{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T21:19:08Z","timestamp":1790284748916,"version":"4.1.0"},"reference-count":72,"publisher":"Oxford University Press (OUP)","issue":"1","license":[{"start":{"date-parts":[[2018,1,1]],"date-time":"2018-01-01T00:00:00Z","timestamp":1514764800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2018,1,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Human CD3+CD4+ Th cells, FOXP3+ T regulatory (Treg) cells, and T regulatory type 1 (Tr1) cells are essential for ensuring peripheral immune response and tolerance, but the diversity of Th, Treg, and Tr1 cell subsets has not been fully characterized. Independent functional characterization of human Th1, Th2, Th17, T follicular helper (Tfh), Treg, and Tr1 cells has helped to define unique surface molecules, transcription factors, and signaling profiles for each subset. However, the adequacy of these markers to recapitulate the whole CD3+CD4+ T cell compartment remains questionable. In this study, we examined CD3+CD4+ T cell populations by single-cell mass cytometry. We characterize the CD3+CD4+ Th, Treg, and Tr1 cell populations simultaneously across 23 memory T cell\u2013associated surface and intracellular molecules. High-dimensional analysis identified several new subsets, in addition to the already defined CD3+CD4+ Th, Treg, and Tr1 cell populations, for a total of 11 Th cell, 4 Treg, and 1 Tr1 cell subsets. Some of these subsets share markers previously thought to be selective for Treg, Th1, Th2, Th17, and Tfh cells, including CD194 (CCR4)+FOXP3+ Treg and CD183 (CXCR3)+T-bet+ Th17 cell subsets. Unsupervised clustering displayed a phenotypic organization of CD3+CD4+ T cells that confirmed their diversity but showed interrelation between the different subsets, including similarity between Th1\u2013Th2\u2013Tfh cell populations and Th17 cells, as well as similarity of Th2 cells with Treg cells. In conclusion, the use of single-cell mass cytometry provides a systems-level characterization of CD3+CD4+ T cells in healthy human blood, which represents an important baseline reference to investigate abnormalities of different subsets in immune-mediated pathologies.<\/jats:p>","DOI":"10.4049\/jimmunol.1701025","type":"journal-article","created":{"date-parts":[[2017,11,27]],"date-time":"2017-11-27T14:10:26Z","timestamp":1511791826000},"page":"336-346","source":"Crossref","is-referenced-by-count":80,"title":["Identity and Diversity of Human Peripheral Th and T Regulatory Cells Defined by Single-Cell Mass Cytometry"],"prefix":"10.1093","volume":"200","author":[{"given":"Matthew A","family":"Kunicki","sequence":"first","affiliation":[{"name":"Division of Stem Cell Transplantation and Regenerative Medicine, Department of Pediatrics, Stanford University , Stanford, CA 94305"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Laura C","family":"Amaya Hernandez","sequence":"additional","affiliation":[{"name":"Division of Stem Cell Transplantation and Regenerative Medicine, Department of Pediatrics, Stanford University , Stanford, CA 94305"},{"name":"Institute for Stem Cell Biology and Regenerative Medicine, Stanford University , Stanford, CA 94305"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kara L","family":"Davis","sequence":"additional","affiliation":[{"name":"Division of Hematology\/Oncology, Department of Pediatrics, Stanford University , Stanford, CA 94305"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rosa","family":"Bacchetta","sequence":"additional","affiliation":[{"name":"Division of Stem Cell Transplantation and Regenerative Medicine, Department of Pediatrics, Stanford University , Stanford, CA 94305"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria-Grazia","family":"Roncarolo","sequence":"additional","affiliation":[{"name":"Division of Stem Cell Transplantation and Regenerative Medicine, Department of Pediatrics, Stanford University , Stanford, CA 94305"},{"name":"Institute for Stem Cell Biology and Regenerative Medicine, Stanford University , Stanford, CA 94305"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2018,1,1]]},"reference":[{"key":"2025030610003556100_r1","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1146\/annurev.iy.07.040189.001045","article-title":"TH1 and TH2 cells: different patterns of lymphokine secretion lead to different functional properties","volume":"7","author":"Mosmann","year":"1989","journal-title":"Annu. Rev. Immunol."},{"key":"2025030610003556100_r2","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1016\/j.coi.2015.08.001","article-title":"Effector T cell differentiation: are master regulators of effector T cells still the masters?","volume":"37","author":"Wang","year":"2015","journal-title":"Curr. Opin. Immunol."},{"key":"2025030610003556100_r3","doi-asserted-by":"crossref","first-page":"252","DOI":"10.1016\/j.smim.2013.10.012","article-title":"The CD4-centered universe of human T cell subsets","volume":"25","author":"Geginat","year":"2013","journal-title":"Semin. Immunol."},{"key":"2025030610003556100_r4","doi-asserted-by":"crossref","first-page":"391","DOI":"10.1111\/j.1600-065X.2008.00634.x","article-title":"CD4+ T-regulatory cells: toward therapy for human diseases","volume":"223","author":"Allan","year":"2008","journal-title":"Immunol. Rev."},{"key":"2025030610003556100_r5","doi-asserted-by":"crossref","first-page":"64","DOI":"10.1038\/icb.2013.55","article-title":"Human T follicular helper (Tfh) cells and disease","volume":"92","author":"Ma","year":"2014","journal-title":"Immunol. Cell Biol."},{"key":"2025030610003556100_r6","doi-asserted-by":"crossref","first-page":"737","DOI":"10.1038\/39614","article-title":"A CD4+ T-cell subset inhibits antigen-specific T-cell responses and prevents colitis","volume":"389","author":"Groux","year":"1997","journal-title":"Nature"},{"key":"2025030610003556100_r7","doi-asserted-by":"crossref","first-page":"1295","DOI":"10.1084\/jem.193.11.1295","article-title":"Human Cd25(+)Cd4(+) T regulatory cells suppress naive and memory T cell proliferation and can be expanded in vitro without loss of function","volume":"193","author":"Levings","year":"2001","journal-title":"J. Exp. Med."},{"key":"2025030610003556100_r8","doi-asserted-by":"crossref","first-page":"330","DOI":"10.1038\/ni904","article-title":"Foxp3 programs the development and function of CD4+CD25+ regulatory T cells","volume":"4","author":"Fontenot","year":"2003","journal-title":"Nat. Immunol."},{"key":"2025030610003556100_r9","doi-asserted-by":"crossref","first-page":"610","DOI":"10.1056\/NEJMra052723","article-title":"The many roles of chemokines and chemokine receptors in inflammation","volume":"354","author":"Charo","year":"2006","journal-title":"N. Engl. J. Med."},{"key":"2025030610003556100_r10","doi-asserted-by":"crossref","first-page":"542","DOI":"10.1016\/j.immuni.2012.03.014","article-title":"JAK and STAT signaling molecules in immunoregulation and immune-mediated disease","volume":"36","author":"O\u2019Shea","year":"2012","journal-title":"Immunity"},{"key":"2025030610003556100_r11","doi-asserted-by":"crossref","first-page":"524","DOI":"10.1038\/nri.2016.75","article-title":"Immunity by equilibrium","volume":"16","author":"Eberl","year":"2016","journal-title":"Nat. Rev. Immunol."},{"key":"2025030610003556100_r12","doi-asserted-by":"crossref","first-page":"811","DOI":"10.1038\/nri2654","article-title":"The functional plasticity of T cell subsets","volume":"9","author":"Bluestone","year":"2009","journal-title":"Nat. Rev. Immunol."},{"key":"2025030610003556100_r13","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-662-09709-0","volume-title":"Redirection of Th1 and Th2 Responses","author":"Coffman","year":"1999","edition":"1st"},{"key":"2025030610003556100_r14","doi-asserted-by":"crossref","first-page":"1331","DOI":"10.1172\/JCI13543","article-title":"Rules of chemokine receptor association with T cell polarization in vivo","volume":"108","author":"Kim","year":"2001","journal-title":"J. Clin. Invest."},{"key":"2025030610003556100_r15","doi-asserted-by":"crossref","first-page":"436","DOI":"10.1016\/j.it.2014.06.002","article-title":"Phenotype and functions of memory Tfh cells in human blood","volume":"35","author":"Schmitt","year":"2014","journal-title":"Trends Immunol."},{"key":"2025030610003556100_r16","doi-asserted-by":"crossref","first-page":"725","DOI":"10.1084\/jem.20040774","article-title":"Chemokine receptor expression identifies Pre-T helper (Th)1, Pre-Th2, and nonpolarized cells among human CD4+ central memory T cells","volume":"200","author":"Rivino","year":"2004","journal-title":"J. Exp. Med."},{"key":"2025030610003556100_r17","doi-asserted-by":"crossref","first-page":"856","DOI":"10.1038\/ni.2947","article-title":"The cytokine TGF-\u03b2 co-opts signaling via STAT3-STAT4 to promote the differentiation of human TFH cells","volume":"15","author":"Schmitt","year":"2014","journal-title":"Nat. Immunol."},{"key":"2025030610003556100_r18","doi-asserted-by":"crossref","first-page":"1903","DOI":"10.1084\/jem.20080397","article-title":"Human interleukin 17\u2013producing cells originate from a CD161+CD4+ T cell precursor","volume":"205","author":"Cosmi","year":"2008","journal-title":"J. Exp. Med."},{"key":"2025030610003556100_r19","doi-asserted-by":"crossref","first-page":"1849","DOI":"10.1084\/jem.20070663","article-title":"Phenotypic and functional features of human Th17 cells","volume":"204","author":"Annunziato","year":"2007","journal-title":"J. Exp. Med."},{"key":"2025030610003556100_r20","doi-asserted-by":"crossref","first-page":"1121","DOI":"10.1016\/j.cell.2006.07.035","article-title":"The orphan nuclear receptor RORgammat directs the differentiation program of proinflammatory IL-17+ T helper cells","volume":"126","author":"Ivanov","year":"2006","journal-title":"Cell"},{"key":"2025030610003556100_r21","doi-asserted-by":"crossref","first-page":"1553","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":"2025030610003556100_r22","doi-asserted-by":"crossref","first-page":"2340","DOI":"10.4049\/jimmunol.175.4.2340","article-title":"The role of ICOS in the CXCR5+ follicular B helper T cell maintenance in vivo","volume":"175","author":"Akiba","year":"2005","journal-title":"J. Immunol."},{"key":"2025030610003556100_r23","doi-asserted-by":"crossref","first-page":"494","DOI":"10.1038\/nm.3109","article-title":"Inadequate T follicular cell help impairs B cell immunity during HIV infection","volume":"19","author":"Cubas","year":"2013","journal-title":"Nat. Med."},{"key":"2025030610003556100_r24","doi-asserted-by":"crossref","first-page":"1701","DOI":"10.1084\/jem.20060772","article-title":"CD127 expression inversely correlates with FoxP3 and suppressive function of human CD4+ Treg cells","volume":"203","author":"Liu","year":"2006","journal-title":"J. Exp. Med."},{"key":"2025030610003556100_r25","doi-asserted-by":"crossref","first-page":"4368","DOI":"10.1182\/blood-2006-11-055756","article-title":"Nonredundant roles for Stat5a\/b in directly regulating Foxp3","volume":"109","author":"Yao","year":"2007","journal-title":"Blood"},{"key":"2025030610003556100_r26","doi-asserted-by":"crossref","first-page":"739","DOI":"10.1038\/nm.3179","article-title":"Coexpression of CD49b and LAG-3 identifies human and mouse T regulatory type 1 cells","volume":"19","author":"Gagliani","year":"2013","journal-title":"Nat. Med."},{"key":"2025030610003556100_r27","doi-asserted-by":"crossref","first-page":"1652","DOI":"10.1002\/eji.201041120","article-title":"Killing of myeloid APCs via HLA class I, CD2 and CD226 defines a novel mechanism of suppression by human Tr1 cells","volume":"41","author":"Magnani","year":"2011","journal-title":"Eur. J. Immunol."},{"key":"2025030610003556100_r28","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1038\/nri.2015.18","article-title":"Harnessing the plasticity of CD4(+) T cells to treat immune-mediated disease","volume":"16","author":"DuPage","year":"2016","journal-title":"Nat. Rev. Immunol."},{"key":"2025030610003556100_r29","doi-asserted-by":"crossref","first-page":"593","DOI":"10.3389\/fimmu.2015.00593","article-title":"Clinical outlook for type-1 and FOXP3(+) T regulatory cell-based therapy","volume":"6","author":"Gregori","year":"2015","journal-title":"Front. Immunol."},{"key":"2025030610003556100_r30","doi-asserted-by":"crossref","first-page":"780","DOI":"10.1016\/j.cell.2016.04.019","article-title":"Mass cytometry: single cells, many features","volume":"165","author":"Spitzer","year":"2016","journal-title":"Cell"},{"key":"2025030610003556100_r31","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1016\/j.cell.2015.05.047","article-title":"Data-driven phenotypic dissection of AML reveals progenitor-like cells that correlate with prognosis","volume":"162","author":"Levine","year":"2015","journal-title":"Cell"},{"key":"2025030610003556100_r32","doi-asserted-by":"crossref","first-page":"1259425","DOI":"10.1126\/science.1259425","article-title":"IMMUNOLOGY. An interactive reference framework for modeling a dynamic immune system","volume":"349","author":"Spitzer","year":"2015","journal-title":"Science"},{"key":"2025030610003556100_r33","doi-asserted-by":"crossref","first-page":"687","DOI":"10.1126\/science.1198704","article-title":"Single-cell mass cytometry of differential immune and drug responses across a human hematopoietic continuum","volume":"332","author":"Bendall","year":"2011","journal-title":"Science"},{"key":"2025030610003556100_r34","doi-asserted-by":"crossref","first-page":"1822","DOI":"10.1016\/j.celrep.2015.05.022","article-title":"Mapping the diversity of follicular helper T cells in human blood and tonsils using high-dimensional mass cytometry analysis","volume":"11","author":"Wong","year":"2015","journal-title":"Cell Reports"},{"key":"2025030610003556100_r35","doi-asserted-by":"crossref","first-page":"2030","DOI":"10.4049\/jimmunol.1500703","article-title":"Phenotypic complexity of the human regulatory T cell compartment revealed by mass cytometry","volume":"195","author":"Mason","year":"2015","journal-title":"J. Immunol."},{"key":"2025030610003556100_r36","doi-asserted-by":"crossref","first-page":"255ra131","DOI":"10.1126\/scitranslmed.3009701","article-title":"Clinical recovery from surgery correlates with single-cell immune signatures","volume":"6","author":"Gaudilli\u00e8re","year":"2014","journal-title":"Sci. Transl. Med."},{"key":"2025030610003556100_r37","doi-asserted-by":"crossref","first-page":"1193","DOI":"10.1182\/blood-2016-03-703702","article-title":"Deep phenotyping of Tregs identifies an immune signature for idiopathic aplastic anemia and predicts response to treatment","volume":"128","author":"Kordasti","year":"2016","journal-title":"Blood"},{"key":"2025030610003556100_r38","doi-asserted-by":"crossref","first-page":"858","DOI":"10.1038\/nbt.2317","article-title":"Multiplexed mass cytometry profiling of cellular states perturbed by small-molecule regulators","volume":"30","author":"Bodenmiller","year":"2012","journal-title":"Nat. Biotechnol."},{"key":"2025030610003556100_r39","first-page":"Unit 10.17","article-title":"Web-based analysis and publication of flow cytometry experiments","volume":"Chapter 10","author":"Kotecha","year":"2010","journal-title":"Curr. Protoc. Cytom."},{"key":"2025030610003556100_r40","doi-asserted-by":"crossref","first-page":"2022","DOI":"10.4049\/jimmunol.1402661","article-title":"Barcoding of live human peripheral blood mononuclear cells for multiplexed mass cytometry","volume":"194","author":"Mei","year":"2015","journal-title":"J. Immunol."},{"key":"2025030610003556100_r41","doi-asserted-by":"crossref","first-page":"545","DOI":"10.1038\/nbt.2594","article-title":"viSNE enables visualization of high dimensional single-cell data and reveals phenotypic heterogeneity of leukemia","volume":"31","author":"Amir","year":"2013","journal-title":"Nat. Biotechnol."},{"key":"2025030610003556100_r42","doi-asserted-by":"crossref","first-page":"636","DOI":"10.1002\/cyto.a.22625","article-title":"FlowSOM: using self-organizing maps for visualization and interpretation of cytometry data","volume":"87","author":"Van Gassen","year":"2015","journal-title":"Cytometry A"},{"key":"2025030610003556100_r43","doi-asserted-by":"crossref","first-page":"310","DOI":"10.1164\/rccm.200502-244OC","article-title":"Role of the chemokine receptors CXCR3 and CCR4 in human pulmonary fibrosis","volume":"173","author":"Pignatti","year":"2006","journal-title":"Am. J. Respir. Crit. Care Med."},{"key":"2025030610003556100_r44","doi-asserted-by":"crossref","first-page":"31","DOI":"10.4103\/2277-9175.108770","article-title":"CCR3, CCR4, CCR5, and CXCR3 expression in peripheral blood CD4+ lymphocytes in gastric cancer patients","volume":"2","author":"Andalib","year":"2013","journal-title":"Adv. Biomed. Res."},{"key":"2025030610003556100_r45","doi-asserted-by":"crossref","first-page":"1373","DOI":"10.1038\/jid.2009.399","article-title":"Circulating Th17, Th22, and Th1 cells are increased in psoriasis","volume":"130","author":"Kagami","year":"2010","journal-title":"J. Invest. Dermatol."},{"key":"2025030610003556100_r46","doi-asserted-by":"crossref","first-page":"440","DOI":"10.1111\/j.1365-2249.2008.03657.x","article-title":"Elevated CCR6+ CD4+ T lymphocytes in tissue compared with blood and induction of CCL20 during the asthmatic late response","volume":"152","author":"Francis","year":"2008","journal-title":"Clin. Exp. Immunol."},{"key":"2025030610003556100_r47","doi-asserted-by":"crossref","first-page":"186","DOI":"10.4049\/jimmunol.162.1.186","article-title":"CC-chemokine receptor 6 is expressed on diverse memory subsets of T cells and determines responsiveness to macrophage inflammatory protein 3 alpha","volume":"162","author":"Liao","year":"1999","journal-title":"J. Immunol."},{"key":"2025030610003556100_r48","doi-asserted-by":"crossref","first-page":"211","DOI":"10.4049\/jimmunol.176.1.211","article-title":"Expression of CD161 (NKR-P1A) defines subsets of human CD4 and CD8 T cells with different functional activities","volume":"176","author":"Takahashi","year":"2006","journal-title":"J. Immunol."},{"key":"2025030610003556100_r49","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1016\/j.jaut.2011.12.009","article-title":"Demethylation analysis of the FOXP3 locus shows quantitative defects of regulatory T cells in IPEX-like syndrome","volume":"38","author":"Barzaghi","year":"2012","journal-title":"J. Autoimmun."},{"key":"2025030610003556100_r50","doi-asserted-by":"crossref","first-page":"830","DOI":"10.1182\/blood.V84.3.830.bloodjournal843830","article-title":"Expression of the G-protein--coupled receptor BLR1 defines mature, recirculating B cells and a subset of T-helper memory cells","volume":"84","author":"F\u00f6rster","year":"1994","journal-title":"Blood"},{"key":"2025030610003556100_r51","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1016\/j.immuni.2011.08.012","article-title":"An essential role of the transcription factor GATA-3 for the function of regulatory T cells","volume":"35","author":"Wang","year":"2011","journal-title":"Immunity"},{"key":"2025030610003556100_r52","doi-asserted-by":"crossref","first-page":"1010","DOI":"10.1038\/ni.2402","article-title":"Transcription factor Foxp3 and its protein partners form a complex regulatory network","volume":"13","author":"Rudra","year":"2012","journal-title":"Nat. Immunol."},{"key":"2025030610003556100_r53","doi-asserted-by":"crossref","first-page":"4793","DOI":"10.1073\/pnas.0900408106","article-title":"Identification of IL-17-producing FOXP3+ regulatory T cells in humans","volume":"106","author":"Voo","year":"2009","journal-title":"Proc Natl Acad Sci U S A."},{"key":"2025030610003556100_r54","doi-asserted-by":"crossref","first-page":"4240","DOI":"10.1182\/blood-2008-10-183251","article-title":"IL-17\u2013producing human peripheral regulatory T cells retain suppressive function","volume":"113","author":"Beriou","year":"2009","journal-title":"Blood"},{"key":"2025030610003556100_r55","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1016\/j.immuni.2008.12.009","article-title":"Global mapping of H3K4me3 and H3K27me3 reveals specificity and plasticity in lineage fate determination of differentiating CD4+ T cells","volume":"30","author":"Wei","year":"2009","journal-title":"Immunity"},{"key":"2025030610003556100_r56","doi-asserted-by":"crossref","first-page":"2031","DOI":"10.1053\/j.gastro.2011.03.009","article-title":"IL-12 converts Foxp3+ regulatory T cells to Foxp3+IFN-\u03b3+ T cells with inhibitory functions during induction of colitis","volume":"140","author":"Feng","year":"2011","journal-title":"Gastroenterology"},{"key":"2025030610003556100_r57","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1016\/j.immuni.2008.11.005","article-title":"Late developmental plasticity in the T helper 17 lineage","volume":"30","author":"Lee","year":"2009","journal-title":"Immunity"},{"key":"2025030610003556100_r58","doi-asserted-by":"crossref","first-page":"2402","DOI":"10.1182\/blood-2012-09-378653","article-title":"Essentials of Th17 cell commitment and plasticity","volume":"121","author":"Muranski","year":"2013","journal-title":"Blood"},{"key":"2025030610003556100_r59","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1016\/j.immuni.2010.12.012","article-title":"Human blood CXCR5(+)CD4(+) T cells are counterparts of T follicular cells and contain specific subsets that differentially support antibody secretion","volume":"34","author":"Morita","year":"2011","journal-title":"Immunity"},{"key":"2025030610003556100_r60","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1167\/iovs.08-2536","article-title":"Human corneal endothelial cells expressing programmed death-ligand 1 (PD-L1) suppress PD-1+ T helper 1 cells by a contact-dependent mechanism","volume":"50","author":"Sugita","year":"2009","journal-title":"Invest. Ophthalmol. Vis. Sci."},{"key":"2025030610003556100_r61","doi-asserted-by":"crossref","first-page":"508","DOI":"10.1158\/0008-5472.CAN-14-1215","article-title":"PD-1\/SHP-2 inhibits Tc1\/Th1 phenotypic responses and the activation of T cells in the tumor microenvironment","volume":"75","author":"Li","year":"2015","journal-title":"Cancer Res."},{"key":"2025030610003556100_r62","doi-asserted-by":"crossref","first-page":"3673","DOI":"10.4049\/jimmunol.1300945","article-title":"The CD226\/CD155 interaction regulates the proinflammatory (Th1\/Th17)\/anti-inflammatory (Th2) balance in humans","volume":"191","author":"Lozano","year":"2013","journal-title":"J. Immunol."},{"key":"2025030610003556100_r63","doi-asserted-by":"crossref","first-page":"3160","DOI":"10.1002\/eji.200939470","article-title":"Abundance of follicular helper T cells in Peyer\u2019s patches is modulated by CD155","volume":"39","author":"Seth","year":"2009","journal-title":"Eur. J. Immunol."},{"key":"2025030610003556100_r64","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1084\/jem.192.2.303","article-title":"Immunologic self-tolerance maintained by CD25(+)CD4(+) regulatory T cells constitutively expressing cytotoxic T lymphocyte-associated antigen 4","volume":"192","author":"Takahashi","year":"2000","journal-title":"J. Exp. Med."},{"key":"2025030610003556100_r65","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1038\/ni.1674","article-title":"The surface protein TIGIT suppresses T cell activation by promoting the generation of mature immunoregulatory dendritic cells","volume":"10","author":"Yu","year":"2009","journal-title":"Nat. Immunol."},{"key":"2025030610003556100_r66","doi-asserted-by":"crossref","first-page":"1778","DOI":"10.1038\/mt.2012.71","article-title":"Enforced IL-10 expression confers type 1 regulatory T cell (Tr1) phenotype and function to human CD4(+) T cells","volume":"20","author":"Andolfi","year":"2012","journal-title":"Mol. Ther."},{"key":"2025030610003556100_r67","doi-asserted-by":"crossref","first-page":"4430","DOI":"10.1182\/blood-2011-11-392324","article-title":"Functionally distinct subsets of human FOXP3+ Treg cells that phenotypically mirror effector Th cells","volume":"119","author":"Duhen","year":"2012","journal-title":"Blood"},{"key":"2025030610003556100_r68","doi-asserted-by":"crossref","first-page":"899","DOI":"10.1016\/j.immuni.2009.03.019","article-title":"Functional delineation and differentiation dynamics of human CD4+ T cells expressing the FoxP3 transcription factor","volume":"30","author":"Miyara","year":"2009","journal-title":"Immunity"},{"key":"2025030610003556100_r69","doi-asserted-by":"crossref","first-page":"639","DOI":"10.1038\/ni1467","article-title":"Surface phenotype and antigenic specificity of human interleukin 17-producing T helper memory cells","volume":"8","author":"Acosta-Rodriguez","year":"2007","journal-title":"Nat. Immunol."},{"key":"2025030610003556100_r70","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1084\/jem.20130301","article-title":"Pro-inflammatory human Th17 cells selectively express P-glycoprotein and are refractory to glucocorticoids","volume":"211","author":"Ramesh","year":"2014","journal-title":"J. Exp. Med."},{"key":"2025030610003556100_r71","doi-asserted-by":"crossref","first-page":"4488","DOI":"10.4049\/jimmunol.177.7.4488","article-title":"The majority of human peripheral blood CD4+CD25highFoxp3+ regulatory T cells bear functional skin-homing receptors","volume":"177","author":"Hirahara","year":"2006","journal-title":"J. Immunol."},{"key":"2025030610003556100_r72","doi-asserted-by":"crossref","first-page":"6329","DOI":"10.1038\/ncomms7329","article-title":"TGF-\u03b23-expressing CD4+CD25(\u2212)LAG3+ regulatory T cells control humoral immune responses","volume":"6","author":"Okamura","year":"2015","journal-title":"Nat. Commun."}],"container-title":["The Journal of Immunology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/jimmunol\/article-pdf\/200\/1\/336\/61511943\/ji1701025.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jimmunol\/article-pdf\/200\/1\/336\/61511943\/ji1701025.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,3,6]],"date-time":"2025-03-06T05:02:53Z","timestamp":1741237373000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/jimmunol\/article\/200\/1\/336\/7962160"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2018,1,1]]},"references-count":72,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2018,1,1]]}},"URL":"https:\/\/doi.org\/10.4049\/jimmunol.1701025","relation":{},"ISSN":["0022-1767","1550-6606"],"issn-type":[{"value":"0022-1767","type":"print"},{"value":"1550-6606","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2018,1]]},"published":{"date-parts":[[2018,1,1]]}}}