{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T21:02:25Z","timestamp":1792789345098,"version":"4.5.0"},"reference-count":223,"publisher":"Wiley","issue":"1","license":[{"start":{"date-parts":[[2026,4,8]],"date-time":"2026-04-08T00:00:00Z","timestamp":1775606400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,4,8]],"date-time":"2026-04-08T00:00:00Z","timestamp":1775606400000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/100010269","name":"Wellcome Trust","doi-asserted-by":"publisher","award":["222623\/Z\/21\/Z"],"award-info":[{"award-number":["222623\/Z\/21\/Z"]}],"id":[{"id":"10.13039\/100010269","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Immunological Reviews"],"published-print":{"date-parts":[[2026,5]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>The thymic medulla is essential for establishing central tolerance, orchestrating the development of a diverse yet self\u2010tolerant T cell repertoire, and preventing autoimmunity. This process is primarily mediated through interactions between developing thymocytes and antigen\u2010presenting cells, including thymic epithelial cells (TECs) and dendritic cells (DCs), with additional regulatory contributions from endothelial cells, mesenchymal cells, and macrophages. Despite its critical role, the complexity of late\u2010stage thymocyte development and the dynamics of their medullary residency remain incompletely understood. Recent advances in single\u2010cell, epigenomic, and transcriptomic technologies have begun to reveal previously unappreciated layers of cellular and molecular heterogeneity within the thymic medulla throughout life. In this review, we explore how the medulla shapes the fate of both conventional and non\u2010conventional T cells, examine the diversity of thymocyte populations it supports, and discuss how this specialized microenvironment adapts during aging and regeneration.<\/jats:p>","DOI":"10.1111\/imr.70118","type":"journal-article","created":{"date-parts":[[2026,4,9]],"date-time":"2026-04-09T04:29:41Z","timestamp":1775708981000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["From Development to Aging: Dynamic Roles of the Thymic Medulla in T Cell Biology"],"prefix":"10.1111","volume":"339","author":[{"given":"Christina","family":"Michalaki","sequence":"first","affiliation":[{"name":"Division of Infection and Immunity University College London  London UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Minahil","family":"Sharjeel","sequence":"additional","affiliation":[{"name":"Division of Infection and Immunity University College London  London UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8461-3044","authenticated-orcid":false,"given":"Jennifer E.","family":"Cowan","sequence":"additional","affiliation":[{"name":"Division of Infection and Immunity University College London  London UK"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,4,8]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1002\/eji.201445108"},{"key":"e_1_2_9_3_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41577\u2010023\u201000911\u20108"},{"key":"e_1_2_9_4_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev\u2010cellbio\u2010112122\u2010023316"},{"key":"e_1_2_9_5_1","doi-asserted-by":"publisher","DOI":"10.1038\/ni.2938"},{"key":"e_1_2_9_6_1","doi-asserted-by":"publisher","DOI":"10.1038\/nri3667"},{"key":"e_1_2_9_7_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2021.646209"},{"key":"e_1_2_9_8_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.926625"},{"key":"e_1_2_9_9_1","doi-asserted-by":"publisher","DOI":"10.15252\/embj.2019101828"},{"key":"e_1_2_9_10_1","doi-asserted-by":"publisher","DOI":"10.1007\/978\u20101\u20100716\u20102740\u20102_2"},{"key":"e_1_2_9_11_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2005.12.011"},{"key":"e_1_2_9_12_1","doi-asserted-by":"publisher","DOI":"10.1002\/bies.202300165"},{"key":"e_1_2_9_13_1","doi-asserted-by":"publisher","DOI":"10.1002\/eji.70114"},{"key":"e_1_2_9_14_1","doi-asserted-by":"publisher","DOI":"10.1182\/bloodadvances.2020003192"},{"key":"e_1_2_9_15_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20161864"},{"key":"e_1_2_9_16_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20040643"},{"key":"e_1_2_9_17_1","doi-asserted-by":"publisher","DOI":"10.1182\/blood\u20102007\u201001\u2010070284"},{"key":"e_1_2_9_18_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0906956106"},{"key":"e_1_2_9_19_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.172.7.3999"},{"key":"e_1_2_9_20_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1400993"},{"key":"e_1_2_9_21_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2008.10.008"},{"key":"e_1_2_9_22_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.0804133"},{"key":"e_1_2_9_23_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0025567"},{"key":"e_1_2_9_24_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.80443"},{"key":"e_1_2_9_25_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20150178"},{"key":"e_1_2_9_26_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20070601"},{"key":"e_1_2_9_27_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41577\u2010020\u20100345\u2010y"},{"key":"e_1_2_9_28_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0708884104"},{"key":"e_1_2_9_29_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20122070"},{"key":"e_1_2_9_30_1","doi-asserted-by":"publisher","DOI":"10.1038\/ni.3419"},{"key":"e_1_2_9_31_1","doi-asserted-by":"publisher","DOI":"10.29245\/2578\u20103009\/2018\/1.1103"},{"key":"e_1_2_9_32_1","doi-asserted-by":"publisher","DOI":"10.1038\/ni1049"},{"key":"e_1_2_9_33_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1900662"},{"key":"e_1_2_9_34_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41590\u2010018\u20100289\u20106"},{"key":"e_1_2_9_35_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2013.02.022"},{"key":"e_1_2_9_36_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.883560"},{"key":"e_1_2_9_37_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2007.11.021"},{"key":"e_1_2_9_38_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2007.11.022"},{"key":"e_1_2_9_39_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.178.1.280"},{"key":"e_1_2_9_40_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41590\u2010023\u201001469\u20102"},{"key":"e_1_2_9_41_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.135.4.793"},{"key":"e_1_2_9_42_1","doi-asserted-by":"publisher","DOI":"10.1007\/s11427\u2010017\u20109298\u20103"},{"key":"e_1_2_9_43_1","doi-asserted-by":"publisher","DOI":"10.1038\/373531a0"},{"key":"e_1_2_9_44_1","doi-asserted-by":"publisher","DOI":"10.1016\/0092\u20108674(95)90416\u20106"},{"key":"e_1_2_9_45_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jaut.2017.03.007"},{"key":"e_1_2_9_46_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2009.04.009"},{"key":"e_1_2_9_47_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2014.08.007"},{"key":"e_1_2_9_48_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20112061"},{"key":"e_1_2_9_49_1","doi-asserted-by":"publisher","DOI":"10.1038\/ni1160"},{"key":"e_1_2_9_50_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41423\u2010019\u20100352\u20108"},{"key":"e_1_2_9_51_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010024\u201051164\u20105"},{"key":"e_1_2_9_52_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.178.2.1208"},{"key":"e_1_2_9_53_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2005.07.005"},{"key":"e_1_2_9_54_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.174.4.1862"},{"key":"e_1_2_9_55_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20102327"},{"key":"e_1_2_9_56_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.182.6.2091"},{"key":"e_1_2_9_57_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20050413"},{"key":"e_1_2_9_58_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1303057"},{"key":"e_1_2_9_59_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.34793"},{"key":"e_1_2_9_60_1","doi-asserted-by":"publisher","DOI":"10.12688\/f1000research.21378.1"},{"key":"e_1_2_9_61_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010018\u201005026\u20106"},{"key":"e_1_2_9_62_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1003965"},{"key":"e_1_2_9_63_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010018\u20100345\u20102"},{"key":"e_1_2_9_64_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.1001255"},{"key":"e_1_2_9_65_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2020.623265"},{"key":"e_1_2_9_66_1","doi-asserted-by":"publisher","DOI":"10.1002\/eji.202350388"},{"key":"e_1_2_9_67_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.173.5.1039"},{"key":"e_1_2_9_68_1","doi-asserted-by":"publisher","DOI":"10.1038\/icb.2008.87"},{"key":"e_1_2_9_69_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.168.5.1929"},{"key":"e_1_2_9_70_1","doi-asserted-by":"publisher","DOI":"10.1016\/0008\u20108749(89)90232\u20103"},{"key":"e_1_2_9_71_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.177.8.5014"},{"key":"e_1_2_9_72_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.177.2.379"},{"key":"e_1_2_9_73_1","doi-asserted-by":"publisher","DOI":"10.1002\/eji.201444529"},{"key":"e_1_2_9_74_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1501226"},{"key":"e_1_2_9_75_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2016.01.003"},{"key":"e_1_2_9_76_1","doi-asserted-by":"publisher","DOI":"10.1016\/s1074\u20107613(00)80095\u20103"},{"key":"e_1_2_9_77_1","doi-asserted-by":"publisher","DOI":"10.1038\/ni.3150"},{"key":"e_1_2_9_78_1","doi-asserted-by":"publisher","DOI":"10.1126\/sciimmunol.adu7341"},{"key":"e_1_2_9_79_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1102964"},{"key":"e_1_2_9_80_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2019.01.001"},{"key":"e_1_2_9_81_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41577\u2010021\u201000519\u2010w"},{"key":"e_1_2_9_82_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20061405"},{"key":"e_1_2_9_83_1","doi-asserted-by":"publisher","DOI":"10.1002\/eji.201747375"},{"key":"e_1_2_9_84_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010023\u201006881\u20100"},{"key":"e_1_2_9_85_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00018\u2010022\u201004328\u20109"},{"key":"e_1_2_9_86_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.58213"},{"key":"e_1_2_9_87_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.181.12.8391"},{"key":"e_1_2_9_88_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00262\u2010022\u201003313\u20102"},{"key":"e_1_2_9_89_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41419\u2010024\u201006806\u20103"},{"key":"e_1_2_9_90_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1101116"},{"key":"e_1_2_9_91_1","doi-asserted-by":"publisher","DOI":"10.1038\/nri2156"},{"key":"e_1_2_9_92_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1301235"},{"key":"e_1_2_9_93_1","doi-asserted-by":"publisher","DOI":"10.1038\/nri3605"},{"key":"e_1_2_9_94_1","doi-asserted-by":"publisher","DOI":"10.1038\/ni.3171"},{"key":"e_1_2_9_95_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010021\u201003531\u20101"},{"key":"e_1_2_9_96_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.it.2021.08.010"},{"key":"e_1_2_9_97_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.181.2.1207"},{"key":"e_1_2_9_98_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2018.00585"},{"key":"e_1_2_9_99_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2025.01.006"},{"key":"e_1_2_9_100_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365\u20103083.1985.tb01916.x"},{"key":"e_1_2_9_101_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010019\u201013465\u2010y"},{"key":"e_1_2_9_102_1","doi-asserted-by":"publisher","DOI":"10.1182\/blood\u20102006\u201002\u2010004531"},{"key":"e_1_2_9_103_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.it.2009.05.001"},{"key":"e_1_2_9_104_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.exger.2006.11.016"},{"key":"e_1_2_9_105_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0601040103"},{"key":"e_1_2_9_106_1","doi-asserted-by":"publisher","DOI":"10.1006\/cimm.1997.1148"},{"key":"e_1_2_9_107_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41577\u2010024\u201001119\u20100"},{"key":"e_1_2_9_108_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.93.15.7634"},{"key":"e_1_2_9_109_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20121716"},{"key":"e_1_2_9_110_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.164.4.1881"},{"key":"e_1_2_9_111_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2020.00897"},{"key":"e_1_2_9_112_1","doi-asserted-by":"publisher","DOI":"10.1182\/blood\u20102008\u201005\u2010156265"},{"key":"e_1_2_9_113_1","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M109.072124"},{"key":"e_1_2_9_114_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1474\u20109726.2010.00559.x"},{"key":"e_1_2_9_115_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2021.621824"},{"key":"e_1_2_9_116_1","doi-asserted-by":"publisher","DOI":"10.1038\/ni.2782"},{"key":"e_1_2_9_117_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1403158"},{"key":"e_1_2_9_118_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature04850"},{"key":"e_1_2_9_119_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature04813"},{"key":"e_1_2_9_120_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1301799110"},{"key":"e_1_2_9_121_1","doi-asserted-by":"publisher","DOI":"10.1002\/eji.201243209"},{"key":"e_1_2_9_122_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1203042"},{"key":"e_1_2_9_123_1","doi-asserted-by":"publisher","DOI":"10.1038\/414763a"},{"key":"e_1_2_9_124_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010023\u201037589\u20104"},{"key":"e_1_2_9_125_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2016.02.080"},{"key":"e_1_2_9_126_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2014.07.029"},{"key":"e_1_2_9_127_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010022\u201004752\u20108"},{"key":"e_1_2_9_128_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.devcel.2023.08.017"},{"key":"e_1_2_9_129_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010020\u201020082\u20107"},{"key":"e_1_2_9_130_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41577\u2010019\u20100238\u20100"},{"key":"e_1_2_9_131_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010018\u20100346\u20101"},{"key":"e_1_2_9_132_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010019\u201011879\u20102"},{"key":"e_1_2_9_133_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1701235"},{"key":"e_1_2_9_134_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1213069109"},{"key":"e_1_2_9_135_1","doi-asserted-by":"publisher","DOI":"10.1038\/283402a0"},{"key":"e_1_2_9_136_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.181.12.8199"},{"key":"e_1_2_9_137_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20080134"},{"key":"e_1_2_9_138_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20080829"},{"key":"e_1_2_9_139_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.2200609"},{"key":"e_1_2_9_140_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2020.00858"},{"key":"e_1_2_9_141_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41598\u2010017\u201005182\u20107"},{"key":"e_1_2_9_142_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.56221"},{"key":"e_1_2_9_143_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010023\u201039722\u20109"},{"key":"e_1_2_9_144_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.176.2.495"},{"key":"e_1_2_9_145_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2015.10.013"},{"key":"e_1_2_9_146_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41590\u2010020\u20100717\u20102"},{"key":"e_1_2_9_147_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2023.1277365"},{"key":"e_1_2_9_148_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1203203"},{"key":"e_1_2_9_149_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.92552"},{"key":"e_1_2_9_150_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.it.2012.03.005"},{"key":"e_1_2_9_151_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20170271"},{"key":"e_1_2_9_152_1","doi-asserted-by":"publisher","DOI":"10.1002\/eji.200737131"},{"key":"e_1_2_9_153_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20070795"},{"key":"e_1_2_9_154_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20062497"},{"key":"e_1_2_9_155_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.60188"},{"key":"e_1_2_9_156_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2013.08.038"},{"key":"e_1_2_9_157_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20092144"},{"key":"e_1_2_9_158_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2012.00019"},{"key":"e_1_2_9_159_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010020\u201016041\u2010x"},{"key":"e_1_2_9_160_1","doi-asserted-by":"publisher","DOI":"10.1111\/imr.70028"},{"key":"e_1_2_9_161_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2022.05.018"},{"key":"e_1_2_9_162_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.73998"},{"key":"e_1_2_9_163_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aay3224"},{"key":"e_1_2_9_164_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010021\u201021346\u20106"},{"key":"e_1_2_9_165_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2023.1186154"},{"key":"e_1_2_9_166_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2024.114072"},{"key":"e_1_2_9_167_1","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.aeb2970"},{"key":"e_1_2_9_168_1","doi-asserted-by":"publisher","DOI":"10.1111\/acel.13624"},{"key":"e_1_2_9_169_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2014.08.070"},{"key":"e_1_2_9_170_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.molimm.2018.04.010"},{"key":"e_1_2_9_171_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1474\u20109726.2011.00773.x"},{"key":"e_1_2_9_172_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41590\u2010024\u201001915\u20109"},{"key":"e_1_2_9_173_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41590\u2010017\u20100006\u2010x"},{"key":"e_1_2_9_174_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.smim.2023.101837"},{"key":"e_1_2_9_175_1","doi-asserted-by":"publisher","DOI":"10.1111\/imr.70050"},{"key":"e_1_2_9_176_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41598\u2010018\u201028111\u20108"},{"key":"e_1_2_9_177_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1714478115"},{"issue":"3","key":"e_1_2_9_178_1","first-page":"280","article-title":"Thymus Size and Age\u2010Related Thymic Involution: Early Programming, Sexual Dimorphism, Progenitors and Stroma","volume":"3","author":"Gui J.","year":"2012","journal-title":"Aging and Disease"},{"key":"e_1_2_9_179_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20050341"},{"key":"e_1_2_9_180_1","doi-asserted-by":"publisher","DOI":"10.1038\/25374"},{"key":"e_1_2_9_181_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.174.11.7446"},{"key":"e_1_2_9_182_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365\u20102249.2010.04209.x"},{"key":"e_1_2_9_183_1","doi-asserted-by":"publisher","DOI":"10.1111\/acel.12865"},{"key":"e_1_2_9_184_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.103614"},{"key":"e_1_2_9_185_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.3003283"},{"key":"e_1_2_9_186_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1302064"},{"key":"e_1_2_9_187_1","doi-asserted-by":"publisher","DOI":"10.1007\/s11357\u2010010\u20109170\u20108"},{"key":"e_1_2_9_188_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20090222"},{"key":"e_1_2_9_189_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20071140"},{"key":"e_1_2_9_190_1","doi-asserted-by":"publisher","DOI":"10.2307\/3284036"},{"key":"e_1_2_9_191_1","doi-asserted-by":"publisher","DOI":"10.1128\/iai.16.2.593\u2010598.1977"},{"key":"e_1_2_9_192_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41587\u2010025\u201002864\u2010w"},{"key":"e_1_2_9_193_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20211239"},{"key":"e_1_2_9_194_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jaci.2023.06.019"},{"key":"e_1_2_9_195_1","doi-asserted-by":"publisher","DOI":"10.1182\/blood\u20102011\u201003\u2010342097"},{"key":"e_1_2_9_196_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1002348"},{"key":"e_1_2_9_197_1","doi-asserted-by":"publisher","DOI":"10.1172\/jci.insight.140313"},{"key":"e_1_2_9_198_1","doi-asserted-by":"publisher","DOI":"10.1038\/ni983"},{"key":"e_1_2_9_199_1","doi-asserted-by":"publisher","DOI":"10.1002\/eji.201545814"},{"key":"e_1_2_9_200_1","doi-asserted-by":"publisher","DOI":"10.1136\/jmg.2011.089532"},{"key":"e_1_2_9_201_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1004623107"},{"key":"e_1_2_9_202_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.200995"},{"key":"e_1_2_9_203_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2024.1423488"},{"key":"e_1_2_9_204_1","doi-asserted-by":"publisher","DOI":"10.1126\/scitranslmed.adp3171"},{"key":"e_1_2_9_205_1","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.173.8.4867"},{"key":"e_1_2_9_206_1","doi-asserted-by":"publisher","DOI":"10.1182\/blood.v100.2.682"},{"key":"e_1_2_9_207_1","doi-asserted-by":"publisher","DOI":"10.1182\/blood\u20102006\u201010\u2010049767"},{"key":"e_1_2_9_208_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0042677"},{"key":"e_1_2_9_209_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010024\u201007238\u2010x"},{"key":"e_1_2_9_210_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010025\u201009873\u20104"},{"key":"e_1_2_9_211_1","doi-asserted-by":"publisher","DOI":"10.1002\/1521\u20104141(200211)32:11<3066::AID\u2010IMMU3066>3.0.CO;2\u2010J"},{"key":"e_1_2_9_212_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1218004"},{"key":"e_1_2_9_213_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.clim.2013.11.002"},{"key":"e_1_2_9_214_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbmt.2019.06.002"},{"key":"e_1_2_9_215_1","doi-asserted-by":"publisher","DOI":"10.1126\/sciimmunol.aal2736"},{"key":"e_1_2_9_216_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2021.109789"},{"key":"e_1_2_9_217_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010023\u201043072\u2010x"},{"key":"e_1_2_9_218_1","doi-asserted-by":"publisher","DOI":"10.1126\/sciimmunol.abq6930"},{"key":"e_1_2_9_219_1","doi-asserted-by":"publisher","DOI":"10.1126\/sciimmunol.abn3286"},{"key":"e_1_2_9_220_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41598\u2010021\u201089836\u20107"},{"key":"e_1_2_9_221_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41590\u2010025\u201002270\u2010z"},{"key":"e_1_2_9_222_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2019.03118"},{"key":"e_1_2_9_223_1","doi-asserted-by":"publisher","DOI":"10.1002\/eji.201646309"},{"key":"e_1_2_9_224_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010020\u201003071\u20100"}],"container-title":["Immunological Reviews"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/imr.70118","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1111\/imr.70118","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/imr.70118","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,27]],"date-time":"2026-05-27T06:46:17Z","timestamp":1779864377000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1111\/imr.70118"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,8]]},"references-count":223,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2026,5]]}},"alternative-id":["10.1111\/imr.70118"],"URL":"https:\/\/doi.org\/10.1111\/imr.70118","archive":["Portico"],"relation":{},"ISSN":["0105-2896","1600-065X"],"issn-type":[{"value":"0105-2896","type":"print"},{"value":"1600-065X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,4,8]]},"assertion":[{"value":"2026-03-05","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-03-12","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-04-08","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e70118"}}