{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T20:28:01Z","timestamp":1791836881608,"version":"4.3.4"},"reference-count":95,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2021,8,18]],"date-time":"2021-08-18T00:00:00Z","timestamp":1629244800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["81971495"],"award-info":[{"award-number":["81971495"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["81600450"],"award-info":[{"award-number":["81600450"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["82071798"],"award-info":[{"award-number":["82071798"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["91442117"],"award-info":[{"award-number":["91442117"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100010014","name":"Six Talent Peaks Project in Jiangsu Province","doi-asserted-by":"publisher","award":["No.2018-WSN-011"],"award-info":[{"award-number":["No.2018-WSN-011"]}],"id":[{"id":"10.13039\/501100010014","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100017330","name":"Foundation of Jiangsu Collaborative Innovation Center of Biomedical Functional Materials","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100017330","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell Dev. Biol."],"abstract":"<jats:p>Regulatory T cells (Tregs) specifically expressing Forkhead box P3 (Foxp3) play roles in suppressing the immune response and maintaining immune homeostasis. After maturation in the thymus, Tregs leave the thymus and migrate to lymphoid tissues or non-lymphoid tissues. Increasing evidence indicates that Tregs with unique characteristics also have significant effects on non-lymphoid peripheral tissues. Tissue-resident Tregs, also called tissue Tregs, do not recirculate in the blood or lymphatics and attain a unique phenotype distinct from common Tregs in circulation. This review first summarizes the phenotype, function, and cytokine expression of these Tregs in visceral adipose tissue, skin, muscle, and other tissues. Then, how Tregs are generated, home, and are attracted to and remain resident in the tissue are discussed. Finally, how an increased understanding of these tissue Tregs might guide clinical treatment is discussed.<\/jats:p>","DOI":"10.3389\/fcell.2021.717903","type":"journal-article","created":{"date-parts":[[2021,8,18]],"date-time":"2021-08-18T05:15:52Z","timestamp":1629263752000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":47,"title":["Tissue Tregs and Maintenance of Tissue Homeostasis"],"prefix":"10.3389","volume":"9","author":[{"given":"Qing","family":"Shao","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jian","family":"Gu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jinren","family":"Zhou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qi","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiangyu","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhenhua","family":"Deng","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ling","family":"Lu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2021,8,18]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"1272","DOI":"10.1038\/nm.3962","article-title":"Hair follicle-derived IL-7 and IL-15 mediate skin-resident memory T cell homeostasis and lymphoma.","volume":"21","author":"Adachi","year":"2015","journal-title":"Nat. Med."},{"key":"B2","doi-asserted-by":"publisher","first-page":"372","DOI":"10.1111\/imm.12791","article-title":"Regulatory T cells in skin.","volume":"152","author":"Ali","year":"2017","journal-title":"Immunology"},{"key":"B3","doi-asserted-by":"publisher","first-page":"1119.e1111","DOI":"10.1016\/j.cell.2017.05.002","article-title":"Regulatory T cells in skin facilitate epithelial stem cell differentiation.","volume":"169","author":"Ali","year":"2017","journal-title":"Cell"},{"key":"B4","doi-asserted-by":"publisher","first-page":"1078","DOI":"10.1016\/j.cell.2015.08.021","article-title":"A distinct function of regulatory T cells in tissue protection.","volume":"162","author":"Arpaia","year":"2015","journal-title":"Cell"},{"key":"B5","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1038\/nature16151","article-title":"Depletion of fat-resident Treg cells prevents age-associated insulin resistance.","volume":"528","author":"Bapat","year":"2015","journal-title":"Nature"},{"key":"B6","doi-asserted-by":"publisher","first-page":"eaaz9631","DOI":"10.1126\/sciimmunol.aaz9631","article-title":"Regulatory T cells in skin injury: at the crossroads of tolerance and tissue repair.","volume":"5","author":"Boothby","year":"2020","journal-title":"Sci. Immunol."},{"key":"B7","doi-asserted-by":"publisher","first-page":"268","DOI":"10.1002\/art.27183","article-title":"Muscle resident macrophages control the immune cell reaction in a mouse model of notexin-induced myoinjury.","volume":"62","author":"Brigitte","year":"2010","journal-title":"Arthritis Rheum"},{"key":"B8","doi-asserted-by":"publisher","first-page":"1282","DOI":"10.1016\/j.cell.2013.10.054","article-title":"A special population of regulatory T cells potentiates muscle repair.","volume":"155","author":"Burzyn","year":"2013","journal-title":"Cell"},{"key":"B9","doi-asserted-by":"publisher","first-page":"748","DOI":"10.1001\/jamadermatol.2014.504","article-title":"Effects of low-dose recombinant interleukin 2 to promote T-regulatory cells in alopecia areata.","volume":"150","author":"Castela","year":"2014","journal-title":"JAMA Dermatol."},{"key":"B10","doi-asserted-by":"publisher","first-page":"482","DOI":"10.1073\/pnas.1423486112","article-title":"Appearance and disappearance of the mRNA signature characteristic of Treg cells in visceral adipose tissue: age, diet, and PPARgamma effects.","volume":"112","author":"Cipolletta","year":"2015","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B11","doi-asserted-by":"publisher","first-page":"549","DOI":"10.1038\/nature11132","article-title":"PPAR-gamma is a major driver of the accumulation and phenotype of adipose tissue Treg cells.","volume":"486","author":"Cipolletta","year":"2012","journal-title":"Nature"},{"key":"B12","doi-asserted-by":"publisher","first-page":"194","DOI":"10.1182\/blood-2006-02-002873","article-title":"IL-15 and dermal fibroblasts induce proliferation of natural regulatory T cells isolated from human skin.","volume":"109","author":"Clark","year":"2007","journal-title":"Blood"},{"key":"B13","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1016\/j.it.2012.11.002","article-title":"Differentiation and function of Foxp3(+) effector regulatory T cells.","volume":"34","author":"Cretney","year":"2013","journal-title":"Trends Immunol."},{"key":"B14","doi-asserted-by":"publisher","first-page":"304","DOI":"10.1038\/ni.2006","article-title":"The transcription factors Blimp-1 and IRF4 jointly control the differentiation and function of effector regulatory T cells.","volume":"12","author":"Cretney","year":"2011","journal-title":"Nat. Immunol."},{"key":"B15","doi-asserted-by":"publisher","first-page":"4530","DOI":"10.4049\/jimmunol.1002031","article-title":"The link between the PDL1 costimulatory pathway and Th17 in fetomaternal tolerance.","volume":"187","author":"D\u2019Addio","year":"2011","journal-title":"J. Immunol."},{"key":"B16","doi-asserted-by":"publisher","first-page":"295.e211","DOI":"10.1016\/j.immuni.2019.12.002","article-title":"Precursors for nonlymphoid-tissue Treg cells reside in secondary lymphoid organs and are programmed by the transcription factor BATF.","volume":"52","author":"Delacher","year":"2020","journal-title":"Immunity"},{"key":"B17","doi-asserted-by":"publisher","first-page":"1160","DOI":"10.1038\/ni.3799","article-title":"Genome-wide DNA-methylation landscape defines specialization of regulatory T cells in tissues.","volume":"18","author":"Delacher","year":"2017","journal-title":"Nat. Immunol."},{"key":"B18","doi-asserted-by":"publisher","first-page":"702.e717","DOI":"10.1016\/j.immuni.2021.03.007","article-title":"Single-cell chromatin accessibility landscape identifies tissue repair program in human regulatory T cells.","volume":"54","author":"Delacher","year":"2021","journal-title":"Immunity"},{"key":"B19","doi-asserted-by":"publisher","first-page":"eaat5861","DOI":"10.1126\/sciimmunol.aat5861","article-title":"Molecular diversification of regulatory T cells in nonlymphoid tissues.","volume":"3","author":"DiSpirito","year":"2018","journal-title":"Sci. Immunol."},{"key":"B20","doi-asserted-by":"publisher","first-page":"674","DOI":"10.1038\/nn.4528","article-title":"Regulatory T cells promote myelin regeneration in the central nervous system.","volume":"20","author":"Dombrowski","year":"2017","journal-title":"Nat. Neurosci."},{"key":"B21","doi-asserted-by":"publisher","first-page":"2954","DOI":"10.2337\/db11-0358","article-title":"Potential role of regulatory T cells in reversing obesity-linked insulin resistance and diabetic nephropathy.","volume":"60","author":"Eller","year":"2011","journal-title":"Diabetes"},{"key":"B22","doi-asserted-by":"publisher","first-page":"387","DOI":"10.1146\/annurev-immunol-032712-100003","article-title":"Immunology of the maternal-fetal interface.","volume":"31","author":"Erlebacher","year":"2013","journal-title":"Annu. Rev. Immunol."},{"key":"B23","doi-asserted-by":"publisher","first-page":"e58463","DOI":"10.7554\/eLife.58463","article-title":"Discovery of surrogate agonists for visceral fat Treg cells that modulate metabolic indices in vivo.","volume":"9","author":"Fernandes","year":"2020","journal-title":"Elife"},{"key":"B24","doi-asserted-by":"publisher","first-page":"930","DOI":"10.1038\/nm.2002","article-title":"Lean, but not obese, fat is enriched for a unique population of regulatory T cells that affect metabolic parameters.","volume":"15","author":"Feuerer","year":"2009","journal-title":"Nat. Med."},{"key":"B25","doi-asserted-by":"publisher","first-page":"703","DOI":"10.1016\/j.neuron.2015.01.013","article-title":"The glia-derived alarmin IL-33 orchestrates the immune response and promotes recovery following CNS injury.","volume":"85","author":"Gadani","year":"2015","journal-title":"Neuron"},{"key":"B26","doi-asserted-by":"publisher","first-page":"1854.e1855","DOI":"10.1016\/j.celrep.2019.01.070","article-title":"Blimp1 prevents methylation of Foxp3 and loss of regulatory T cell identity at sites of inflammation.","volume":"26","author":"Garg","year":"2019","journal-title":"Cell Rep."},{"key":"B27","doi-asserted-by":"publisher","first-page":"3349","DOI":"10.1016\/j.ajpath.2015.08.010","article-title":"Enhancement of muscle T regulatory cells and improvement of muscular dystrophic process in mdx mice by blockade of extracellular ATP\/P2X Axis.","volume":"185","author":"Gazzerro","year":"2015","journal-title":"Am. J. Pathol."},{"key":"B28","doi-asserted-by":"publisher","first-page":"107874","DOI":"10.1016\/j.celrep.2020.107874","article-title":"Regulatory T cells play a role in a subset of idiopathic preterm labor\/birth and adverse neonatal outcomes.","volume":"32","author":"Gomez-Lopez","year":"2020","journal-title":"Cell Rep."},{"key":"B29","doi-asserted-by":"publisher","first-page":"4483","DOI":"10.4049\/jimmunol.1300212","article-title":"Cutting Edge: memory regulatory t cells require IL-7 and not IL-2 for their maintenance in peripheral tissues.","volume":"190","author":"Gratz","year":"2013","journal-title":"J. Immunol."},{"key":"B30","doi-asserted-by":"publisher","first-page":"106027","DOI":"10.1016\/j.intimp.2019.106027","article-title":"Brain Foxp3(+) regulatory T cells can be expanded by Interleukin-33 in mouse ischemic stroke.","volume":"81","author":"Guo","year":"2020","journal-title":"Int. Immunopharmacol."},{"key":"B31","doi-asserted-by":"publisher","first-page":"eabb8620","DOI":"10.1126\/sciimmunol.abb8620","article-title":"Sex drives Tregs into fat.","volume":"5","author":"Habrylo","year":"2020","journal-title":"Sci. Immunol."},{"key":"B32","doi-asserted-by":"publisher","first-page":"4777","DOI":"10.4049\/jimmunol.1500020","article-title":"IL-33 reverses an obesity-induced deficit in visceral adipose tissue ST2+ T regulatory cells and ameliorates adipose tissue inflammation and insulin resistance.","volume":"194","author":"Han","year":"2015","journal-title":"J. Immunol."},{"key":"B33","doi-asserted-by":"publisher","first-page":"eaat6280","DOI":"10.1126\/science.aat6280","article-title":"Commensal-specific T cell plasticity promotes rapid tissue adaptation to injury.","volume":"363","author":"Harrison","year":"2019","journal-title":"Science"},{"key":"B34","doi-asserted-by":"publisher","first-page":"1098","DOI":"10.3389\/fimmu.2019.01098","article-title":"Functional phenotypic diversity of regulatory T cells remaining in inflamed skin.","volume":"10","author":"Ikebuchi","year":"2019","journal-title":"Front. Immunol."},{"key":"B35","doi-asserted-by":"publisher","first-page":"9765","DOI":"10.1073\/pnas.0908771107","article-title":"Induction of regulatory T cells decreases adipose inflammation and alleviates insulin resistance in ob\/ob mice.","volume":"107","author":"Ilan","year":"2010","journal-title":"Proc. Natl. Acad. Sci."},{"key":"B36","doi-asserted-by":"publisher","first-page":"246","DOI":"10.1038\/s41586-018-0824-5","article-title":"Brain regulatory T cells suppress astrogliosis and potentiate neurological recovery.","volume":"565","author":"Ito","year":"2019","journal-title":"Nature"},{"key":"B37","doi-asserted-by":"publisher","first-page":"531","DOI":"10.1146\/annurev.immunol.25.022106.141623","article-title":"Regulatory T cells: mechanisms of differentiation and function.","volume":"30","author":"Josefowicz","year":"2012","journal-title":"Annu. Rev. Immunol."},{"key":"B38","doi-asserted-by":"publisher","first-page":"eaaw2910","DOI":"10.1126\/sciimmunol.aaw2910","article-title":"Regulatory T cells in skin are uniquely poised to suppress profibrotic immune responses.","volume":"4","author":"Kalekar","year":"2019","journal-title":"Sci. Immunol."},{"key":"B39","doi-asserted-by":"publisher","first-page":"1456","DOI":"10.1126\/science.1237013","article-title":"GPR15-mediated homing controls immune homeostasis in the large intestine mucosa.","volume":"340","author":"Kim","year":"2013","journal-title":"Science"},{"key":"B40","doi-asserted-by":"publisher","first-page":"543","DOI":"10.1016\/j.cmet.2015.03.005","article-title":"Antigen- and cytokine-driven accumulation of regulatory T cells in visceral adipose tissue of lean mice.","volume":"21","author":"Kolodin","year":"2015","journal-title":"Cell Metab."},{"key":"B41","doi-asserted-by":"publisher","first-page":"423","DOI":"10.1038\/nm1564","article-title":"Myelin-specific regulatory T cells accumulate in the CNS but fail to control autoimmune inflammation.","volume":"13","author":"Korn","year":"2007","journal-title":"Nat. Med."},{"key":"B42","doi-asserted-by":"publisher","first-page":"163","DOI":"10.1186\/s40478-014-0163-1","article-title":"Treg cells mediate recovery from EAE by controlling effector T cell proliferation and motility in the CNS.","volume":"2","author":"Koutrolos","year":"2014","journal-title":"Acta Neuropathol. Commun."},{"key":"B43","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1002\/mus.880160116","article-title":"Distinguishing unloading- versus reloading-induced changes in rat soleus muscle.","volume":"16","author":"Krippendorf","year":"1993","journal-title":"Muscle Nerve"},{"key":"B44","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1016\/j.immuni.2016.01.009","article-title":"Poor Repair of skeletal muscle in aging mice reflects a defect in local, interleukin-33-dependent accumulation of regulatory T cells.","volume":"44","author":"Kuswanto","year":"2016","journal-title":"Immunity"},{"key":"B45","doi-asserted-by":"publisher","first-page":"1835","DOI":"10.4049\/jimmunol.181.3.1835","article-title":"Functional regulatory T cells accumulate in aged hosts and promote chronic infectious disease reactivation.","volume":"181","author":"Lages","year":"2008","journal-title":"J. Immunol."},{"key":"B46","doi-asserted-by":"publisher","first-page":"285.e212","DOI":"10.1016\/j.cell.2018.05.004","article-title":"TCR transgenic mice reveal stepwise, multi-site acquisition of the distinctive Fat-Treg phenotype.","volume":"174","author":"Li","year":"2018","journal-title":"Cell"},{"key":"B47","doi-asserted-by":"publisher","first-page":"e2025197118","DOI":"10.1073\/pnas.2025197118","article-title":"PPARgamma marks splenic precursors of multiple nonlymphoid-tissue Treg compartments.","volume":"118","author":"Li","year":"2021","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B48","doi-asserted-by":"publisher","first-page":"613","DOI":"10.1038\/mi.2016.84","article-title":"Regulatory T cells decrease invariant natural killer T cell-mediated pregnancy loss in mice.","volume":"10","author":"Li","year":"2017","journal-title":"Mucosal Immunol."},{"key":"B49","doi-asserted-by":"publisher","first-page":"e129756","DOI":"10.1172\/jci.insight.129756","article-title":"Regulatory T cells use arginase 2 to enhance their metabolic fitness in tissues.","volume":"4","author":"Lowe","year":"2019","journal-title":"JCI Insight"},{"key":"B50","doi-asserted-by":"publisher","first-page":"527","DOI":"10.1016\/j.cmet.2015.03.001","article-title":"Regulation of obesity-related insulin resistance with gut anti-inflammatory agents.","volume":"21","author":"Luck","year":"2015","journal-title":"Cell Metab."},{"key":"B51","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1038\/ni.3047","article-title":"Regulatory iNKT cells lack expression of the transcription factor PLZF and control the homeostasis of T(reg) cells and macrophages in adipose tissue.","volume":"16","author":"Lynch","year":"2015","journal-title":"Nat. Immunol."},{"key":"B52","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1186\/s13024-020-00375-7","article-title":"Harnessing regulatory T cell neuroprotective activities for treatment of neurodegenerative disorders.","volume":"15","author":"Machhi","year":"2020","journal-title":"Mol. Neurodegener."},{"key":"B53","doi-asserted-by":"publisher","first-page":"eaao6923","DOI":"10.1126\/sciimmunol.aao6923","article-title":"RORalpha-expressing T regulatory cells restrain allergic skin inflammation.","volume":"3","author":"Malhotra","year":"2018","journal-title":"Sci. Immunol."},{"key":"B54","doi-asserted-by":"publisher","first-page":"R832","DOI":"10.1152\/ajpregu.00797.2009","article-title":"Regulation of skeletal muscle regeneration by CCR2-activating chemokines is directly related to macrophage recruitment.","volume":"299","author":"Martinez","year":"2010","journal-title":"Am. J. Physiol. Regul. Integr. Comp. Physiol."},{"key":"B55","doi-asserted-by":"publisher","first-page":"655.e","DOI":"10.1016\/j.immuni.2019.02.013","article-title":"Treg-cell control of a CXCL5-IL-17 inflammatory axis promotes hair-follicle-stem-cell differentiation during skin-barrier repair.","volume":"50","author":"Mathur","year":"2019","journal-title":"Immunity"},{"key":"B56","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1016\/j.immuni.2015.05.019","article-title":"Interleukin-33 and Interferon-gamma counter-regulate group 2 innate lymphoid cell activation during immune perturbation.","volume":"43","author":"Molofsky","year":"2015","journal-title":"Immunity"},{"key":"B57","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1038\/nri.2015.3","article-title":"Tissue-resident memory T cells: local specialists in immune defence.","volume":"16","author":"Mueller","year":"2015","journal-title":"Nat. Rev. Immunol."},{"key":"B58","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1046\/j.0022-202x.2001.01377.x","article-title":"A comprehensive guide for the accurate classification of murine hair follicles in distinct hair cycle stages.","volume":"117","author":"Muller-Rover","year":"2001","journal-title":"J. Invest. Dermatol."},{"key":"B59","doi-asserted-by":"publisher","first-page":"2010","DOI":"10.4049\/jimmunol.1502139","article-title":"Cutting edge: regulatory T cells facilitate cutaneous wound healing.","volume":"196","author":"Nosbaum","year":"2016","journal-title":"J. Immunol."},{"key":"B60","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1016\/j.celrep.2016.08.092","article-title":"Rapid and efficient generation of regulatory T cells to commensal antigens in the periphery.","volume":"17","author":"Nutsch","year":"2016","journal-title":"Cell Rep."},{"key":"B61","doi-asserted-by":"publisher","first-page":"989","DOI":"10.1126\/science.aac4263","article-title":"Mucosal immunology. The microbiota regulates type 2 immunity through RORgammat(+) T cells.","volume":"349","author":"Ohnmacht","year":"2015","journal-title":"Science"},{"key":"B62","doi-asserted-by":"publisher","first-page":"363","DOI":"10.1038\/nm.2627","article-title":"The cellular and signalling networks linking the immune system and metabolism in disease.","volume":"18","author":"Osborn","year":"2012","journal-title":"Nat. Med."},{"key":"B63","doi-asserted-by":"publisher","first-page":"442","DOI":"10.1038\/s41586-020-03071-0","article-title":"RANK links thymic regulatory T cells to fetal loss and gestational diabetes in pregnancy.","volume":"589","author":"Paolino","year":"2021","journal-title":"Nature"},{"key":"B64","doi-asserted-by":"publisher","first-page":"1268","DOI":"10.3389\/fimmu.2019.01268","article-title":"Thrombospondin-1\/CD47 interaction regulates Th17 and Treg differentiation in psoriasis.","volume":"10","author":"Rodriguez-Jimenez","year":"2019","journal-title":"Front. Immunol."},{"key":"B65","doi-asserted-by":"publisher","first-page":"20","DOI":"10.1016\/j.cell.2013.12.012","article-title":"What we talk about when we talk about fat.","volume":"156","author":"Rosen","year":"2014","journal-title":"Cell"},{"key":"B66","doi-asserted-by":"publisher","first-page":"2352","DOI":"10.1038\/s41467-019-10203-2","article-title":"Bcl10-controlled Malt1 paracaspase activity is key for the immune suppressive function of regulatory T cells.","volume":"10","author":"Rosenbaum","year":"2019","journal-title":"Nat. Commun."},{"key":"B67","doi-asserted-by":"publisher","first-page":"102","DOI":"10.1038\/nature11462","article-title":"Pregnancy imprints regulatory memory that sustains anergy to fetal antigen.","volume":"490","author":"Rowe","year":"2012","journal-title":"Nature"},{"key":"B68","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1016\/j.cell.2012.05.031","article-title":"Extrathymic generation of regulatory T cells in placental mammals mitigates maternal-fetal conflict.","volume":"150","author":"Samstein","year":"2012","journal-title":"Cell"},{"key":"B69","doi-asserted-by":"publisher","first-page":"1027","DOI":"10.1172\/JCI72932","article-title":"Memory regulatory T cells reside in human skin.","volume":"124","author":"Sanchez Rodriguez","year":"2014","journal-title":"J. Clin. Invest."},{"key":"B70","doi-asserted-by":"publisher","first-page":"467.e465","DOI":"10.1016\/j.chom.2017.03.001","article-title":"Commensal microbes and hair follicle morphogenesis coordinately drive treg migration into neonatal skin.","volume":"21","author":"Scharschmidt","year":"2017","journal-title":"Cell Host Microbe"},{"key":"B71","doi-asserted-by":"publisher","first-page":"1011","DOI":"10.1016\/j.immuni.2015.10.016","article-title":"A wave of regulatory T cells into neonatal skin mediates tolerance to commensal microbes.","volume":"43","author":"Scharschmidt","year":"2015","journal-title":"Immunity"},{"key":"B72","doi-asserted-by":"publisher","first-page":"ra12","DOI":"10.1126\/scisignal.2001270","article-title":"ATP inhibits the generation and function of regulatory T cells through the activation of purinergic P2X receptors.","volume":"4","author":"Schenk","year":"2011","journal-title":"Sci. Signal"},{"key":"B73","doi-asserted-by":"publisher","first-page":"564","DOI":"10.1038\/nature13577","article-title":"The alarmin IL-33 promotes regulatory T-cell function in the intestine.","volume":"513","author":"Schiering","year":"2014","journal-title":"Nature"},{"key":"B74","doi-asserted-by":"publisher","first-page":"993","DOI":"10.1126\/science.aaa9420","article-title":"MUCOSAL IMMUNOLOGY. Individual intestinal symbionts induce a distinct population of RORgamma(+) regulatory T cells.","volume":"349","author":"Sefik","year":"2015","journal-title":"Science"},{"key":"B75","doi-asserted-by":"publisher","first-page":"1527.e8","DOI":"10.1016\/j.immuni.2021.04.022","article-title":"Treg cell-derived osteopontin promotes microglia-mediated white matter repair after ischemic stroke.","volume":"54","author":"Shi","year":"2021","journal-title":"Immunity"},{"key":"B76","doi-asserted-by":"publisher","first-page":"1865","DOI":"10.3389\/fimmu.2019.01865","article-title":"IL-21 induces an imbalance of Th17\/Treg cells in moderate-to-severe plaque psoriasis patients.","volume":"10","author":"Shi","year":"2019","journal-title":"Front. Immunol."},{"key":"B77","doi-asserted-by":"publisher","first-page":"20696","DOI":"10.1073\/pnas.2000372117","article-title":"Proenkephalin(+) regulatory T cells expanded by ultraviolet B exposure maintain skin homeostasis with a healing function.","volume":"117","author":"Shime","year":"2020","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B78","doi-asserted-by":"publisher","first-page":"E512","DOI":"10.1152\/ajpendo.00341.2020","article-title":"Visceral adipose tissue mesenchymal stromal cells in the intersection of immunology and metabolism.","volume":"320","author":"Spallanzani","year":"2021","journal-title":"Am. J. Physiol. Endocrinol. Metab."},{"key":"B79","doi-asserted-by":"publisher","first-page":"410","DOI":"10.1016\/j.cmet.2008.04.004","article-title":"Beneficial effects of subcutaneous fat transplantation on metabolism.","volume":"7","author":"Tran","year":"2008","journal-title":"Cell Metab."},{"key":"B80","doi-asserted-by":"publisher","first-page":"1711","DOI":"10.3389\/fimmu.2020.01711","article-title":"Adoptive Cell transfer of regulatory T cells exacerbates hepatic steatosis in high-fat high-fructose diet-fed mice.","volume":"11","author":"Van Herck","year":"2020","journal-title":"Front. Immunol."},{"key":"B81","doi-asserted-by":"publisher","first-page":"581","DOI":"10.1038\/s41586-020-2040-3","article-title":"Sex-specific adipose tissue imprinting of regulatory T cells.","volume":"579","author":"Vasanthakumar","year":"2020","journal-title":"Nature"},{"key":"B82","doi-asserted-by":"publisher","first-page":"2906","DOI":"10.1016\/j.celrep.2017.08.068","article-title":"The TNF receptor superfamily-NF-kappaB axis is critical to maintain effector regulatory T cells in lymphoid and non-lymphoid tissues.","volume":"20","author":"Vasanthakumar","year":"2017","journal-title":"Cell Rep."},{"key":"B83","doi-asserted-by":"publisher","first-page":"544","DOI":"10.1038\/ni0515-544d","article-title":"Erratum: the transcriptional regulators IRF4, BATF and IL-33 orchestrate development and maintenance of adipose tissue-resident regulatory T cells.","volume":"16","author":"Vasanthakumar","year":"2015","journal-title":"Nat. Immunol."},{"key":"B84","doi-asserted-by":"publisher","first-page":"258ra142","DOI":"10.1126\/scitranslmed.3009925","article-title":"Regulatory T cells suppress muscle inflammation and injury in muscular dystrophy.","volume":"6","author":"Villalta","year":"2014","journal-title":"Sci. Transl. Med."},{"key":"B85","doi-asserted-by":"publisher","first-page":"5402","DOI":"10.1073\/pnas.1922559117","article-title":"Neuronal, stromal, and T-regulatory cell crosstalk in murine skeletal muscle.","volume":"117","author":"Wang","year":"2020","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B86","doi-asserted-by":"publisher","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":"B87","doi-asserted-by":"publisher","first-page":"678","DOI":"10.1111\/acel.12350","article-title":"Increases of M2a macrophages and fibrosis in aging muscle are influenced by bone marrow aging and negatively regulated by muscle-derived nitric oxide.","volume":"14","author":"Wang","year":"2015","journal-title":"Aging Cell"},{"key":"B88","doi-asserted-by":"publisher","first-page":"426","DOI":"10.1182\/blood-2006-01-0177","article-title":"Regulatory T-cell compartmentalization and trafficking.","volume":"108","author":"Wei","year":"2006","journal-title":"Blood"},{"key":"B89","doi-asserted-by":"publisher","first-page":"495","DOI":"10.1038\/s41586-018-0846-z","article-title":"Mitochondrial complex III is essential for suppressive function of regulatory T cells.","volume":"565","author":"Weinberg","year":"2019","journal-title":"Nature"},{"key":"B90","doi-asserted-by":"publisher","first-page":"4503","DOI":"10.1172\/JCI57456","article-title":"GATA3 controls Foxp3(+) regulatory T cell fate during inflammation in mice.","volume":"121","author":"Wohlfert","year":"2011","journal-title":"J. Clin. Invest."},{"key":"B91","doi-asserted-by":"publisher","first-page":"5488","DOI":"10.4049\/jimmunol.1400985","article-title":"Homeostasis of thymus-derived Foxp3+ regulatory T cells is controlled by ultraviolet B exposure in the skin.","volume":"193","author":"Yamazaki","year":"2014","journal-title":"J. Immunol."},{"key":"B92","doi-asserted-by":"publisher","first-page":"589","DOI":"10.1126\/science.aaa7017","article-title":"Immune tolerance. Regulatory T cells generated early in life play a distinct role in maintaining self-tolerance.","volume":"348","author":"Yang","year":"2015","journal-title":"Science"},{"key":"B93","doi-asserted-by":"publisher","first-page":"2011","DOI":"10.1038\/s41467-019-09541-y","article-title":"Loss of TET2 and TET3 in regulatory T cells unleashes effector function.","volume":"10","author":"Yue","year":"2019","journal-title":"Nat. Commun."},{"key":"B94","doi-asserted-by":"publisher","first-page":"710","DOI":"10.1038\/cmi.2017.70","article-title":"The altered PD-1\/PD-L1 pathway delivers the \u2018one-two punch\u2019 effects to promote the Treg\/Th17 imbalance in pre-eclampsia.","volume":"15","author":"Zhang","year":"2018","journal-title":"Cell Mol. Immunol."},{"key":"B95","doi-asserted-by":"publisher","first-page":"eaay6191","DOI":"10.1126\/sciadv.aay6191","article-title":"The obesity-induced adipokine sST2 exacerbates adipose Treg and ILC2 depletion and promotes insulin resistance.","volume":"6","author":"Zhao","year":"2020","journal-title":"Sci. Adv."}],"container-title":["Frontiers in Cell and Developmental Biology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcell.2021.717903\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,8,18]],"date-time":"2021-08-18T05:15:55Z","timestamp":1629263755000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcell.2021.717903\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,8,18]]},"references-count":95,"alternative-id":["10.3389\/fcell.2021.717903"],"URL":"https:\/\/doi.org\/10.3389\/fcell.2021.717903","relation":{},"ISSN":["2296-634X"],"issn-type":[{"value":"2296-634X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,8,18]]},"article-number":"717903"}}