{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T21:38:30Z","timestamp":1792791510420,"version":"4.5.0"},"reference-count":51,"publisher":"Springer Science and Business Media LLC","issue":"18","license":[{"start":{"date-parts":[[2025,8,1]],"date-time":"2025-08-01T00:00:00Z","timestamp":1754006400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2025,8,1]],"date-time":"2025-08-01T00:00:00Z","timestamp":1754006400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"government of india","id":[{"id":"https:\/\/ror.org\/036h6g940","id-type":"ROR","asserted-by":"crossref"}]},{"DOI":"10.13039\/100008732","name":"Uehara Memorial Foundation","doi-asserted-by":"publisher","award":["201830047"],"award-info":[{"award-number":["201830047"]}],"id":[{"id":"10.13039\/100008732","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100005156","name":"Alexander von Humboldt-Stiftung","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100005156","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004543","name":"China Scholarship Council","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100004543","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002347","name":"Federal Ministry of Education and Research","doi-asserted-by":"crossref","award":["01KX1012"],"award-info":[{"award-number":["01KX1012"]}],"id":[{"id":"10.13039\/501100002347","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["209933838"],"award-info":[{"award-number":["209933838"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100031584","name":"Deutsches Zentrum f\u00fcr Diabetesforschung","doi-asserted-by":"publisher","award":["82DZD00601"],"award-info":[{"award-number":["82DZD00601"]}],"id":[{"id":"10.13039\/100031584","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100018694","name":"EC | Horizon Europe | Excellent Science | HORIZON EUROPE Marie Sklodowska-Curie Actions","doi-asserted-by":"publisher","award":["101120466"],"award-info":[{"award-number":["101120466"]}],"id":[{"id":"10.13039\/100018694","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["EMBO J"],"published-print":{"date-parts":[[2025,9,15]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Adipose tissue is a central organiser of systemic lipid homeostasis and a pharmacological target in obesity, orchestrating cellular responses to environmental cues. Nutritionally regulated adipose and cardiac enriched protein (NRAC) is a small adipocyte-specific transmembrane protein with unknown function. Here, we show that Nrac directly interacts with scavenger receptor CD36 via its first transmembrane domain. Forming a complex with CD36 and caveolin-1 under low extracellular fatty acid (FA) concentrations, NRAC modulates CD36-dependent fatty acid uptake in adipocytes. Upon increase in extracellular FA levels, NRAC is ubiquitinated and internalised, leading to CD36\u2019s dissociation from caveolin-1 and clathrin-mediated endocytosis. This results in increased fatty acid uptake into fat cells, adipocyte hypertrophy, increased fat mass and elevated lipid clearance from the blood in chow-diet-fed mice. Finally, human\n                    <jats:italic>NRAC<\/jats:italic>\n                    expression and the intronic SNP rs12878589 are associated with body fat distribution and obesity. Together, these findings reveal a novel regulatory mechanism by which adipocytes sense and respond to extracellular fatty acid availability to fine-tune lipid uptake and storage at cellular and organismal level.\n                  <\/jats:p>","DOI":"10.1038\/s44318-025-00520-2","type":"journal-article","created":{"date-parts":[[2025,8,1]],"date-time":"2025-08-01T09:40:20Z","timestamp":1754041220000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":7,"title":["NRAC controls CD36-mediated fatty acid uptake in adipocytes and lipid clearance in vivo"],"prefix":"10.1038","volume":"44","author":[{"given":"Inderjeet","family":"Singh","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7950-8952","authenticated-orcid":false,"given":"Yasuhiro","family":"Onogi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7630-5447","authenticated-orcid":false,"given":"Filipe","family":"Menezes","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dina","family":"Khasanova","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lingru","family":"Kang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chenxi","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-5752-7527","authenticated-orcid":false,"given":"Julio","family":"Ruiz-Trave","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1039-8879","authenticated-orcid":false,"given":"Sapna","family":"Sharma","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-2204-8636","authenticated-orcid":false,"given":"Ahmed","family":"Khalil","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0004-4115-6201","authenticated-orcid":false,"given":"Valentin K","family":"Reichenbach","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yingzi","family":"Shi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrew","family":"Flatley","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaocheng","family":"Yan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andreas","family":"Israel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6856-5468","authenticated-orcid":false,"given":"Nathalia R V","family":"Dragano","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Juan Antonio","family":"Aguilar-Pimentel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anne","family":"Hoffmann","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5160-4571","authenticated-orcid":false,"given":"Adhideb","family":"Ghosh","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-3360-5455","authenticated-orcid":false,"given":"Falko","family":"No\u00e9","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3862-6805","authenticated-orcid":false,"given":"Christian","family":"Wolfrum","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sebastian","family":"Cucuruz","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ann-Christine","family":"K\u00f6nig","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0635-3507","authenticated-orcid":false,"given":"Ingo","family":"Burtscher","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1630-6827","authenticated-orcid":false,"given":"Stefanie M","family":"Hauck","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Heiko","family":"Lickert","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7682-5840","authenticated-orcid":false,"given":"Susanna M","family":"Hofmann","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3981-367X","authenticated-orcid":false,"given":"Regina","family":"Feederle","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5461-3214","authenticated-orcid":false,"given":"Sonja C","family":"Schriever","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7065-2862","authenticated-orcid":false,"given":"Rene","family":"Hernandez-Bautista","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2098-0654","authenticated-orcid":false,"given":"Gencer","family":"Sancar","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alberto","family":"Cebrian-Serrano","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6855-0012","authenticated-orcid":false,"given":"Igor","family":"Tetko","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Helmut","family":"Fuchs","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Val\u00e9rie","family":"Gailus-Durner","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Matthias","family":"Bl\u00fcher","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Martin","family":"Hrab\u011b de Angelis","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7575-0920","authenticated-orcid":false,"given":"Siegfried","family":"Ussar","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,8,1]]},"reference":[{"key":"520_CR1","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1016\/j.softx.2015.06.001","volume":"1-2","author":"MJ Abraham","year":"2015","unstructured":"Abraham MJ, Murtola T, Schulz R, P\u00e1ll S, Smith JC, Hess B, Lindahl E (2015) GROMACS: high performance molecular simulations through multi-level parallelism from laptops to supercomputers. SoftwareX 1-2:19\u201325","journal-title":"SoftwareX"},{"key":"520_CR2","doi-asserted-by":"crossref","first-page":"2828","DOI":"10.2337\/db14-1409","volume":"64","author":"AH Ali","year":"2015","unstructured":"Ali AH, Mundi M, Koutsari C, Bernlohr DA, Jensen MD (2015) Adipose tissue free fatty acid storage in vivo: effects of insulin versus niacin as a control for suppression of lipolysis. Diabetes 64:2828\u20132835","journal-title":"Diabetes"},{"key":"520_CR3","doi-asserted-by":"crossref","first-page":"585","DOI":"10.1111\/j.1463-1326.2006.00558.x","volume":"8","author":"AH Bakker","year":"2006","unstructured":"Bakker AH, Nijhuis J, Buurman WA, van Dielen FM, Greve JW (2006) Low number of omental preadipocytes with high leptin and low adiponectin secretion is associated with high fasting plasma glucose levels in obese subjects. Diab Obes Metab 8:585\u2013588","journal-title":"Diab Obes Metab"},{"key":"520_CR4","doi-asserted-by":"crossref","first-page":"200","DOI":"10.1038\/nm.2297","volume":"17","author":"A Bartelt","year":"2011","unstructured":"Bartelt A, Bruns OT, Reimer R, Hohenberg H, Ittrich H, Peldschus K, Kaul MG, Tromsdorf UI, Weller H, Waurisch C et al (2011) Brown adipose tissue activity controls triglyceride clearance. Nat Med 17:200\u2013205","journal-title":"Nat Med"},{"key":"520_CR5","doi-asserted-by":"crossref","first-page":"bnaa004","DOI":"10.1210\/endrev\/bnaa004","volume":"41","author":"M Bluher","year":"2020","unstructured":"Bluher M (2020) Metabolically healthy obesity. Endocr Rev 41:bnaa004","journal-title":"Endocr Rev"},{"key":"520_CR6","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1038\/s41569-022-00676-y","volume":"19","author":"J Boren","year":"2022","unstructured":"Boren J, Taskinen MR, Bjornson E, Packard CJ (2022) Metabolism of triglyceride-rich lipoproteins in health and dyslipidaemia. Nat Rev Cardiol 19:577\u2013592","journal-title":"Nat Rev Cardiol"},{"key":"520_CR7","doi-asserted-by":"crossref","DOI":"10.1038\/s41467-022-34532-x","volume":"13","author":"PM Brailey","year":"2022","unstructured":"Brailey PM, Evans L, Lopez-Rodriguez JC, Sinadinos A, Tyrrel V, Kelly G, O\u2019Donnell V, Ghazal P, John S, Barral P (2022) CD1d-dependent rewiring of lipid metabolism in macrophages regulates innate immune responses. Nat Commun 13:6723","journal-title":"Nat Commun"},{"key":"520_CR8","doi-asserted-by":"crossref","first-page":"525","DOI":"10.1038\/nbt.3519","volume":"34","author":"NL Bray","year":"2016","unstructured":"Bray NL, Pimentel H, Melsted P, Pachter L (2016) Near-optimal probabilistic RNA-seq quantification. Nat Biotechnol 34:525\u2013527","journal-title":"Nat Biotechnol"},{"key":"520_CR9","doi-asserted-by":"crossref","first-page":"D609","DOI":"10.1093\/nar\/gkae1010","volume":"53","author":"TU Consortium","year":"2024","unstructured":"Consortium TU (2024) UniProt: the Universal Protein Knowledgebase in 2025. Nucleic Acids Res 53:D609\u2013D617","journal-title":"Nucleic Acids Res"},{"key":"520_CR10","doi-asserted-by":"crossref","first-page":"e147057","DOI":"10.1172\/jci.insight.147057","volume":"6","author":"AC Daquinag","year":"2021","unstructured":"Daquinag AC, Gao Z, Fussell C, Immaraj L, Pasqualini R, Arap W, Akimzhanov AM, Febbraio M, Kolonin MG (2021) Fatty acid mobilization from adipose tissue is mediated by CD36 posttranslational modifications and intracellular trafficking. JCI Insight 6:e147057","journal-title":"JCI Insight"},{"issue":"Suppl","key":"520_CR11","doi-asserted-by":"crossref","first-page":"S86","DOI":"10.1194\/jlr.R800085-JLR200","volume":"50","author":"IJ Goldberg","year":"2009","unstructured":"Goldberg IJ, Eckel RH, Abumrad NA (2009) Regulation of fatty acid uptake into tissues: lipoprotein lipase- and CD36-mediated pathways. J Lipid Res 50(Suppl):S86\u201390","journal-title":"J Lipid Res"},{"key":"520_CR12","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1186\/1743-7075-4-22","volume":"4","author":"AM Gonzales","year":"2007","unstructured":"Gonzales AM, Orlando RA (2007) Role of adipocyte-derived lipoprotein lipase in adipocyte hypertrophy. Nutr Metab 4:22","journal-title":"Nutr Metab"},{"key":"520_CR13","doi-asserted-by":"crossref","first-page":"1209","DOI":"10.1111\/eci.12519","volume":"45","author":"SM Grundy","year":"2015","unstructured":"Grundy SM (2015) Adipose tissue and metabolic syndrome: too much, too little or neither. Eur J Clin Invest 45:1209\u20131217","journal-title":"Eur J Clin Invest"},{"key":"520_CR14","doi-asserted-by":"crossref","first-page":"2806","DOI":"10.3390\/biomedicines11102806","volume":"11","author":"T Hagemann","year":"2023","unstructured":"Hagemann T, Czechowski P, Ghosh A, Sun W, Dong H, Noe F, Wolfrum C, Bluher M, Hoffmann A (2023) Laminin alpha4 expression in human adipose tissue depots and its association with obesity and obesity related traits. Biomedicines 11:2806","journal-title":"Biomedicines"},{"key":"520_CR15","doi-asserted-by":"crossref","unstructured":"Hallgren J, Tsirigos KD, Pedersen MD, Almagro Armenteros JJ, Marcatili P, Nielsen H, Krogh A, Winther O (2022) DeepTMHMM predicts alpha and beta transmembrane proteins using deep neural networks. Preprint at https:\/\/www.biorxiv.org\/content\/10.1101\/2022.04.08.487609v1","DOI":"10.1101\/2022.04.08.487609"},{"key":"520_CR16","volume":"11","author":"JW Hao","year":"2020","unstructured":"Hao JW, Wang J, Guo H, Zhao YY, Sun HH, Li YF, Lai XY, Zhao N, Wang X, Xie C et al (2020) CD36 facilitates fatty acid uptake by dynamic palmitoylation-regulated endocytosis. Nat Commun 11:4765","journal-title":"Nat Commun"},{"key":"520_CR17","doi-asserted-by":"crossref","first-page":"5","DOI":"10.2337\/db07-0403","volume":"57","author":"SM Hofmann","year":"2008","unstructured":"Hofmann SM, Perez-Tilve D, Greer TM, Coburn BA, Grant E, Basford JE, Tschop MH, Hui DY (2008) Defective lipid delivery modulates glucose tolerance and metabolic response to diet in apolipoprotein E-deficient mice. Diabetes 57:5\u201312","journal-title":"Diabetes"},{"key":"520_CR18","doi-asserted-by":"crossref","first-page":"3219","DOI":"10.1038\/s41596-024-01011-0","volume":"19","author":"RV Honorato","year":"2024","unstructured":"Honorato RV, Trellet ME, Jimenez-Garcia B, Schaarschmidt JJ, Giulini M, Reys V, Koukos PI, Rodrigues J, Karaca E, van Zundert GCP et al (2024) The HADDOCK2.4 web server for integrative modeling of biomolecular complexes. Nat Protoc 19:3219\u20133241","journal-title":"Nat Protoc"},{"key":"520_CR19","volume":"7","author":"FL Hsieh","year":"2016","unstructured":"Hsieh FL, Turner L, Bolla JR, Robinson CV, Lavstsen T, Higgins MK (2016) The structural basis for CD36 binding by the malaria parasite. Nat Commun 7:12837","journal-title":"Nat Commun"},{"key":"520_CR20","doi-asserted-by":"crossref","first-page":"2135","DOI":"10.1002\/jcc.23354","volume":"34","author":"J Huang","year":"2013","unstructured":"Huang J, MacKerell AD Jr (2013) CHARMM36 all-atom additive protein force field: validation based on comparison to NMR data. J Comput Chem 34:2135\u20132145","journal-title":"J Comput Chem"},{"key":"520_CR21","volume":"14","author":"DW Jeong","year":"2023","unstructured":"Jeong DW, Park JW, Kim KS, Kim J, Huh J, Seo J, Kim YL, Cho JY, Lee KW, Fukuda J, Chun YS (2023) Palmitoylation-driven PHF2 ubiquitination remodels lipid metabolism through the SREBP1c axis in hepatocellular carcinoma. Nat Commun 14:6370","journal-title":"Nat Commun"},{"key":"520_CR22","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1038\/s41586-021-03819-2","volume":"596","author":"J Jumper","year":"2021","unstructured":"Jumper J, Evans R, Pritzel A, Green T, Figurnov M, Ronneberger O, Tunyasuvunakool K, Bates R, Zidek A, Potapenko A et al (2021) Highly accurate protein structure prediction with AlphaFold. Nature 596:583\u2013589","journal-title":"Nature"},{"key":"520_CR23","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1016\/j.molmet.2019.04.009","volume":"25","author":"AG Kerr","year":"2019","unstructured":"Kerr AG, Sinha I, Dadvar S, Arner P, Dahlman I (2019) Epigenetic regulation of diabetogenic adipose morphology. Mol Metab 25:159\u2013167","journal-title":"Mol Metab"},{"key":"520_CR24","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/j.cmet.2021.12.012","volume":"34","author":"S Klein","year":"2022","unstructured":"Klein S, Gastaldelli A, Yki-Jarvinen H, Scherer PE (2022) Why does obesity cause diabetes? Cell Metab 34:11\u201320","journal-title":"Cell Metab"},{"key":"520_CR25","doi-asserted-by":"crossref","first-page":"E506","DOI":"10.1152\/ajpendo.00586.2009","volume":"299","author":"N Kloting","year":"2010","unstructured":"Kloting N, Fasshauer M, Dietrich A, Kovacs P, Schon MR, Kern M, Stumvoll M, Bluher M (2010) Insulin-sensitive obesity. Am J Physiol Endocrinol Metab 299:E506\u2013515","journal-title":"Am J Physiol Endocrinol Metab"},{"key":"520_CR26","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1038\/256495a0","volume":"256","author":"G Kohler","year":"1975","unstructured":"Kohler G, Milstein C (1975) Continuous cultures of fused cells secreting antibody of predefined specificity. Nature 256:495\u2013497","journal-title":"Nature"},{"key":"520_CR27","doi-asserted-by":"crossref","first-page":"508","DOI":"10.1038\/s41586-022-05473-8","volume":"613","author":"MI Kurki","year":"2023","unstructured":"Kurki MI, Karjalainen J, Palta P, Sipila TP, Kristiansson K, Donner KM, Reeve MP, Laivuori H, Aavikko M, Kaunisto MA et al (2023) FinnGen provides genetic insights from a well-phenotyped isolated population. Nature 613:508\u2013518","journal-title":"Nature"},{"key":"520_CR28","doi-asserted-by":"crossref","first-page":"475","DOI":"10.1159\/000494295","volume":"11","author":"J Langhardt","year":"2018","unstructured":"Langhardt J, Flehmig G, Kloting N, Lehmann S, Ebert T, Kern M, Schon MR, Gartner D, Lohmann T, Dressler M et al (2018) Effects of weight loss on glutathione peroxidase 3 serum concentrations and adipose tissue expression in human obesity. Obes Facts 11:475\u2013490","journal-title":"Obes Facts"},{"key":"520_CR29","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1021\/acs.jctc.5b00935","volume":"12","author":"J Lee","year":"2016","unstructured":"Lee J, Cheng X, Swails JM, Yeom MS, Eastman PK, Lemkul JA, Wei S, Buckner J, Jeong JC, Qi Y et al (2016) CHARMM-GUI input generator for NAMD, GROMACS, AMBER, OpenMM, and CHARMM\/OpenMM simulations using the CHARMM36 additive force field. J Chem Theory Comput 12:405\u2013413","journal-title":"J Chem Theory Comput"},{"key":"520_CR30","doi-asserted-by":"crossref","first-page":"5068","DOI":"10.33011\/livecoms.1.1.5068","volume":"1","author":"JA Lemkul","year":"2018","unstructured":"Lemkul JA (2018) From proteins to perturbed hamiltonians: a suite of tutorials for the GROMACS-2018 molecular simulation package [Article v1.0]. Living J Comput Mol Sci 1:5068","journal-title":"Living J Comput Mol Sci"},{"key":"520_CR31","doi-asserted-by":"crossref","first-page":"1069","DOI":"10.15252\/embr.201438775","volume":"15","author":"Y Li","year":"2014","unstructured":"Li Y, Fromme T, Schweizer S, Schottl T, Klingenspor M (2014) Taking control over intracellular fatty acid levels is essential for the analysis of thermogenic function in cultured primary brown and brite\/beige adipocytes. EMBO Rep 15:1069\u20131076","journal-title":"EMBO Rep"},{"key":"520_CR32","doi-asserted-by":"crossref","first-page":"D370","DOI":"10.1093\/nar\/gkr703","volume":"40","author":"MA Lomize","year":"2012","unstructured":"Lomize MA, Pogozheva ID, Joo H, Mosberg HI, Lomize AL (2012) OPM database and PPM web server: resources for positioning of proteins in membranes. Nucleic Acids Res 40:D370\u2013376","journal-title":"Nucleic Acids Res"},{"key":"520_CR33","doi-asserted-by":"crossref","DOI":"10.1038\/srep14841","volume":"5","author":"A Mardinoglu","year":"2015","unstructured":"Mardinoglu A, Heiker JT, Gartner D, Bjornson E, Schon MR, Flehmig G, Kloting N, Krohn K, Fasshauer M, Stumvoll M et al (2015) Extensive weight loss reveals distinct gene expression changes in human subcutaneous and visceral adipose tissue. Sci Rep 5:14841","journal-title":"Sci Rep"},{"key":"520_CR51","doi-asserted-by":"crossref","unstructured":"Mauvais-Jarvis F, Clegg DJ, Hevener AL (2013) The role of estrogens in control of energy balance and glucose homeostasis. Endocr Rev 34:309\u2013338","DOI":"10.1210\/er.2012-1055"},{"key":"520_CR34","volume":"32","author":"EC Meng","year":"2023","unstructured":"Meng EC, Goddard TD, Pettersen EF, Couch GS, Pearson ZJ, Morris JH, Ferrin TE (2023) UCSF ChimeraX: tools for structure building and analysis. Protein Sci 32:e4792","journal-title":"Protein Sci"},{"key":"520_CR35","doi-asserted-by":"crossref","first-page":"679","DOI":"10.1038\/s41592-022-01488-1","volume":"19","author":"M Mirdita","year":"2022","unstructured":"Mirdita M, Schutze K, Moriwaki Y, Heo L, Ovchinnikov S, Steinegger M (2022) ColabFold: making protein folding accessible to all. Nat Methods 19:679\u2013682","journal-title":"Nat Methods"},{"key":"520_CR36","doi-asserted-by":"crossref","first-page":"276","DOI":"10.1038\/s41574-021-00471-8","volume":"17","author":"P Morigny","year":"2021","unstructured":"Morigny P, Boucher J, Arner P, Langin D (2021) Lipid and glucose metabolism in white adipocytes: pathways, dysfunction and therapeutics. Nat Rev Endocrinol 17:276\u2013295","journal-title":"Nat Rev Endocrinol"},{"key":"520_CR37","doi-asserted-by":"crossref","first-page":"2509","DOI":"10.1042\/BCJ20190462","volume":"477","author":"Y Onogi","year":"2020","unstructured":"Onogi Y, Khalil A, Ussar S (2020) Identification and characterization of adipose surface epitopes. Biochem J 477:2509\u20132541","journal-title":"Biochem J"},{"key":"520_CR38","doi-asserted-by":"crossref","DOI":"10.1038\/s41467-023-39752-3","volume":"14","author":"VS Peche","year":"2023","unstructured":"Peche VS, Pietka TA, Jacome-Sosa M, Samovski D, Palacios H, Chatterjee-Basu G, Dudley AC, Beatty W, Meyer GA, Goldberg IJ, Abumrad NA (2023) Endothelial cell CD36 regulates membrane ceramide formation, exosome fatty acid transfer and circulating fatty acid levels. Nat Commun 14:4029","journal-title":"Nat Commun"},{"key":"520_CR39","doi-asserted-by":"crossref","first-page":"e88535","DOI":"10.1371\/journal.pone.0088535","volume":"9","author":"JM Peterson","year":"2014","unstructured":"Peterson JM, Seldin MM, Tan SY, Wong GW (2014) CTRP2 overexpression improves insulin and lipid tolerance in diet-induced obese mice. PLoS ONE 9:e88535","journal-title":"PLoS ONE"},{"key":"520_CR40","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1038\/nprot.2014.006","volume":"9","author":"S Picelli","year":"2014","unstructured":"Picelli S, Faridani OR, Bjorklund AK, Winberg G, Sagasser S, Sandberg R (2014) Full-length RNA-seq from single cells using Smart-seq2. Nat Protoc 9:171\u2013181","journal-title":"Nat Protoc"},{"key":"520_CR41","doi-asserted-by":"crossref","first-page":"633","DOI":"10.1038\/nrendo.2017.90","volume":"13","author":"SM Reilly","year":"2017","unstructured":"Reilly SM, Saltiel AR (2017) Adapting to obesity with adipose tissue inflammation. Nat Rev Endocrinol 13:633\u2013643","journal-title":"Nat Rev Endocrinol"},{"key":"520_CR42","volume":"19","author":"Y Song","year":"2018","unstructured":"Song Y, Milon B, Ott S, Zhao X, Sadzewicz L, Shetty A, Boger ET, Tallon LJ, Morell RJ, Mahurkar A, Hertzano R (2018) A comparative analysis of library prep approaches for sequencing low input translatome samples. BMC Genomics 19:696","journal-title":"BMC Genomics"},{"key":"520_CR43","doi-asserted-by":"crossref","first-page":"W228","DOI":"10.1093\/nar\/gkac278","volume":"50","author":"V Thumuluri","year":"2022","unstructured":"Thumuluri V, Almagro Armenteros JJ, Johansen AR, Nielsen H, Winther O (2022) DeepLoc 2.0: multi-label subcellular localization prediction using protein language models. Nucleic Acids Res 50:W228\u2013W234","journal-title":"Nucleic Acids Res"},{"key":"520_CR44","doi-asserted-by":"crossref","first-page":"4587","DOI":"10.1021\/acs.jcim.3c01835","volume":"64","author":"L Urvas","year":"2024","unstructured":"Urvas L, Chiesa L, Bret G, Jacquemard C, Kellenberger E (2024) Benchmarking AlphaFold-generated structures of chemokine-chemokine receptor complexes. J Chem Inf Model 64:4587\u20134600","journal-title":"J Chem Inf Model"},{"key":"520_CR45","doi-asserted-by":"crossref","first-page":"247ra103","DOI":"10.1126\/scitranslmed.3008490","volume":"6","author":"S Ussar","year":"2014","unstructured":"Ussar S, Lee KY, Dankel SN, Boucher J, Haering MF, Kleinridders A, Thomou T, Xue R, Macotela Y, Cypess AM et al (2014) ASC-1, PAT2, and P2RX5 are cell surface markers for white, beige, and brown adipocytes. Sci Transl Med 6:247ra103","journal-title":"Sci Transl Med"},{"key":"520_CR46","doi-asserted-by":"crossref","DOI":"10.1016\/j.xpro.2023.102693","volume":"4","author":"T Villanueva-Carmona","year":"2023","unstructured":"Villanueva-Carmona T, Cedo L, Nunez-Roa C, Maymo-Masip E, Vendrell J, Fernandez-Veledo S (2023) Protocol for the in vitro isolation and culture of mature adipocytes and white adipose tissue explants from humans and mice. STAR Protoc 4:102693","journal-title":"STAR Protoc"},{"key":"520_CR47","doi-asserted-by":"crossref","first-page":"384","DOI":"10.1016\/j.devcel.2023.12.005","volume":"59","author":"J Wang","year":"2024","unstructured":"Wang J, Li DL, Zheng LF, Ren S, Huang ZQ, Tao Y, Liu Z, Shang Y, Pang D, Guo H et al (2024) Dynamic palmitoylation of STX11 controls injury-induced fatty acid uptake to promote muscle regeneration. Dev Cell 59:384\u2013399.e385","journal-title":"Dev Cell"},{"key":"520_CR48","doi-asserted-by":"crossref","first-page":"btac749","DOI":"10.1093\/bioinformatics\/btac749","volume":"39","author":"D Yu","year":"2023","unstructured":"Yu D, Chojnowski G, Rosenthal M, Kosinski J (2023) AlphaPulldown\u2014a Python package for protein-protein interaction screens using AlphaFold-Multimer. Bioinformatics 39:btac749","journal-title":"Bioinformatics"},{"key":"520_CR49","doi-asserted-by":"crossref","first-page":"45931","DOI":"10.1074\/jbc.M307722200","volume":"278","author":"Y Zeng","year":"2003","unstructured":"Zeng Y, Tao N, Chung KN, Heuser JE, Lublin DM (2003) Endocytosis of oxidized low density lipoprotein through scavenger receptor CD36 utilizes a lipid raft pathway that does not require caveolin-1. J Biol Chem 278:45931\u201345936","journal-title":"J Biol Chem"},{"key":"520_CR50","doi-asserted-by":"crossref","first-page":"e46254","DOI":"10.1371\/journal.pone.0046254","volume":"7","author":"R Zhang","year":"2012","unstructured":"Zhang R, Yao F, Gao F, Abou-Samra AB (2012) Nrac, a novel nutritionally-regulated adipose and cardiac-enriched gene. PLoS ONE 7:e46254","journal-title":"PLoS ONE"}],"container-title":["The EMBO Journal"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1038\/s44318-025-00520-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1038\/s44318-025-00520-2","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1038\/s44318-025-00520-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T12:25:06Z","timestamp":1786969506000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1038\/s44318-025-00520-2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8,1]]},"references-count":51,"journal-issue":{"issue":"18","published-print":{"date-parts":[[2025,9,15]]}},"alternative-id":["520"],"URL":"https:\/\/doi.org\/10.1038\/s44318-025-00520-2","relation":{},"ISSN":["0261-4189","1460-2075"],"issn-type":[{"value":"0261-4189","type":"print"},{"value":"1460-2075","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,8,1]]},"assertion":[{"value":"4 November 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"15 July 2025","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"17 July 2025","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"1 August 2025","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"MB received honoraria as a consultant and speaker from Amgen, AstraZeneca, Bayer, Boehringer-Ingelheim, Lilly, Novo Nordisk, Novartis, and Sanofi. All other authors declare no conflict of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Disclosure and competing interests statement"}}],"article-number":"5"}}