{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,24]],"date-time":"2026-07-24T14:31:15Z","timestamp":1784903475379,"version":"3.55.0"},"reference-count":77,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2021,11,3]],"date-time":"2021-11-03T00:00:00Z","timestamp":1635897600000},"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>The mucosal immune system is the first line of defense against pathogens. Germinal centers (GCs) in the Peyer\u2019s patches (PPs) of the small intestine are constantly generated through stimulation of the microbiota. In this study, we investigated the role of \u03b3\u03b4 T cells in the GC reactions in PPs. Most \u03b3\u03b4 T cells in PPs localized in the GCs and expressed a TCR composed of V\u03b31 and V\u03b46 chains. By using mice with partial and total \u03b3\u03b4 T cell deficiencies, we found that V\u03b31<jats:sup>+<\/jats:sup>\/V\u03b46<jats:sup>+<\/jats:sup>T cells can produce high amounts of IL-4, which drives the proliferation of GC B cells as well as the switch of GC B cells towards IgA. Therefore, we conclude that \u03b3\u03b4 T cells play a role in sustaining gut homeostasis and symbiosis<jats:italic>via<\/jats:italic>supporting the GC reactions in PPs.<\/jats:p>","DOI":"10.3389\/fimmu.2021.729607","type":"journal-article","created":{"date-parts":[[2021,11,3]],"date-time":"2021-11-03T06:58:43Z","timestamp":1635922723000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":21,"title":["IL-4-Producing V\u03b31+\/V\u03b46+ \u03b3\u03b4 T Cells Sustain Germinal Center Reactions in Peyer\u2019s Patches of Mice"],"prefix":"10.3389","volume":"12","author":[{"given":"Leon","family":"Ullrich","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yvonne","family":"Lueder","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anna-Lena","family":"Juergens","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anneke","family":"Wilharm","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Joana","family":"Barros-Martins","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anja","family":"Bubke","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Abdi","family":"Demera","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Koichi","family":"Ikuta","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gwendolyn Elena","family":"Patzer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anika","family":"Janssen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Inga","family":"Sandrock","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Immo","family":"Prinz","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Francesca","family":"Rampoldi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2021,11,3]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2016.00342","article-title":"Development and Function of Secondary and Tertiary Lymphoid Organs in the Small Intestine and the Colon","volume":"7","author":"Buettner","year":"2016","journal-title":"Front Immunol"},{"key":"B2","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1038\/mi.2012.96","article-title":"Guarding the Perimeter: Protection of the Mucosa by Tissue-Resident Memory T Cells","volume":"6","author":"Cauley","year":"2013","journal-title":"Mucosal Immunol"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.1111\/imr.12400","article-title":"Peyer\u2019s Patches: Organizing B-Cell Responses at the Intestinal Frontier","volume":"271","author":"Reboldi","year":"2016","journal-title":"Immunol Rev"},{"key":"B4","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":"B5","article-title":"Beitrag Zur Cytologie Der Keimzentren Der Lymphknoten","volume":"20","author":"R\u00f6hlich","year":"1930","journal-title":"Z Mikrosk Anat Forsch"},{"key":"B6","article-title":"Long-Term Antigen Retention by Dendritic Cells in the Popliteal Lymph Node of Immunized Mice","volume":"43","author":"Mandel","year":"1981","journal-title":"Immunology"},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.1016\/S0092-8674(00)00078-7","article-title":"Class Switch Recombination and Hypermutation Require Activation-Induced Cytidine Deaminase (AID), a Potential RNA Editing Enzyme","volume":"102","author":"Muramatsu","year":"2000","journal-title":"Cell"},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.1126\/science.288.5474.2222","article-title":"A Primitive T Cell-Independent Mechanism of Intestinal Mucosal IgA Responses to Commensal Bacteria","volume":"288","author":"Macpherson","year":"2000","journal-title":"Science"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.1038\/nri3322","article-title":"New Concepts in the Generation and Functions of IgA","volume":"12","author":"Pabst","year":"2012","journal-title":"Nat Rev Immunol"},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.1111\/j.1600-065X.2011.01072.x","article-title":"The Habitat, Double Life, Citizenship, and Forgetfulness of IgA","volume":"245","author":"Macpherson","year":"2012","journal-title":"Immunol Rev"},{"key":"B11","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1084\/jem.181.1.41","article-title":"Impaired Mucosal Immune Responses in Interleukin 4-Targeted Mice","volume":"181","author":"Vajdy","year":"1995","journal-title":"J Exp Med"},{"key":"B12","doi-asserted-by":"crossref","DOI":"10.4049\/jimmunol.154.7.3156","article-title":"Novel In Vitro Model for High-Rate IgA Class Switching","volume":"154","author":"McIntyre","year":"1995","journal-title":"J Immunol"},{"key":"B13","doi-asserted-by":"crossref","DOI":"10.4049\/jimmunol.139.8.2669","article-title":"Interleukin 5 and Interleukin 4 Produced by Peyer\u2019s Patch T Cells Selectively Enhance Immunoglobulin A Expression","volume":"139","author":"Murray","year":"1987","journal-title":"J Immunol"},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.173.5.3305","article-title":"TGF-\u03b2 Receptor Signaling Is Critical for Mucosal IgA Responses","volume":"173","author":"Borsutzky","year":"2004","journal-title":"J Immunol"},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1084\/jem.170.3.1039","article-title":"Transforming Growth Factor Beta Specifically Enhances IgA Production by Lipopolysaccharide-Stimulated Murine B Lymphocytes","volume":"170","author":"Coffman","year":"1989","journal-title":"J Exp Med"},{"key":"B16","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1016\/j.smim.2007.12.001","article-title":"Follicular Dendritic Cell Networks of Primary Follicles and Germinal Centers: Phenotype and Function","volume":"20","author":"Allen","year":"2008","journal-title":"Semin Immunol"},{"key":"B17","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/S1074-7613(01)00257-6","article-title":"CXCL13 Is Required for B1 Cell Homing, Natural Antibody Production, and Body Cavity Immunity","volume":"16","author":"Ansel","year":"2002","journal-title":"Immunity"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.1016\/S0092-8674(00)81798-5","article-title":"A Putative Chemokine Receptor, BLR1, Directs B Cell Migration to Defined Lymphoid Organs and Specific Anatomic Compartments of the Spleen","volume":"87","author":"F\u00f6rster","year":"1996","journal-title":"Cell"},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.179.8.5099","article-title":"Role of CXCR5 and CCR7 in Follicular Th Cell Positioning and Appearance of a Programmed Cell Death Gene-1 High Germinal Center-Associated Subpopulation","volume":"179","author":"Haynes","year":"2007","journal-title":"J Immunol"},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.1126\/science.1136736","article-title":"Imaging of Germinal Center Selection Events During Affinity Maturation","volume":"315","author":"Allen","year":"2007","journal-title":"Science (80-)"},{"key":"B21","doi-asserted-by":"publisher","first-page":"592","DOI":"10.1016\/j.cell.2010.10.032","article-title":"Germinal Center Dynamics Revealed by Multiphoton Microscopy With a Photoactivatable Fluorescent Reporter","volume":"143","author":"Victora","year":"2010","journal-title":"Cell"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.1038\/nri3804","article-title":"Dynamics of B Cells in Germinal Centres","volume":"15","author":"De Silva","year":"2015","journal-title":"Nat Rev Immunol"},{"key":"B23","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1016\/s0165-2478(97)87000-9","article-title":"Somatic Mutation Leads to Efficient Affinity Maturation When Centrocytes Recycle Back to Centroblasts","volume":"56","author":"Oprea","year":"1997","journal-title":"Immunol Lett"},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.1038\/ni1100","article-title":"Germinal Center Dark and Light Zone Organization Is Mediated by CXCR4 and CXCR5","volume":"5","author":"Allen","year":"2004","journal-title":"Nat Immunol"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.1038\/ni.1814","article-title":"Multiple Layers of B Cell Memory With Different Effector Functions","volume":"10","author":"Dogan","year":"2009","journal-title":"Nat Immunol"},{"key":"B26","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1146\/annurev.iy.11.040193.000405","article-title":"Immunological Memory","volume":"11","author":"Gray","year":"1993","journal-title":"Annu Rev Immunol"},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.1038\/nri1572","article-title":"Regulation of Plasma-Cell Development","volume":"5","author":"Shapiro-Shelef","year":"2005","journal-title":"Nat Rev Immunol"},{"key":"B28","doi-asserted-by":"publisher","DOI":"10.3390\/cells9030743","article-title":"Revisiting the Interaction of \u03b3\u03b4 T-Cells and B-Cells","volume":"9","author":"Rampoldi","year":"2020","journal-title":"Cells"},{"key":"B29","doi-asserted-by":"crossref","DOI":"10.1016\/bs.ai.2017.01.002","volume-title":"\u03b3\u03b4 T Cells and B Cells","author":"Born","year":"2017"},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2014.00486","article-title":"Ontogeny of Innate T Lymphocytes - Some Innate Lymphocytes Are More Innate Than Others","volume":"5","author":"Vermijlen","year":"2014","journal-title":"Front Immunol"},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.1084\/jem.172.4.1225","article-title":"Late Dominance of the Inflammatory Process in Murine Influenza by Gamma\/Delta + T Cells","volume":"172","author":"Carding","year":"1990","journal-title":"J Exp Med"},{"key":"B32","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0903895106","article-title":"TCR-Inducible PLZF Transcription Factor Required for Innate Phenotype of a Subset of T Cells With Restricted TCR Diversity","volume":"106","author":"Kreslavsky","year":"2009","journal-title":"Proc Natl Acad Sci"},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.1038\/nri2781","article-title":"\u03b3\u03b4 T Cell Effector Functions: A Blend of Innate Programming and Acquired Plasticity","volume":"10","author":"Bonneville","year":"2010","journal-title":"Nat Rev Immunol"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.165.8.4174","article-title":"V\u03b31 + T Cells Suppress and V\u03b34 + T Cells Promote Susceptibility to Coxsackievirus B3-Induced Myocarditis in Mice","volume":"165","author":"Huber","year":"2000","journal-title":"J Immunol"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1002\/eji.1830230151","article-title":"NK1.1+ CD4+ CD8- Thymocytes With Specific Lymphokine Secretion","volume":"23","author":"Arase","year":"1993","journal-title":"Eur J Immunol"},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.171.12.6488","article-title":"The Interaction of \u03b3\u03b4 T Cells With Activated Macrophages Is a Property of the V\u03b31 Subset","volume":"171","author":"Dalton","year":"2003","journal-title":"J Immunol"},{"key":"B37","doi-asserted-by":"crossref","DOI":"10.4049\/jimmunol.163.6.3076","article-title":"IL-4-Producing Gamma Delta T Cells That Express a Very Restricted TCR Repertoire Are Preferentially Localized in Liver and Spleen","volume":"163","author":"Gerber","year":"1999","journal-title":"J Immunol"},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.1016\/j.micinf.2006.11.015","article-title":"Accumulation of Gamma\/Delta T Cells in the Lungs and Their Roles in Neutrophil-Mediated Host Defense Against Pneumococcal Infection","volume":"9","author":"Nakasone","year":"2007","journal-title":"Microbes Infect"},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.0903332","article-title":"Essential Role of IL-17A in the Formation of a Mycobacterial Infection-Induced Granuloma in the Lung","volume":"184","author":"Okamoto Yoshida","year":"2010","journal-title":"J Immunol"},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.1016\/j.it.2012.11.004","article-title":"The Natural and the Inducible: Interleukin (IL)-17-Producing \u03b3\u03b4 T Cells","volume":"34","author":"Chien","year":"2013","journal-title":"Trends Immunol"},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1415107111","article-title":"\u03b3\u03b4 T Cells Affect IL-4 Production and B-Cell Tolerance","volume":"112","author":"Huang","year":"2015","journal-title":"Proc Natl Acad Sci"},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20181439","article-title":"Genetic Models Reveal Origin, Persistence and Non-Redundant Functions of IL-17\u2013Producing \u03b3\u03b4 T Cells","volume":"215","author":"Sandrock","year":"2018","journal-title":"J Exp Med"},{"key":"B43","doi-asserted-by":"publisher","first-page":"995","DOI":"10.1038\/ni1371","article-title":"Visualization of the Earliest Steps of \u03b3\u03b4 T Cell Development in the Adult Thymus","volume":"7","author":"Prinz","year":"2006","journal-title":"Nat Immunol"},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.1016\/0092-8674(93)90112-4","article-title":"T Cell Receptor \u03b4 Gene Mutant Mice: Independent Generation of \u03b1\u03b2 T Cells and Programmed Rearrangements of \u03b3\u03b4 TCR Genes","volume":"72","author":"Itohara","year":"1993","journal-title":"Cell"},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.175.3.1741","article-title":"Delineation of the Function of a Major \u03b3\u03b4 T Cell Subset During Infection","volume":"175","author":"Andrew","year":"2005","journal-title":"J Immunol"},{"key":"B46","doi-asserted-by":"crossref","DOI":"10.4049\/jimmunol.158.9.4223","article-title":"Developmentally Ordered V-J Recombination in Mouse T Cell Receptor Gamma Locus Is Not Perturbed by Targeted Deletion of the Vgamma4 Gene","volume":"158","author":"Sunaga","year":"1997","journal-title":"J Immunol"},{"key":"B47","doi-asserted-by":"publisher","first-page":"68","DOI":"10.1189\/jlb.0703326","article-title":"Subset-Specific, Uniform Activation Among V\u03b36\/V\u03b41 + \u03b3\u03b4 T Cells Elicited by Inflammation","volume":"75","author":"Roark","year":"2004","journal-title":"J Leukoc Biol"},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.1038\/s41385-019-0175-z","article-title":"BLT1 Mediates Commensal Bacteria-Dependent Innate Immune Signals to Enhance Antigen-Specific Intestinal IgA Responses","volume":"12","author":"Nagatake","year":"2019","journal-title":"Mucosal Immunol"},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.1016\/j.it.2011.09.005","article-title":"T Cell Receptor Signalling in \u03b3\u03b4 Cell Development: Strength Isn\u2019t Everything","volume":"32","author":"Turchinovich","year":"2011","journal-title":"Trends Immunol"},{"key":"B50","doi-asserted-by":"publisher","DOI":"10.1002\/eji.200939922","article-title":"CCR6 and NK1.1 Distinguish Between IL-17A and IFN-\u03b3-Producing \u03b3\u03b4 Effector T Cells","volume":"39","author":"Haas","year":"2009","journal-title":"Eur J Immunol"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.1002\/eji.201142109","article-title":"Innate Nkt\u03b3\u03b4 and Nkt\u03b1\u03b2 Cells Exert Similar Functions and Compete for a Thymic Niche","volume":"42","author":"Pereira","year":"2012","journal-title":"Eur J Immunol"},{"key":"B52","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2014.10.004","article-title":"T Follicular Helper Cell Differentiation, Function, and Roles in Disease","volume":"41","author":"Crotty","year":"2014","journal-title":"Immunity"},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2012-03-415380","article-title":"Identification of Human Germinal Center Light and Dark Zone Cells and Their Relationship to Human B-Cell Lymphomas","volume":"120","author":"Victora","year":"2012","journal-title":"Blood"},{"key":"B54","doi-asserted-by":"publisher","DOI":"10.1002\/aja.1001700315","article-title":"Functional Anatomy of Germinal Centers","volume":"170","author":"Nieuwenhuis","year":"1984","journal-title":"Am J Anat"},{"key":"B55","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":"B56","doi-asserted-by":"publisher","DOI":"10.1016\/S0167-5699(98)01327-9","article-title":"The Dynamic Structure of the Germinal Center","volume":"19","author":"Camacho","year":"1998","journal-title":"Immunol Today"},{"key":"B57","doi-asserted-by":"publisher","DOI":"10.1038\/291238a0","article-title":"Aromatic-Dependent Salmonella Typhimurium Are Non-Virulent and Effective as Live Vaccines","volume":"291","author":"Hoiseth","year":"1981","journal-title":"Nature"},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.1016\/j.coi.2017.03.007","article-title":"Deconstructing the Germinal Center, One Cell at a Time","volume":"45","author":"Dufaud","year":"2017","journal-title":"Curr Opin Immunol"},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1501064","article-title":"\u03b3\u03b4 T Cells Shape Preimmune Peripheral B Cell Populations","volume":"196","author":"Huang","year":"2015","journal-title":"J Immunol"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.1038\/icb.2009.105","article-title":"Balanced Approach of \u03b3\u03b4 T Cells to Type 2 Immunity","volume":"88","author":"Born","year":"2010","journal-title":"Immunol Cell Biol"},{"key":"B61","doi-asserted-by":"publisher","first-page":"3151","DOI":"10.1038\/s41467-018-05487-9","article-title":"\u03b3\u03b4 T Cells Control Humoral Immune Response by Inducing T Follicular Helper Cell Differentiation","volume":"9","author":"Rezende","year":"2018","journal-title":"Nat Commun"},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.1002\/eji.201344203","article-title":"Voehringer D. B-Cell-Intrinsic STAT6 Signaling Controls Germinal Center Formation","volume":"44","author":"Turqueti-Neves","year":"2014","journal-title":"Eur J Immunol"},{"key":"B63","doi-asserted-by":"publisher","DOI":"10.1111\/imcb.12283","article-title":"BAFF, IL-4 and IL-21 Separably Program Germinal Center-Like Phenotype Acquisition, BCL6 Expression, Proliferation and Survival of CD40L-Activated B Cells In Vitro","volume":"97","author":"Robinson","year":"2019","journal-title":"Immunol Cell Biol"},{"key":"B64","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20090313","article-title":"IL-4\u2013Producing CD4+ T Cells in Reactive Lymph Nodes During Helminth Infection Are T Follicular Helper Cells","volume":"206","author":"King","year":"2009","journal-title":"J Exp Med"},{"key":"B65","doi-asserted-by":"publisher","DOI":"10.1038\/ni.1715","article-title":"Cytokine-Secreting Follicular T Cells Shape the Antibody Repertoire","volume":"10","author":"Reinhardt","year":"2009","journal-title":"Nat Immunol"},{"key":"B66","doi-asserted-by":"publisher","DOI":"10.1038\/ni.2552","article-title":"Plasticity of TH17 Cells in Peyer\u2019s Patches Is Responsible for the Induction of T Cell\u2013Dependent IgA Responses","volume":"14","author":"Hirota","year":"2013","journal-title":"Nat Immunol"},{"key":"B67","doi-asserted-by":"publisher","DOI":"10.1126\/science.1169152","article-title":"Preferential Generation of Follicular B Helper T Cells From Foxp3+ T Cells in Gut Peyer\u2019s Patches","volume":"323","author":"Tsuji","year":"2009","journal-title":"Science"},{"key":"B68","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1038\/icb.2013.54","article-title":"Gut T FH and IgA: Key Players for Regulation of Bacterial Communities and Immune Homeostasis","volume":"92","author":"Kato","year":"2014","journal-title":"Immunol Cell Biol"},{"key":"B69","doi-asserted-by":"publisher","DOI":"10.1111\/imr.12031","article-title":"Vinuesa CG. T-Cell Subsets in the Germinal Center","volume":"252","author":"Ramiscal","year":"2013","journal-title":"Immunol Rev"},{"key":"B70","doi-asserted-by":"publisher","DOI":"10.1016\/S0960-9822(02)70718-5","article-title":"\u03b3\u03b4 T Cell Help of B Cells Is Induced by Repeated Parasitic Infection, in the Absence of Other T Cells","volume":"6","author":"Pao","year":"1996","journal-title":"Curr Biol"},{"key":"B71","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.171.10.4979","article-title":"Cutting Edge: \u03b3\u03b4 T Cells Provide Help to B Cells With Altered Clonotypes and Are Capable of Inducing Ig Gene Hypermutation","volume":"171","author":"Zheng","year":"2003","journal-title":"J Immunol"},{"key":"B72","doi-asserted-by":"publisher","DOI":"10.1084\/jem.183.4.1929","article-title":"Gamma\/Delta T Cell-Deficient Mice Have Impaired Mucosal Immunoglobulin A Responses","volume":"183","author":"Fujihashi","year":"1996","journal-title":"J Exp Med"},{"key":"B73","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.177.8.5290","article-title":"CXCR5 Identifies a Subset of V\u03b39v\u03b42 T Cells Which Secrete IL-4 and IL-10 and Help B Cells for Antibody Production","volume":"177","author":"Caccamo","year":"2006","journal-title":"J Immunol"},{"key":"B74","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2003-04-1016","article-title":"Flexible Migration Program Regulates \u03b3\u03b4 T-Cell Involvement in Humoral Immunity","volume":"102","author":"Brandes","year":"2003","journal-title":"Blood"},{"key":"B75","doi-asserted-by":"publisher","DOI":"10.1084\/jem.192.11.1545","article-title":"Follicular B Helper T Cells Express Cxc Chemokine Receptor 5, Localize to B Cell Follicles, and Support Immunoglobulin Production","volume":"192","author":"Breitfeld","year":"2000","journal-title":"J Exp Med"},{"key":"B76","doi-asserted-by":"publisher","DOI":"10.1084\/jem.192.11.1553","article-title":"Cxc Chemokine Receptor 5 Expression Defines Follicular Homing T Cells With B Cell Helper Function","volume":"192","author":"Schaerli","year":"2000","journal-title":"J Exp Med"},{"key":"B77","doi-asserted-by":"publisher","DOI":"10.1046\/j.1365-2249.2001.01494.x","article-title":"Oligoclonally Expanding \u03b3\u03b4 T Lymphocytes Induce IgA Switching in IgA Nephropathy","volume":"124","author":"Toyabe","year":"2001","journal-title":"Clin Exp Immunol"}],"container-title":["Frontiers in Immunology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2021.729607\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,1,14]],"date-time":"2023-01-14T10:57:03Z","timestamp":1673693823000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2021.729607\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,11,3]]},"references-count":77,"alternative-id":["10.3389\/fimmu.2021.729607"],"URL":"https:\/\/doi.org\/10.3389\/fimmu.2021.729607","relation":{},"ISSN":["1664-3224"],"issn-type":[{"value":"1664-3224","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,11,3]]},"article-number":"729607"}}