{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,28]],"date-time":"2026-10-28T00:54:42Z","timestamp":1793148882690,"version":"4.5.0"},"reference-count":85,"publisher":"American Society for Clinical Investigation","issue":"6","license":[{"start":{"date-parts":[[2026,3,23]],"date-time":"2026-03-23T00:00:00Z","timestamp":1774224000000},"content-version":"vor","delay-in-days":41,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100031158","name":"Penn Diabetes Research Center","doi-asserted-by":"crossref","award":["Pilot Grant"],"award-info":[{"award-number":["Pilot Grant"]}],"id":[{"id":"10.13039\/100031158","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100021041","name":"American College of Allergy, Asthma, and Immunology","doi-asserted-by":"crossref","award":["Junior Faculty Grant"],"award-info":[{"award-number":["Junior Faculty Grant"]}],"id":[{"id":"10.13039\/100021041","id-type":"DOI","asserted-by":"crossref"}]},{"name":"Children\u2019s Hospital of Philadelphia Research","award":["N\/A"],"award-info":[{"award-number":["N\/A"]}]},{"name":"University of Pennsylvania Theo Wilson Graduate Research Fellowship","award":["N\/A"],"award-info":[{"award-number":["N\/A"]}]},{"name":"American Heart Association Predoctoral Fellowship","award":["24PRE1191220"],"award-info":[{"award-number":["24PRE1191220"]}]},{"DOI":"10.13039\/100007457","name":"The JPB Foundation","doi-asserted-by":"crossref","award":["N\/A"],"award-info":[{"award-number":["N\/A"]}],"id":[{"id":"10.13039\/100007457","id-type":"DOI","asserted-by":"crossref"}]},{"name":"University of Pennsylvania Michael Brown Graduate Research Fellowship","award":["N\/A"],"award-info":[{"award-number":["N\/A"]}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["K08 DK116668,R01 HL162715,5T32 DK007314-43,R01 DK49780,T32 GM007170,R35 HL166663"],"award-info":[{"award-number":["K08 DK116668,R01 HL162715,5T32 DK007314-43,R01 DK49780,T32 GM007170,R35 HL166663"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,3,23]]},"abstract":"<jats:p>Dysfunctional white adipose tissue contributes to the development of obesity-related morbidities, including insulin resistance, dyslipidemia, and other metabolic disorders. Adipose tissue macrophages (ATMs) accumulate in obesity and play both beneficial and harmful roles in the maintenance of adipose tissue homeostasis and function. Despite their importance, the molecules and mechanisms that regulate these diverse functions are not well understood. Lipid-associated macrophages (LAMs), the dominant subset of obesity-associated ATMs, accumulate in crown-like structures and are characterized by a metabolically activated and proinflammatory phenotype. We previously identified CD9 as a surface marker of LAMs. However, the contribution of CD9 to the activation and function of LAMs during obesity is unknown. Using a myeloid-specific CD9-KO model, we show that CD9 supports ATM-adipocyte adhesion and crown-like structure formation. Furthermore, CD9 promotes the expression of profibrotic and extracellular matrix remodeling genes. Loss of myeloid CD9 reduces adipose tissue fibrosis, increases visceral adipose tissue accumulation, and improves global metabolic outcomes during diet-induced obesity. These results identify CD9 as a causal regulator of pathogenic LAM functions, highlighting CD9 as a potential therapeutic target for treating obesity-associated metabolic disease.<\/jats:p>","DOI":"10.1172\/jci.insight.193837","type":"journal-article","created":{"date-parts":[[2026,2,10]],"date-time":"2026-02-10T17:00:15Z","timestamp":1770742815000},"source":"Crossref","is-referenced-by-count":7,"title":["CD9 regulates macrophage-mediated remodeling of adipose tissue in obesity"],"prefix":"10.1172","volume":"11","author":[{"given":"Julia","family":"Chini","sequence":"first","affiliation":[{"name":"Department of Pediatrics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA."},{"name":"Division of Allergy and Immunology, Children\u2019s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA."},{"name":"Institute for Immunology and Immune Health, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nicole","family":"DeMarco","sequence":"additional","affiliation":[{"name":"Division of Allergy and Immunology, Children\u2019s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dana V.","family":"Mitchell","sequence":"additional","affiliation":[{"name":"Department of Biomedical and Health Informatics, Children\u2019s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sam J.","family":"McCright","sequence":"additional","affiliation":[{"name":"Department of Pediatrics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA."},{"name":"Division of Allergy and Immunology, Children\u2019s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA."},{"name":"Institute for Immunology and Immune Health, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kaitlyn M.","family":"Shen","sequence":"additional","affiliation":[{"name":"Departments of Medicine and Cell and Developmental Biology, Penn Cardiovascular Institute, Penn Epigenetics Institute,"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Divyansi","family":"Pandey","sequence":"additional","affiliation":[{"name":"Department of Pediatrics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA."},{"name":"Division of Allergy and Immunology, Children\u2019s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA."},{"name":"Institute for Immunology and Immune Health, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rachel L.","family":"Clement","sequence":"additional","affiliation":[{"name":"Department of Pediatrics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA."},{"name":"Division of Allergy and Immunology, Children\u2019s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA."},{"name":"Institute for Immunology and Immune Health, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jessica","family":"Miller","sequence":"additional","affiliation":[{"name":"Division of Allergy and Immunology, Children\u2019s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rajan","family":"Jain","sequence":"additional","affiliation":[{"name":"Departments of Medicine and Cell and Developmental Biology, Penn Cardiovascular Institute, Penn Epigenetics Institute,"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3302-4610","authenticated-orcid":true,"given":"Deanne M.","family":"Taylor","sequence":"additional","affiliation":[{"name":"Department of Pediatrics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA."},{"name":"Department of Biomedical and Health Informatics, Children\u2019s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mitchell A.","family":"Lazar","sequence":"additional","affiliation":[{"name":"Department of Medicine, Division of Endocrinology, and"},{"name":"Institute for Diabetes, Obesity and Metabolism, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9286-4268","authenticated-orcid":true,"given":"David A.","family":"Hill","sequence":"additional","affiliation":[{"name":"Department of Pediatrics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA."},{"name":"Division of Allergy and Immunology, Children\u2019s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA."},{"name":"Institute for Immunology and Immune Health, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA."},{"name":"Institute for Diabetes, Obesity and Metabolism, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA."}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"232","published-online":{"date-parts":[[2026,2,10]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.1172\/JCI45887"},{"key":"B2","doi-asserted-by":"publisher","DOI":"10.1111\/dom.15637"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.1016\/j.molmet.2015.12.001"},{"key":"B4","doi-asserted-by":"publisher","DOI":"10.20900\/immunometab20190016"},{"key":"B5","doi-asserted-by":"publisher","DOI":"10.2337\/db16-0500"},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-30646-4"},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.2337\/db07-0767"},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.1038\/s41419-021-03872-9"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.1016\/j.cmet.2014.08.010"},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2017.08.096"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2019.05.054"},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1126\/science.aaw2586"},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.1172\/JCI29881"},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.2337\/db08-0872"},{"key":"B15","doi-asserted-by":"crossref","unstructured":"Askia H, et al. Obesity-elicited macrophages shape CD9hi progenitor fate to promote adipose tissue fibrosis and dysfunction [Preprint]. https:\/\/doi.org\/10.1101\/2023.09.26.559540 Posted on bioRxiv September 28, 2023","DOI":"10.1101\/2023.09.26.559540"},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms5982"},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.1016\/j.jbc.2022.102322"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0031274"},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.1210\/me.2008-0183"},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.1016\/j.cmet.2013.11.001"},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.2337\/db13-1869"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.2337\/db20-0572"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1802611115"},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0029092"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.166.5.3256"},{"key":"B26","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2018.02474"},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-021-24384-2"},{"key":"B28","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-15459-7"},{"key":"B29","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2567.2008.02945.x"},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2018.00863"},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.1091\/mbc.e12-06-0481"},{"key":"B32","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-49632-z"},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.1002\/path.5392"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.1161\/CIRCRESAHA.119.315175"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2020.06.003"},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2020.08.004"},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.1016\/j.molcel.2019.07.028"},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2020.108626"},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-019-1631-3"},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.1126\/sciimmunol.add8945"},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.1172\/jci.insight.171701"},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.1126\/science.aau0964"},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-021-99090-6"},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.1172\/JCI200319246"},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.1172\/JCI200319451"},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.1194\/jlr.M800019-JLR200"},{"key":"B47","doi-asserted-by":"publisher","DOI":"10.1023\/A:1008942828960"},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.1182\/blood.V91.2.419"},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2018.00659"},{"key":"B50","doi-asserted-by":"publisher","DOI":"10.3389\/fnut.2022.828522"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.2337\/db22-0192"},{"key":"B52","doi-asserted-by":"publisher","DOI":"10.1002\/oby.20634"},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.2337\/db10-1411"},{"key":"B54","doi-asserted-by":"publisher","DOI":"10.1038\/ni.3728"},{"key":"B55","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-022-04518-2"},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.26508\/lsa.202101286"},{"key":"B57","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2405746121"},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-57368-w"},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2021.719979"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.1194\/jlr.M500294-JLR200"},{"key":"B61","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-52265-8"},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.1194\/jlr.M011338"},{"key":"B63","doi-asserted-by":"publisher","DOI":"10.1016\/j.cmet.2020.11.008"},{"key":"B64","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0163120"},{"key":"B65","doi-asserted-by":"publisher","DOI":"10.3389\/fcvm.2021.757926"},{"key":"B66","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamcr.2015.05.018"},{"key":"B67","doi-asserted-by":"publisher","DOI":"10.1111\/imcb.12616"},{"key":"B68","doi-asserted-by":"publisher","DOI":"10.1530\/JOE-23-0180"},{"key":"B69","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-physiol-060721-092930"},{"key":"B70","doi-asserted-by":"publisher","DOI":"10.1016\/j.bcp.2024.116324"},{"key":"B71","doi-asserted-by":"publisher","DOI":"10.15252\/embr.202152835"},{"key":"B72","doi-asserted-by":"publisher","DOI":"10.1038\/s41420-024-02247-1"},{"key":"B73","doi-asserted-by":"publisher","DOI":"10.1016\/j.cmet.2017.01.010"},{"key":"B74","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-14263-2"},{"key":"B75","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.180.8.5707"},{"key":"B76","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1949.sp004363"},{"key":"B77","doi-asserted-by":"publisher","DOI":"10.1016\/j.cmet.2018.06.019"},{"key":"B78","doi-asserted-by":"publisher","DOI":"10.1038\/s41587-020-0439-x"},{"key":"B79","unstructured":"FastQC: A Quality Control tool for High Throughput Sequence Data; 2010. http:\/\/www.bioinformatics.babraham.ac.uk\/projects\/fastqc\/"},{"key":"B80","doi-asserted-by":"publisher","DOI":"10.14806\/ej.17.1.200"},{"key":"B81","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/bts635"},{"key":"B82","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2105-12-323"},{"key":"B83","unstructured":"Picard. Github Repository. Broad Institute; 2019. http:\/\/broadinstitute.github.io\/picard\/"},{"key":"B84","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btp616"},{"key":"B85","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkae456"}],"container-title":["JCI Insight"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/insight.jci.org\/articles\/view\/193837\/files\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,15]],"date-time":"2026-05-15T19:27:38Z","timestamp":1778873258000},"score":1,"resource":{"primary":{"URL":"https:\/\/insight.jci.org\/articles\/view\/193837"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,10]]},"references-count":85,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2026,3,23]]},"published-print":{"date-parts":[[2026,3,23]]}},"alternative-id":["10.1172\/jci.insight.193837"],"URL":"https:\/\/doi.org\/10.1172\/jci.insight.193837","relation":{},"ISSN":["2379-3708"],"issn-type":[{"value":"2379-3708","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,10]]},"article-number":"e193837"}}