{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,7]],"date-time":"2026-08-07T15:29:41Z","timestamp":1786116581917,"version":"build-2736575974"},"reference-count":66,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,5,18]],"date-time":"2026-05-18T00:00:00Z","timestamp":1779062400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,8,6]],"date-time":"2026-08-06T00:00:00Z","timestamp":1785974400000},"content-version":"vor","delay-in-days":80,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Commun Biol"],"DOI":"10.1038\/s42003-026-10299-5","type":"journal-article","created":{"date-parts":[[2026,5,18]],"date-time":"2026-05-18T11:35:06Z","timestamp":1779104106000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Aberrant activation of tissue-resident lymphocytes and pro-fibrotic GZMK\u207a CD8\u207a T cells in IPF"],"prefix":"10.1038","volume":"9","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1215-5822","authenticated-orcid":false,"given":"Fei","family":"Gao","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zitao","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yanjing","family":"Su","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuilian","family":"Chi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tao","family":"Yan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dong","family":"Wei","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yanjun","family":"Huang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jingbo","family":"Shao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhenhang","family":"Dai","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0842-2254","authenticated-orcid":false,"given":"Man","family":"Huang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0415-6080","authenticated-orcid":false,"given":"Chaoqun","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2127-1788","authenticated-orcid":false,"given":"Jingyu","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,5,18]]},"reference":[{"key":"10299_CR1","doi-asserted-by":"crossref","unstructured":"Gray, J. I. & Farber, D. L. Tissue-resident immune cells in humans. Annu. Rev. Immunol. 40, 195\u2013220 (2022).","DOI":"10.1146\/annurev-immunol-093019-112809"},{"key":"10299_CR2","doi-asserted-by":"publisher","first-page":"886","DOI":"10.1016\/j.immuni.2014.12.007","volume":"41","author":"JM Schenkel","year":"2014","unstructured":"Schenkel, J. M. & Masopust, D. Tissue-resident memory T cells. Immunity 41, 886\u2013897 (2014).","journal-title":"Immunity"},{"key":"10299_CR3","doi-asserted-by":"publisher","DOI":"10.1126\/sciimmunol.abm7996","volume":"7","author":"H Narasimhan","year":"2022","unstructured":"Narasimhan, H., Wu, Y., Goplen, N. P. & Sun, J. Immune determinants of chronic sequelae after respiratory viral infection. Sci. Immunol. 7, eabm7996 (2022).","journal-title":"Sci. Immunol."},{"key":"10299_CR4","doi-asserted-by":"publisher","first-page":"277","DOI":"10.1146\/annurev-immunol-101921-040450","volume":"41","author":"X Wei","year":"2023","unstructured":"Wei, X., Narasimhan, H., Zhu, B. & Sun, J. Host recovery from respiratory viral infection. Annu. Rev. Immunol. 41, 277\u2013300 (2023).","journal-title":"Annu. Rev. Immunol."},{"key":"10299_CR5","doi-asserted-by":"publisher","first-page":"576","DOI":"10.1016\/j.immuni.2023.01.032","volume":"56","author":"C Wang","year":"2023","unstructured":"Wang, C. et al. Dysregulated lung stroma drives emphysema exacerbation by potentiating resident lymphocytes to suppress an epithelial stem cell reservoir. Immunity 56, 576\u2013591.e510 (2023).","journal-title":"Immunity"},{"key":"10299_CR6","doi-asserted-by":"publisher","DOI":"10.1126\/scitranslmed.adn0136","volume":"16","author":"C Li","year":"2024","unstructured":"Li, C. et al. Comparative single-cell analysis reveals IFN-gamma as a driver of respiratory sequelae after acute COVID-19. Sci. Transl. Med. 16, eadn0136 (2024).","journal-title":"Sci. Transl. Med."},{"key":"10299_CR7","doi-asserted-by":"publisher","first-page":"1206","DOI":"10.1164\/rccm.2202055","volume":"165","author":"RM Strieter","year":"2002","unstructured":"Strieter, R. M. Con: Inflammatory mechanisms are not a minor component of the pathogenesis of idiopathic pulmonary fibrosis. Am. J. Respir. Crit. Care Med. 165, 1206\u20131207 (2002).","journal-title":"Am. J. Respir. Crit. Care Med."},{"key":"10299_CR8","doi-asserted-by":"publisher","first-page":"1811","DOI":"10.1056\/NEJMra1705751","volume":"378","author":"DJ Lederer","year":"2018","unstructured":"Lederer, D. J. & Martinez, F. J. Idiopathic pulmonary fibrosis. N. Engl. J. Med. 378, 1811\u20131823 (2018).","journal-title":"N. Engl. J. Med."},{"key":"10299_CR9","doi-asserted-by":"publisher","first-page":"1176","DOI":"10.1513\/AnnalsATS.201404-145OC","volume":"11","author":"JP Hutchinson","year":"2014","unstructured":"Hutchinson, J. P., McKeever, T. M., Fogarty, A. W., Navaratnam, V. & Hubbard, R. B. Increasing global mortality from idiopathic pulmonary fibrosis in the twenty-first century. Ann. Am. Thorac. Soc. 11, 1176\u20131185 (2014).","journal-title":"Ann. Am. Thorac. Soc."},{"key":"10299_CR10","doi-asserted-by":"publisher","DOI":"10.1016\/j.pharmthera.2020.107798","volume":"222","author":"P Spagnolo","year":"2021","unstructured":"Spagnolo, P. et al. Idiopathic pulmonary fibrosis: disease mechanisms and drug development. Pharm. Ther. 222, 107798 (2021).","journal-title":"Pharm. Ther."},{"key":"10299_CR11","doi-asserted-by":"publisher","DOI":"10.1016\/j.pharmthera.2023.108562","volume":"252","author":"SE Mutsaers","year":"2023","unstructured":"Mutsaers, S. E., Miles, T., Prele, C. M. & Hoyne, G. F. Emerging role of immune cells as drivers of pulmonary fibrosis. Pharm. Ther. 252, 108562 (2023).","journal-title":"Pharm. Ther."},{"key":"10299_CR12","doi-asserted-by":"publisher","unstructured":"Celada, L. J. et al. PD-1 up-regulation on CD4(+) T cells promotes pulmonary fibrosis through STAT3-mediated IL-17A and TGF-beta1 production. Sci. Transl. Med. 10, https:\/\/doi.org\/10.1126\/scitranslmed.aar8356 (2018).","DOI":"10.1126\/scitranslmed.aar8356"},{"key":"10299_CR13","doi-asserted-by":"publisher","first-page":"1469","DOI":"10.1038\/s41590-019-0494-y","volume":"20","author":"T Ichikawa","year":"2019","unstructured":"Ichikawa, T. et al. CD103(hi) T(reg) cells constrain lung fibrosis induced by CD103(lo) tissue-resident pathogenic CD4 T cells. Nat. Immunol. 20, 1469\u20131480 (2019).","journal-title":"Nat. Immunol."},{"key":"10299_CR14","doi-asserted-by":"publisher","unstructured":"Goplen, N. P. et al. Tissue-resident CD8(+) T cells drive age-associated chronic lung sequelae after viral pneumonia. Sci. Immunol. 5, https:\/\/doi.org\/10.1126\/sciimmunol.abc4557 (2020).","DOI":"10.1126\/sciimmunol.abc4557"},{"key":"10299_CR15","doi-asserted-by":"publisher","first-page":"3506","DOI":"10.1016\/j.cell.2024.05.015","volume":"187","author":"M Kamiya","year":"2024","unstructured":"Kamiya, M. et al. Immune mechanisms in fibrotic interstitial lung disease. Cell 187, 3506\u20133530 (2024).","journal-title":"Cell"},{"key":"10299_CR16","doi-asserted-by":"publisher","first-page":"1207","DOI":"10.1164\/rccm.202012-4358ED","volume":"203","author":"S Soma","year":"2021","unstructured":"Soma, S. & Harriff, M. J. Ex vivo lung perfusion provides new insights into human lung-resident immune cell localization and functional interactions. Am. J. Respir. Crit. Care Med. 203, 1207\u20131208 (2021).","journal-title":"Am. J. Respir. Crit. Care Med."},{"key":"10299_CR17","doi-asserted-by":"publisher","first-page":"124","DOI":"10.1016\/j.immuni.2023.11.018","volume":"57","author":"T Torcellan","year":"2024","unstructured":"Torcellan, T. et al. Circulating NK cells establish tissue residency upon acute infection of skin and mediate accelerated effector responses to secondary infection. Immunity 57, 124\u2013140.e127 (2024).","journal-title":"Immunity"},{"key":"10299_CR18","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1016\/j.immuni.2020.11.005","volume":"54","author":"DA Mogilenko","year":"2021","unstructured":"Mogilenko, D. A. et al. Comprehensive profiling of an aging immune system reveals clonal GZMK(+) CD8(+) T cells as conserved hallmark of inflammaging. Immunity 54, 99\u2013115.e112 (2021).","journal-title":"Immunity"},{"key":"10299_CR19","doi-asserted-by":"publisher","first-page":"1112","DOI":"10.1038\/cdd.2011.5","volume":"18","author":"LT Joeckel","year":"2011","unstructured":"Joeckel, L. T. et al. Mouse granzyme K has pro-inflammatory potential. Cell Death Differ. 18, 1112\u20131119 (2011).","journal-title":"Cell Death Differ."},{"key":"10299_CR20","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/icb.2017.35","volume":"95","author":"LT Joeckel","year":"2017","unstructured":"Joeckel, L. T., Allison, C. C., Pellegrini, M., Bird, C. H. & Bird, P. I. Granzyme K-deficient mice show no evidence of impaired antiviral immunity. Immunol. Cell Biol. 95, 676\u2013683 (2017).","journal-title":"Immunol. Cell Biol."},{"key":"10299_CR21","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.aba1972","volume":"6","author":"AC Habermann","year":"2020","unstructured":"Habermann, A. C. et al. Single-cell RNA sequencing reveals profibrotic roles of distinct epithelial and mesenchymal lineages in pulmonary fibrosis. Sci. Adv. 6, eaba1972 (2020).","journal-title":"Sci. Adv."},{"key":"10299_CR22","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.aba1983","volume":"6","author":"TS Adams","year":"2020","unstructured":"Adams, T. S. et al. Single-cell RNA-seq reveals ectopic and aberrant lung-resident cell populations in idiopathic pulmonary fibrosis. Sci. Adv. 6, eaba1983 (2020).","journal-title":"Sci. Adv."},{"key":"10299_CR23","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms14532","volume":"8","author":"MJ Schafer","year":"2017","unstructured":"Schafer, M. J. et al. Cellular senescence mediates fibrotic pulmonary disease. Nat. Commun. 8, 14532 (2017).","journal-title":"Nat. Commun."},{"key":"10299_CR24","doi-asserted-by":"publisher","unstructured":"Lee, J. Y. et al. An in vivo screening platform identifies senolytic compounds that target p16INK4a+ fibroblasts in lung fibrosis. J. Clin. Investig. 134, https:\/\/doi.org\/10.1172\/JCI173371 (2024).","DOI":"10.1172\/JCI173371"},{"key":"10299_CR25","doi-asserted-by":"publisher","first-page":"970","DOI":"10.4049\/jimmunol.1202805","volume":"190","author":"SK Bromley","year":"2013","unstructured":"Bromley, S. K., Yan, S., Tomura, M., Kanagawa, O. & Luster, A. D. Recirculating memory T cells are a unique subset of CD4+ T cells with a distinct phenotype and migratory pattern. J. Immunol. 190, 970\u2013976 (2013).","journal-title":"J. Immunol."},{"key":"10299_CR26","doi-asserted-by":"publisher","first-page":"1174","DOI":"10.1038\/ni730","volume":"2","author":"D Tzachanis","year":"2001","unstructured":"Tzachanis, D. et al. Tob is a negative regulator of activation that is expressed in anergic and quiescent T cells. Nat. Immunol. 2, 1174\u20131182 (2001).","journal-title":"Nat. Immunol."},{"key":"10299_CR27","doi-asserted-by":"publisher","first-page":"615","DOI":"10.2337\/db10-1574","volume":"61","author":"T Marker","year":"2012","unstructured":"Marker, T. et al. Heat shock protein 60 as a mediator of adipose tissue inflammation and insulin resistance. Diabetes 61, 615\u2013625 (2012).","journal-title":"Diabetes"},{"key":"10299_CR28","doi-asserted-by":"publisher","first-page":"75","DOI":"10.3389\/fimmu.2012.00075","volume":"3","author":"SK Calderwood","year":"2012","unstructured":"Calderwood, S. K., Murshid, A. & Gong, J. Heat shock proteins: conditional mediators of inflammation in tumor immunity. Front Immunol. 3, 75 (2012).","journal-title":"Front Immunol."},{"key":"10299_CR29","doi-asserted-by":"publisher","unstructured":"Unterman, A. et al. Single-cell profiling reveals immune aberrations in progressive idiopathic pulmonary fibrosis. Am. J. Respir. Crit. Care Med. https:\/\/doi.org\/10.1164\/rccm.202306-0979OC (2024).","DOI":"10.1164\/rccm.202306-0979OC"},{"key":"10299_CR30","doi-asserted-by":"publisher","first-page":"899","DOI":"10.1016\/j.immuni.2009.03.019","volume":"30","author":"M Miyara","year":"2009","unstructured":"Miyara, M. et al. Functional delineation and differentiation dynamics of human CD4+ T cells expressing the FoxP3 transcription factor. Immunity 30, 899\u2013911 (2009).","journal-title":"Immunity"},{"key":"10299_CR31","doi-asserted-by":"publisher","first-page":"291","DOI":"10.1038\/s41590-018-0051-0","volume":"19","author":"D Zemmour","year":"2018","unstructured":"Zemmour, D. et al. Single-cell gene expression reveals a landscape of regulatory T cell phenotypes shaped by the TCR. Nat. Immunol. 19, 291\u2013301 (2018).","journal-title":"Nat. Immunol."},{"key":"10299_CR32","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.aaw8330","volume":"5","author":"Y Elyahu","year":"2019","unstructured":"Elyahu, Y. et al. Aging promotes reorganization of the CD4 T cell landscape toward extreme regulatory and effector phenotypes. Sci. Adv. 5, eaaw8330 (2019).","journal-title":"Sci. Adv."},{"key":"10299_CR33","doi-asserted-by":"publisher","first-page":"775","DOI":"10.1016\/j.cell.2008.05.009","volume":"133","author":"S Sakaguchi","year":"2008","unstructured":"Sakaguchi, S., Yamaguchi, T., Nomura, T. & Ono, M. Regulatory T cells and immune tolerance. Cell 133, 775\u2013787 (2008).","journal-title":"Cell"},{"key":"10299_CR34","doi-asserted-by":"publisher","first-page":"330","DOI":"10.1038\/ni904","volume":"4","author":"JD Fontenot","year":"2003","unstructured":"Fontenot, J. D., Gavin, M. A. & Rudensky, A. Y. Foxp3 programs the development and function of CD4+CD25+ regulatory T cells. Nat. Immunol. 4, 330\u2013336 (2003).","journal-title":"Nat. Immunol."},{"key":"10299_CR35","doi-asserted-by":"publisher","first-page":"1524","DOI":"10.1073\/pnas.0910341107","volume":"107","author":"N Jain","year":"2010","unstructured":"Jain, N., Nguyen, H., Chambers, C. & Kang, J. Dual function of CTLA-4 in regulatory T cells and conventional T cells to prevent multiorgan autoimmunity. Proc. Natl. Acad. Sci. USA 107, 1524\u20131528 (2010).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10299_CR36","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1111\/j.1600-065X.2011.01004.x","volume":"241","author":"G Cheng","year":"2011","unstructured":"Cheng, G., Yu, A. & Malek, T. R. T-cell tolerance and the multi-functional role of IL-2R signaling in T-regulatory cells. Immunol. Rev. 241, 63\u201376 (2011).","journal-title":"Immunol. Rev."},{"key":"10299_CR37","doi-asserted-by":"publisher","first-page":"1121","DOI":"10.1164\/rccm.200812-1936OC","volume":"179","author":"I Kotsianidis","year":"2009","unstructured":"Kotsianidis, I. et al. Global impairment of CD4+CD25+FOXP3+ regulatory T cells in idiopathic pulmonary fibrosis. Am. J. Respir. Crit. Care Med 179, 1121\u20131130 (2009).","journal-title":"Am. J. Respir. Crit. Care Med"},{"key":"10299_CR38","doi-asserted-by":"publisher","unstructured":"Prele, C. M. et al. Plasma cell but not CD20-mediated B-cell depletion protects from bleomycin-induced lung fibrosis. Eur. Respir. J. 60, https:\/\/doi.org\/10.1183\/13993003.01469-2021 (2022).","DOI":"10.1183\/13993003.01469-2021"},{"key":"10299_CR39","doi-asserted-by":"publisher","first-page":"722","DOI":"10.1165\/rcmb.2020-0387OC","volume":"64","author":"MF Ali","year":"2021","unstructured":"Ali, M. F. et al. Antifibrotics modify B-cell-induced fibroblast migration and activation in patients with idiopathic pulmonary fibrosis. Am. J. Respir. Cell Mol. Biol. 64, 722\u2013733 (2021).","journal-title":"Am. J. Respir. Cell Mol. Biol."},{"key":"10299_CR40","doi-asserted-by":"publisher","first-page":"2089","DOI":"10.4049\/jimmunol.1203476","volume":"191","author":"J Xue","year":"2013","unstructured":"Xue, J. et al. Plasma B lymphocyte stimulator and B cell differentiation in idiopathic pulmonary fibrosis patients. J. Immunol. 191, 2089\u20132095 (2013).","journal-title":"J. Immunol."},{"key":"10299_CR41","doi-asserted-by":"publisher","first-page":"290","DOI":"10.1038\/ni.2527","volume":"14","author":"V Peperzak","year":"2013","unstructured":"Peperzak, V. et al. Mcl-1 is essential for the survival of plasma cells. Nat. Immunol. 14, 290\u2013297 (2013).","journal-title":"Nat. Immunol."},{"key":"10299_CR42","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-15647-5","volume":"11","author":"T Tsukui","year":"2020","unstructured":"Tsukui, T. et al. Collagen-producing lung cell atlas identifies multiple subsets with distinct localization and relevance to fibrosis. Nat. Commun. 11, 1920 (2020).","journal-title":"Nat. Commun."},{"key":"10299_CR43","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-021-21246-9","volume":"12","author":"S Jin","year":"2021","unstructured":"Jin, S. et al. Inference and analysis of cell-cell communication using CellChat. Nat. Commun. 12, 1088 (2021).","journal-title":"Nat. Commun."},{"key":"10299_CR44","doi-asserted-by":"publisher","first-page":"1633","DOI":"10.2353\/ajpath.2006.060501","volume":"169","author":"H Yasuoka","year":"2006","unstructured":"Yasuoka, H. et al. Insulin-like growth factor-binding protein-5 induces pulmonary fibrosis and triggers mononuclear cellular infiltration. Am. J. Pathol. 169, 1633\u20131642 (2006).","journal-title":"Am. J. Pathol."},{"key":"10299_CR45","doi-asserted-by":"publisher","first-page":"160","DOI":"10.1038\/ki.2011.305","volume":"81","author":"H Yokoi","year":"2012","unstructured":"Yokoi, H. et al. Pleiotrophin triggers inflammation and increased peritoneal permeability leading to peritoneal fibrosis. Kidney Int. 81, 160\u2013169 (2012).","journal-title":"Kidney Int."},{"key":"10299_CR46","doi-asserted-by":"publisher","first-page":"740","DOI":"10.1016\/j.ajpath.2016.12.006","volume":"187","author":"T Masuda","year":"2017","unstructured":"Masuda, T. et al. Growth factor midkine promotes T-cell activation through nuclear factor of activated T cells signaling and Th1 cell differentiation in Lupus Nephritis. Am. J. Pathol. 187, 740\u2013751 (2017).","journal-title":"Am. J. Pathol."},{"key":"10299_CR47","doi-asserted-by":"publisher","first-page":"1400","DOI":"10.1038\/mi.2017.3","volume":"10","author":"FM Verhamme","year":"2017","unstructured":"Verhamme, F. M. et al. Elevated GDF-15 contributes to pulmonary inflammation upon cigarette smoke exposure. Mucosal Immunol. 10, 1400\u20131411 (2017).","journal-title":"Mucosal Immunol."},{"key":"10299_CR48","doi-asserted-by":"publisher","first-page":"785","DOI":"10.4049\/jimmunol.145.3.785","volume":"145","author":"LS Davis","year":"1990","unstructured":"Davis, L. S., Oppenheimer-Marks, N., Bednarczyk, J. L., McIntyre, B. W. & Lipsky, P. E. Fibronectin promotes proliferation of naive and memory T cells by signaling through both the VLA-4 and VLA-5 integrin molecules. J. Immunol. 145, 785\u2013793 (1990).","journal-title":"J. Immunol."},{"key":"10299_CR49","doi-asserted-by":"publisher","DOI":"10.1126\/scisignal.abn9405","volume":"16","author":"M Weiss","year":"2023","unstructured":"Weiss, M. et al. Adhesion to laminin-1 and collagen IV induces the formation of Ca(2+) microdomains that sensitize mouse T cells for activation. Sci. Signal 16, eabn9405 (2023).","journal-title":"Sci. Signal"},{"key":"10299_CR50","doi-asserted-by":"publisher","first-page":"509","DOI":"10.1083\/jcb.200109098","volume":"157","author":"Z Li","year":"2002","unstructured":"Li, Z. et al. Interactions of thrombospondins with alpha4beta1 integrin and CD47 differentially modulate T cell behavior. J. Cell Biol. 157, 509\u2013519 (2002).","journal-title":"J. Cell Biol."},{"key":"10299_CR51","doi-asserted-by":"publisher","first-page":"176","DOI":"10.1038\/emm.2007.20","volume":"39","author":"KI Oh","year":"2007","unstructured":"Oh, K. I. et al. CD99 activates T cells via a costimulatory function that promotes raft association of TCR complex and tyrosine phosphorylation of TCR z. eta. Exp. Mol. Med. 39, 176\u2013184 (2007).","journal-title":"eta. Exp. Mol. Med."},{"key":"10299_CR52","doi-asserted-by":"publisher","first-page":"1070","DOI":"10.4049\/jimmunol.177.2.1070","volume":"177","author":"M Capasso","year":"2006","unstructured":"Capasso, M. et al. Costimulation via CD55 on human CD4+ T cells mediated by CD97. J. Immunol. 177, 1070\u20131077 (2006).","journal-title":"J. Immunol."},{"key":"10299_CR53","doi-asserted-by":"publisher","first-page":"1230","DOI":"10.1164\/rccm.202006-2403OC","volume":"203","author":"ME Snyder","year":"2021","unstructured":"Snyder, M. E. et al. Human lung-resident macrophages colocalize with and provide costimulation to PD1(hi) Tissue-resident memory T cells. Am. J. Respir. Crit. Care Med. 203, 1230\u20131244 (2021).","journal-title":"Am. J. Respir. Crit. Care Med."},{"key":"10299_CR54","doi-asserted-by":"publisher","first-page":"682","DOI":"10.1038\/nature04444","volume":"439","author":"DL Barber","year":"2006","unstructured":"Barber, D. L. et al. Restoring function in exhausted CD8 T cells during chronic viral infection. Nature 439, 682\u2013687 (2006).","journal-title":"Nature"},{"key":"10299_CR55","doi-asserted-by":"publisher","first-page":"684","DOI":"10.1165\/rcmb.2017-0326OC","volume":"58","author":"K Ni","year":"2018","unstructured":"Ni, K. et al. PD-1\/PD-L1 pathway mediates the alleviation of pulmonary fibrosis by human mesenchymal stem cells in humanized mice. Am. J. Respir. Cell Mol. Biol. 58, 684\u2013695 (2018).","journal-title":"Am. J. Respir. Cell Mol. Biol."},{"key":"10299_CR56","doi-asserted-by":"publisher","unstructured":"Wang, Z. et al. PD-1(hi) CD8(+) resident memory T cells balance immunity and fibrotic sequelae. Sci. Immunol. 4, https:\/\/doi.org\/10.1126\/sciimmunol.aaw1217 (2019).","DOI":"10.1126\/sciimmunol.aaw1217"},{"key":"10299_CR57","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1186\/1465-9921-6-81","volume":"6","author":"Z Daniil","year":"2005","unstructured":"Daniil, Z. et al. CD8+ T lymphocytes in lung tissue from patients with idiopathic pulmonary fibrosis. Respir. Res. 6, 81 (2005).","journal-title":"Respir. Res."},{"key":"10299_CR58","doi-asserted-by":"publisher","first-page":"535","DOI":"10.1084\/jem.20092121","volume":"207","author":"MS Wilson","year":"2010","unstructured":"Wilson, M. S. et al. Bleomycin and IL-1beta-mediated pulmonary fibrosis is IL-17A dependent. J. Exp. Med. 207, 535\u2013552 (2010).","journal-title":"J. Exp. Med."},{"key":"10299_CR59","doi-asserted-by":"publisher","first-page":"516","DOI":"10.3389\/fimmu.2016.00516","volume":"7","author":"A Adegunsoye","year":"2016","unstructured":"Adegunsoye, A. et al. Skewed lung CCR4 to CCR6 CD4(+) T cell ratio in idiopathic pulmonary fibrosis is associated with pulmonary function. Front Immunol. 7, 516 (2016).","journal-title":"Front Immunol."},{"key":"10299_CR60","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-020-78140-5","volume":"10","author":"HC Kim","year":"2020","unstructured":"Kim, H. C. et al. Histologic features suggesting connective tissue disease in idiopathic pulmonary fibrosis. Sci. Rep. 10, 21137 (2020).","journal-title":"Sci. Rep."},{"key":"10299_CR61","doi-asserted-by":"publisher","first-page":"3574","DOI":"10.1002\/art.23995","volume":"58","author":"K Komura","year":"2008","unstructured":"Komura, K. et al. CD19 regulates the development of bleomycin-induced pulmonary fibrosis in a mouse model. Arthritis Rheum. 58, 3574\u20133584 (2008).","journal-title":"Arthritis Rheum."},{"key":"10299_CR62","doi-asserted-by":"publisher","first-page":"793","DOI":"10.1016\/j.cmet.2024.01.015","volume":"36","author":"L Yu","year":"2024","unstructured":"Yu, L. et al. IgG is an aging factor that drives adipose tissue fibrosis and metabolic decline. Cell Metab. 36, 793\u2013807.e795 (2024).","journal-title":"Cell Metab."},{"key":"10299_CR63","doi-asserted-by":"publisher","first-page":"209","DOI":"10.1038\/nprot.2014.005","volume":"9","author":"KG Anderson","year":"2014","unstructured":"Anderson, K. G. et al. Intravascular staining for discrimination of vascular and tissue leukocytes. Nat. Protoc. 9, 209\u2013222 (2014).","journal-title":"Nat. Protoc."},{"key":"10299_CR64","doi-asserted-by":"publisher","unstructured":"Tkachev, V. et al. Spatiotemporal single-cell profiling reveals that invasive and tissue-resident memory donor CD8(+) T cells drive gastrointestinal acute graft-versus-host disease. Sci. Transl. Med. 13, https:\/\/doi.org\/10.1126\/scitranslmed.abc0227 (2021).","DOI":"10.1126\/scitranslmed.abc0227"},{"key":"10299_CR65","doi-asserted-by":"publisher","unstructured":"Potter, E. L. et al. Measurement of leukocyte trafficking kinetics in macaques by serial intravascular staining. Sci. Transl. Med. 13, https:\/\/doi.org\/10.1126\/scitranslmed.abb4582 (2021).","DOI":"10.1126\/scitranslmed.abb4582"},{"key":"10299_CR66","doi-asserted-by":"publisher","first-page":"10","DOI":"10.1038\/s41556-021-00809-4","volume":"24","author":"JJ Kathiriya","year":"2022","unstructured":"Kathiriya, J. J. et al. Human alveolar type 2 epithelium transdifferentiates into metaplastic KRT5 basal cells. Nat. Cell Biol. 24, 10\u201323 (2022).","journal-title":"Nat. Cell Biol."}],"container-title":["Communications Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s42003-026-10299-5","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-026-10299-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-026-10299-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,6]],"date-time":"2026-08-06T11:03:22Z","timestamp":1786014202000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s42003-026-10299-5"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,18]]},"references-count":66,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["10299"],"URL":"https:\/\/doi.org\/10.1038\/s42003-026-10299-5","relation":{"has-preprint":[{"id-type":"doi","id":"10.21203\/rs.3.rs-7500111\/v1","asserted-by":"object"}]},"ISSN":["2399-3642"],"issn-type":[{"value":"2399-3642","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,18]]},"assertion":[{"value":"31 August 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 May 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"18 May 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Ethics"}}],"article-number":"1051"}}