{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T21:35:23Z","timestamp":1790804123791,"version":"4.1.0"},"reference-count":64,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2025,8,21]],"date-time":"2025-08-21T00:00:00Z","timestamp":1755734400000},"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. Immunol."],"abstract":"<jats:p>Immunoglobulins (IGs) made by chronic lymphocytic leukemia (CLL) B cells are unique in that they bind themselves (homo-dimerize). This interaction leads to signal transduction with functional consequences that depend on the affinity of homo-dimerization. We have studied the antigen-binding properties of the IGs from a subset of patients with CLL (Subset #4) that homo-dimerize at high affinity. Previously, we had found that subset #4 IGs bound viable lymphocytes. Our new studies, probing an array of &amp;gt;8,000 antigens, indicate that these IGs also bind influenza virus. Because of the IGs high-affinity homo-dimerization, we asked if the defined foreign- and self-antigenic interactions were mediated by conventional B-cell receptor (BCR) domains or a non-conventional receptor created by homo-dimerization. The studies indicated the latter since abrogation of homo-dimerization eliminated binding to influenza virus and its hemagglutinin and to viable lymphocytes. Using these findings, we modeled a developmental path whereby a naive IgM<jats:sup>+<\/jats:sup> B cell with subset #4 heavy and light chain variable domains used the conventional BCR to interact with auto- and foreign antigens and acquire homo-dimerization capacity to create the non-conventional antigen-receptor when transitioning to a leukemic cell. Future studies will determine if this process is an idiosyncratic occurrence or a physiologic principle.<\/jats:p>","DOI":"10.3389\/fimmu.2025.1607189","type":"journal-article","created":{"date-parts":[[2025,8,21]],"date-time":"2025-08-21T05:28:24Z","timestamp":1755754104000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":2,"title":["Conventional and non-conventional antigen-binding sites promote the development and function of chronic lymphocytic leukemia stereotyped subset #4 clones"],"prefix":"10.3389","volume":"16","author":[{"given":"Yun","family":"Liu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dzmitry","family":"Padhorny","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rosa","family":"Catera","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Antonella","family":"Nicolo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiao-Jie","family":"Yan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stan Xiaogang","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anastasia","family":"Iatrou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrea N.","family":"Mazzarello","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Noemi","family":"Destefani","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Steven L.","family":"Allen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jonathan E.","family":"Kolitz","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kanti R.","family":"Rai","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Massimo","family":"Degano","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paolo P.","family":"Ghia","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Charles C.","family":"Chu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Florian","family":"Krammer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hassan","family":"Jumaa","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kostas","family":"Stamatopoulos","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dima","family":"Kozakov","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nicholas","family":"Chiorazzi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2025,8,21]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"787","DOI":"10.1016\/j.cell.2004.05.014","article-title":"Survival of resting mature B lymphocytes depends on BCR signaling via the Igalpha\/beta heterodimer","volume":"117","author":"Kraus","year":"2004","journal-title":"Cell"},{"key":"B2","doi-asserted-by":"publisher","DOI":"10.1016\/S0092-8674(00)80373-6","article-title":"In vivo ablation of surface immunoglobulin on mature B cells by inducible gene targeting results in rapid cell death","volume":"90","author":"Lam","year":"1997","journal-title":"Cell"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.iy.12.040194.001001","article-title":"Germinal centers","volume":"12","author":"MacLennan","year":"1994","journal-title":"Annu Rev Immunol"},{"key":"B4","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1034\/j.1600-065X.2003.00058.x","article-title":"Extrafollicular antibody responses","volume":"194","author":"MacLennan","year":"2003","journal-title":"Immunol Rev"},{"key":"B5","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-immunol-020711-075032","article-title":"Germinal centers","volume":"30","author":"Victora","year":"2012","journal-title":"Annu Rev Immunol"},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1038\/ni959","article-title":"The nonimmunoglobulin portion of lambda5 mediates cell-autonomous pre-B cell receptor signaling","volume":"4","author":"Ohnishi","year":"2003","journal-title":"Nat Immunol"},{"key":"B7","doi-asserted-by":"publisher","first-page":"323","DOI":"10.1016\/j.immuni.2007.01.012","article-title":"Conventional light chains inhibit the autonomous signaling capacity of the B cell receptor","volume":"26","author":"Meixlsperger","year":"2007","journal-title":"Immunity"},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2008.10.013","article-title":"Autoreactive B cell receptors mimic autonomous pre-B cell receptor signaling and induce proliferation of early B cells","volume":"29","author":"K\u00f6hler","year":"2008","journal-title":"Immunity"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.1038\/nature11309","article-title":"Chronic lymphocytic leukaemia is driven by antigen-independent cell-autonomous signalling","volume":"489","author":"Duhren-von Minden","year":"2012","journal-title":"Nature"},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.1016\/S1074-7613(00)80029-1","article-title":"Negative selection of immature B cells by receptor editing or deletion is determined by site of antigen encounter","volume":"10","author":"Sandel","year":"1999","journal-title":"Immunity"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.1016\/j.it.2011.06.004","article-title":"BAFF and selection of autoreactive B cells","volume":"32","author":"Liu","year":"2011","journal-title":"Trends Immunol"},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.050552997","article-title":"A B-cell receptor-specific selection step governs immature to mature B cell differentiation","volume":"97","author":"Levine","year":"2000","journal-title":"Proc Natl Acad Sci U S A"},{"key":"B13","doi-asserted-by":"publisher","first-page":"429","DOI":"10.12688\/f1000research","article-title":"Role of B-cell receptors for B-cell development and antigen-induced differentiation","volume":"7","author":"Yam-Puc","year":"2018","journal-title":"F1000Res"},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2019.03.016","article-title":"B cell responses: cell interaction dynamics and decisions","volume":"177","author":"Cyster","year":"2019","journal-title":"Cell"},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20230941","article-title":"Antigen-independent, autonomous B cell receptor signaling drives activated B cell DLBCL","volume":"221","author":"Eken","year":"2024","journal-title":"J Exp Med"},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2012-09-454439","article-title":"CLL B-cell receptors can recognize themselves: alternative epitopes and structural clues for autostimulatory mechanisms in CLL","volume":"121","author":"Binder","year":"2013","journal-title":"Blood"},{"key":"B17","doi-asserted-by":"publisher","first-page":"15746","DOI":"10.1038\/ncomms15746","article-title":"Distinct homotypic B-cell receptor interactions shape the outcome of chronic lymphocytic leukaemia","volume":"8","author":"Minici","year":"2017","journal-title":"Nat Commun"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.immunol.21.120601.141018","article-title":"B cell chronic lymphocytic leukemia: lessons learned from studies of the B cell antigen receptor","volume":"21","author":"Chiorazzi","year":"2003","journal-title":"Annu Rev Immunol"},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.1172\/JCI3009","article-title":"Chronic lymphocytic leukemia B cells express restricted sets of mutated and unmutated antigen receptors","volume":"102","author":"Fais","year":"1998","journal-title":"J Clin Invest"},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2002-11-3485","article-title":"Chronic lymphocytic leukemias utilizing the VH3\u201321 gene display highly restricted V{lambda}2\u201314 gene use and homologous CDR3s: implicating recognition of a common antigen epitope","volume":"101","author":"Tobin","year":"2003","journal-title":"Blood"},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2004-01-0132","article-title":"Subsets with restricted immunoglobulin gene rearrangement features indicate a role for antigen selection in the development of chronic lymphocytic leukemia","volume":"104","author":"Tobin","year":"2004","journal-title":"Blood"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.1172\/JCI19399","article-title":"Remarkably similar antigen receptors among a subset of patients with chronic lymphocytic leukemia","volume":"113","author":"Ghiotto","year":"2004","journal-title":"J Clin Invest"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20040544","article-title":"Multiple distinct sets of stereotyped antigen receptors indicate a role for antigen in promoting chronic lymphocytic leukemia","volume":"200","author":"Messmer","year":"2004","journal-title":"J Exp Med"},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.1053\/j.seminhematol.2023.12.005","article-title":"B-cell receptor immunoglobulin stereotypy in chronic lymphocytic leukemia: Key to understanding disease biology and stratifying patients","volume":"61","author":"Agathangelidis","year":"2024","journal-title":"Semin Hematol"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2007-07-099564","article-title":"Stereotyped patterns of somatic hypermutation in subsets of patients with chronic lymphocytic leukemia: implications for the role of antigen selection in leukemogenesis","volume":"111","author":"Murray","year":"2008","journal-title":"Blood"},{"key":"B26","doi-asserted-by":"publisher","DOI":"10.3324\/haematol.2016.141812","article-title":"Different spectra of recurrent gene mutations in subsets of chronic lymphocytic leukemia harboring stereotyped B-cell receptors","volume":"101","author":"Sutton","year":"2016","journal-title":"Haematologica"},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.1158\/1078-0432.CCR-16-3100","article-title":"Chronic lymphocytic leukemia with mutated IGHV4\u201334 receptors: shared and distinct immunogenetic features and clinical outcomes","volume":"23","author":"Xochelli","year":"2017","journal-title":"Clin Cancer Res"},{"key":"B28","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2006-03-012948","article-title":"Over 20% of patients with chronic lymphocytic leukemia carry stereotyped receptors: Pathogenetic implications and clinical correlations","volume":"109","author":"Stamatopoulos","year":"2007","journal-title":"Blood"},{"key":"B29","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2009-05-221309","article-title":"Extensive intraclonal diversification in a subgroup of chronic lymphocytic leukemia patients with stereotyped IGHV4\u201334 receptors: implications for ongoing interactions with antigen","volume":"114","author":"Sutton","year":"2009","journal-title":"Blood"},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2008-07-166868","article-title":"Evidence for the significant role of immunoglobulin light chains in antigen recognition and selection in chronic lymphocytic leukemia","volume":"113","author":"Hadzidimitriou","year":"2009","journal-title":"Blood"},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.1158\/1078-0432.CCR-13-1993","article-title":"IgG-switched CLL has a distinct immunogenetic signature from the common MD variant: ontogenetic implications","volume":"20","author":"Vardi","year":"2014","journal-title":"Clin Cancer Res"},{"key":"B32","doi-asserted-by":"publisher","DOI":"10.1038\/leu.2014.196","article-title":"Recurrent mutations refine prognosis in chronic lymphocytic leukemia","volume":"29","author":"Baliakas","year":"2014","journal-title":"Leukemia"},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2007-09-111344","article-title":"Constitutive activation of distinct BCR-signaling pathways in a subset of CLL patients: a molecular signature of anergy","volume":"112","author":"Muzio","year":"2008","journal-title":"Blood"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1502297","article-title":"B Cell Anergy Modulated by TLR1\/2 and the miR-17 approximately 92 Cluster Underlies the Indolent Clinical Course of Chronic Lymphocytic Leukemia Stereotyped Subset 4","volume":"196","author":"Ntoufa","year":"2016","journal-title":"J Immunol"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2007-11-125450","article-title":"A new perspective: molecular motifs on oxidized LDL, apoptotic cells, and bacteria are targets for chronic lymphocytic leukemia antibodies","volume":"111","author":"Lanemo Myhrinder","year":"2008","journal-title":"Blood"},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.2119\/2008-00102.Catera","article-title":"Chronic lymphocytic leukemia cells recognize conserved epitopes associated with apoptosis and oxidation","volume":"14","author":"Catera","year":"2008","journal-title":"Mol Med"},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2009-09-244251","article-title":"Many chronic lymphocytic leukemia antibodies recognize apoptotic cells with exposed nonmuscle myosin heavy chain IIA: implications for patient outcome and cell of origin","volume":"115","author":"Chu","year":"2010","journal-title":"Blood"},{"key":"B38","doi-asserted-by":"publisher","first-page":"1","DOI":"10.2119\/molmed.2017.00003","article-title":"Binding of CLL subset 4 B-cell receptor immunoglobulins to viable human memory B lymphocytes requires a distinctive IGKV somatic mutation","volume":"23","author":"Catera","year":"2017","journal-title":"Mol Med"},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1016\/0896-8411(88)90068-6","article-title":"Chronic lymphocytic leukemic cells secrete multispecific autoantibodies","volume":"1","author":"Broker","year":"1988","journal-title":"J Autoimmun"},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.1084\/jem.169.1.255","article-title":"Production of autoantibodies by CD5-expressing B lymphocytes from patients with chronic lymphocytic leukemia","volume":"169","author":"Sthoeger","year":"1989","journal-title":"J Exp Med"},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.1182\/blood.V76.3.562.562","article-title":"Evidence that chronic lymphocytic leukemia B lymphocytes are frequently committed to production of natural autoantibodies","volume":"76","author":"Borche","year":"1990","journal-title":"Blood"},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.1172\/JCI24387","article-title":"Unmutated and mutated chronic lymphocytic leukemias derive from self-reactive B cell precursors despite expressing different antibody reactivity","volume":"115","author":"Herve","year":"2005","journal-title":"J Clin Invest"},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2009-01-197822","article-title":"Characterization of structurally defined epitopes recognized by monoclonal antibodies produced by chronic lymphocytic leukemia B cells","volume":"114","author":"Seiler","year":"2009","journal-title":"Blood"},{"key":"B44","doi-asserted-by":"publisher","first-page":"e15992","DOI":"10.1371\/journal.pone.0015992","article-title":"Stereotypical chronic lymphocytic leukemia B-cell receptors recognize survival promoting antigens on stromal cells","volume":"5","author":"Binder","year":"2011","journal-title":"PloS One"},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2012-08-447904","article-title":"Autoantigenic targets of B-cell receptors derived from chronic lymphocytic leukemias bind to and induce proliferation of leukemic cells","volume":"121","author":"Zwick","year":"2013","journal-title":"Blood"},{"key":"B46","doi-asserted-by":"publisher","first-page":"e90725","DOI":"10.1371\/journal.pone.0090725","article-title":"IGHV1\u201369 B cell chronic lymphocytic leukemia antibodies cross-react with HIV-1 and hepatitis C virus antigens as well as intestinal commensal bacteria","volume":"9","author":"Hwang","year":"2014","journal-title":"PloS One"},{"key":"B47","doi-asserted-by":"publisher","first-page":"44","DOI":"10.1016\/j.clim.2016.08.020","article-title":"Chronic lymphocytic leukemia immunoglobulins display bacterial reactivity that converges and diverges from auto-\/poly-reactivity and IGHV mutation status","volume":"172","author":"Hatzi","year":"2016","journal-title":"Clin Immunol"},{"key":"B48","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1084\/jem.20121801","article-title":"A mutated B cell chronic lymphocytic leukemia subset that recognizes and responds to fungi","volume":"210","author":"Hoogeboom","year":"2013","journal-title":"J Exp Med"},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2008-06-162024","article-title":"Chronic lymphocytic leukemia antibodies with a common stereotypic rearrangement recognize nonmuscle myosin heavy chain IIA","volume":"112","author":"Chu","year":"2008","journal-title":"Blood"},{"key":"B50","doi-asserted-by":"publisher","first-page":"e3942","DOI":"10.1371\/journal.pone.0003942","article-title":"Heterosubtypic neutralizing monoclonal antibodies cross-protective against H5N1 and H1N1 recovered from human IgM+ memory B cells","volume":"3","author":"Throsby","year":"2008","journal-title":"PloS One"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2017.11.056","article-title":"Structural determination of the broadly reactive anti-IGHV1-69 anti-idiotypic antibody G6 and its idiotope","volume":"21","author":"Avnir","year":"2017","journal-title":"Cell Rep"},{"key":"B52","doi-asserted-by":"publisher","DOI":"10.1016\/0378-1135(92)90058-2","article-title":"The pathogenic determinant of influenza virus","volume":"33","author":"Rott","year":"1992","journal-title":"Vet Microbiol"},{"key":"B53","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1007\/82_2014_384","article-title":"The hemagglutinin: a determinant of pathogenicity","volume":"385","author":"Bottcher-Friebertshauser","year":"2014","journal-title":"Curr Top Microbiol Immunol"},{"key":"B54","doi-asserted-by":"publisher","first-page":"9","DOI":"10.3201\/eid1201.051254","article-title":"Influenza pandemics of the 20th century","volume":"12","author":"Kilbourne","year":"2006","journal-title":"Emerg Infect Dis"},{"key":"B55","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1126\/science.1176225","article-title":"Antigenic and genetic characteristics of swine-origin 2009 A(H1N1) influenza viruses circulating in humans","volume":"325","author":"Garten","year":"2009","journal-title":"Science"},{"key":"B56","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1007\/978-1-0716-2609-2_1","article-title":"Antibody sequence and structure analyses using IMGT((R)): 30 years of immunoinformatics","volume":"2552","author":"Lefranc","year":"2023","journal-title":"Methods Mol Biol"},{"key":"B57","doi-asserted-by":"publisher","DOI":"10.1038\/nprot.2016.169","article-title":"The ClusPro web server for protein-protein docking","volume":"12","author":"Kozakov","year":"2017","journal-title":"Nat Protoc"},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.1126\/science.1171491","article-title":"Antibody recognition of a highly conserved influenza virus epitope","volume":"324","author":"Ekiert","year":"2009","journal-title":"Science"},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.1126\/science.1222908","article-title":"Highly conserved protective epitopes on influenza B viruses","volume":"337","author":"Dreyfus","year":"2012","journal-title":"Science"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.1126\/scitranslmed.abg4535","article-title":"First exposure to the pandemic H1N1 virus induced broadly neutralizing antibodies targeting hemagglutinin head epitopes","volume":"13","author":"Guthmiller","year":"2021","journal-title":"Sci Transl Med"},{"key":"B61","doi-asserted-by":"publisher","first-page":"337","DOI":"10.1016\/j.immuni.2019.07.001","article-title":"Class-switch recombination occurs infrequently in germinal centers","volume":"51","author":"Roco","year":"2019","journal-title":"Immunity"},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.1126\/science.1086907","article-title":"Predominant autoantibody production by early human B cell precursors","volume":"301","author":"Wardemann","year":"2003","journal-title":"Science"},{"key":"B63","doi-asserted-by":"publisher","DOI":"10.1038\/nrd4529","article-title":"Advances in the development of influenza virus vaccines","volume":"14","author":"Krammer","year":"2015","journal-title":"Nat Rev Drug Discov"},{"key":"B64","doi-asserted-by":"publisher","DOI":"10.1172\/JCI149308","article-title":"B cell receptor isotypes differentially associate with cell signaling, kinetics, and outcome in chronic lymphocytic leukemia","volume":"132","author":"Mazzarello","year":"2022","journal-title":"J Clin Invest"}],"container-title":["Frontiers in Immunology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2025.1607189\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,8,21]],"date-time":"2025-08-21T05:28:25Z","timestamp":1755754105000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2025.1607189\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8,21]]},"references-count":64,"alternative-id":["10.3389\/fimmu.2025.1607189"],"URL":"https:\/\/doi.org\/10.3389\/fimmu.2025.1607189","relation":{},"ISSN":["1664-3224"],"issn-type":[{"value":"1664-3224","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,8,21]]},"article-number":"1607189"}}