/j.cellimm.2026.105144_bb0125","doi-asserted-by":"crossref","first-page":"777","DOI":"10.1084\/jem.20072513","article-title":"CXCR4 is required for the quiescence of primitive hematopoietic cells","volume":"205","author":"Nie","year":"2008","journal-title":"J. Exp. Med."},{"key":"10.1016\/j.cellimm.2026.105144_bb0130","doi-asserted-by":"crossref","DOI":"10.1016\/j.celrep.2021.108748","article-title":"Dynamic adoption of anergy by antigen-exhausted CD4(+) T cells","volume":"34","author":"Trefzer","year":"2021","journal-title":"Cell Rep."},{"key":"10.1016\/j.cellimm.2026.105144_bb0135","doi-asserted-by":"crossref","first-page":"5225","DOI":"10.4049\/jimmunol.181.8.5225","article-title":"Alterations of the medullaru epithelial compartment in the Aire-deficient thymus: implications for programs of thymic epithelial differentiation","volume":"181","author":"Dooley","year":"2008","journal-title":"J. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0140","doi-asserted-by":"crossref","first-page":"1875","DOI":"10.1084\/jem.20030152","article-title":"Conversion of peripheral CD4+CD25- naive T cells to CD4+CD25+ regulatory T cells by TGF-beta induction of transcription factor Foxp3","volume":"198","author":"Chen","year":"2003","journal-title":"J. Exp. Med."},{"key":"10.1016\/j.cellimm.2026.105144_bb0145","doi-asserted-by":"crossref","first-page":"888","DOI":"10.1038\/ni.2370","article-title":"Deciphering the transcriptional network of the dendritic cell lineage","volume":"13","author":"Miller","year":"2012","journal-title":"Nat. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0150","doi-asserted-by":"crossref","first-page":"987","DOI":"10.1084\/jem.20101773","article-title":"Viral persistence redirects CD4 T cell differentiation toward T follicular helper cells","volume":"208","author":"Fahey","year":"2011","journal-title":"J. Exp. Med."},{"key":"10.1016\/j.cellimm.2026.105144_bb0155","doi-asserted-by":"crossref","first-page":"1509","DOI":"10.4049\/jimmunol.182.3.1509","article-title":"B and T lymphocyte attenuator regulates B cell receptor signaling by targeting Syk and BLNK","volume":"182","author":"Vendel","year":"2009","journal-title":"J. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0160","doi-asserted-by":"crossref","first-page":"935","DOI":"10.1093\/intimm\/9.7.935","article-title":"Immobilized anti-CD3 mAb induces anergy in murine naive and memory CD4+ T cells in vitro","volume":"9","author":"Chai","year":"1997","journal-title":"Int. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0165","doi-asserted-by":"crossref","first-page":"2005","DOI":"10.1084\/jem.186.12.2005","article-title":"Self-reactive B cells are not eliminated or inactivated by autoantigen expressed on thyroid epithelial cells","volume":"186","author":"Akkaraju","year":"1997","journal-title":"J. Exp. Med."},{"key":"10.1016\/j.cellimm.2026.105144_bb0170","doi-asserted-by":"crossref","DOI":"10.1126\/sciimmunol.aba1896","article-title":"Absence of MHC class II on cDC1 dendritic cells triggers fatal autoimmunity to a cross-presented self-antigen","volume":"5","author":"Wohn","year":"2020","journal-title":"Sci Immunol"},{"key":"10.1016\/j.cellimm.2026.105144_bb0175","doi-asserted-by":"crossref","first-page":"1066","DOI":"10.1016\/j.immuni.2016.10.008","article-title":"Immunomodulatory functions of BTLA and HVEM govern induction of Extrathymic regulatory T cells and tolerance by dendritic cells","volume":"45","author":"Jones","year":"2016","journal-title":"Immunity"},{"key":"10.1016\/j.cellimm.2026.105144_bb0180","doi-asserted-by":"crossref","first-page":"1420","DOI":"10.1002\/eji.201040930","article-title":"Steady state migratory RelB+ langerin+ dermal dendritic cells mediate peripheral induction of antigen-specific CD4+ CD25+ Foxp3+ regulatory T cells","volume":"41","author":"Azukizawa","year":"2011","journal-title":"Eur. J. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0185","doi-asserted-by":"crossref","first-page":"670","DOI":"10.1038\/ni944","article-title":"BTLA is a lymphocyte inhibitory receptor with similarities to CTLA-4 and PD-1","volume":"4","author":"Watanabe","year":"2003","journal-title":"Nat. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0190","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1038\/s41577-024-00994-x","article-title":"The regulation and differentiation of regulatory T cells and their dysfunction in autoimmune diseases","volume":"24","author":"Sumida","year":"2024","journal-title":"Nat. Rev. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0195","doi-asserted-by":"crossref","first-page":"3677","DOI":"10.1172\/JCI57379","article-title":"Desmoglein 3-specific CD4+ T cells induce pemphigus vulgaris and interface dermatitis in mice","volume":"121","author":"Takahashi","year":"2011","journal-title":"J. Clin. Invest."},{"key":"10.1016\/j.cellimm.2026.105144_bb0200","doi-asserted-by":"crossref","first-page":"582","DOI":"10.4049\/jimmunol.2100781","article-title":"IgM to IgG class switching is a necessary step for pemphigus phenotype induction in Desmoglein 3-specific B cell receptor Knock-in mouse","volume":"208","author":"Nomura","year":"2022","journal-title":"J. Immunol."},{"key":"10.1016\/j.cellimm.2026.105144_bb0205","doi-asserted-by":"crossref","DOI":"10.3389\/fimmu.2022.931821","article-title":"Herpesvirus entry mediator on T cells as a protective factor for myasthenia gravis: a Mendelian randomization study","volume":"13","author":"Zhong","year":"2022","journal-title":"Front. Immunol."}],"container-title":["Cellular Immunology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0008874926000857?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0008874926000857?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,8,27]],"date-time":"2026-08-27T00:35:09Z","timestamp":1787790909000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0008874926000857"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9]]},"references-count":41,"alternative-id":["S0008874926000857"],"URL":"https:\/\/doi.org\/10.1016\/j.cellimm.2026.105144","relation":{},"ISSN":["0008-8749"],"issn-type":[{"value":"0008-8749","type":"print"}],"subject":[],"published":{"date-parts":[[2026,9]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"HVEM expressed in CD4 T cells promotes self-tolerance to an epidermal antigen through Treg expansion and modulation of B cell responses","name":"articletitle","label":"Article Title"},{"value":"Cellular Immunology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.cellimm.2026.105144","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 The Authors. Published by Elsevier Inc.","name":"copyright","label":"Copyright"}],"article-number":"105144"}}