{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,10]],"date-time":"2026-02-10T16:42:36Z","timestamp":1770741756000,"version":"3.49.0"},"reference-count":48,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2021,9,1]],"date-time":"2021-09-01T00:00:00Z","timestamp":1630454400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2021,9,1]],"date-time":"2021-09-01T00:00:00Z","timestamp":1630454400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2021,9,1]],"date-time":"2021-09-01T00:00:00Z","timestamp":1630454400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-017"},{"start":{"date-parts":[[2021,9,1]],"date-time":"2021-09-01T00:00:00Z","timestamp":1630454400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2021,9,1]],"date-time":"2021-09-01T00:00:00Z","timestamp":1630454400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-012"},{"start":{"date-parts":[[2021,9,1]],"date-time":"2021-09-01T00:00:00Z","timestamp":1630454400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2021,9,1]],"date-time":"2021-09-01T00:00:00Z","timestamp":1630454400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-004"}],"funder":[{"DOI":"10.13039\/501100014188","name":"Ministry of Science and ICT, South Korea","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100014188","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003725","name":"National Research Foundation of Korea","doi-asserted-by":"publisher","award":["2019R1A2C2002163"],"award-info":[{"award-number":["2019R1A2C2002163"]}],"id":[{"id":"10.13039\/501100003725","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["The International Journal of Biochemistry & Cell Biology"],"published-print":{"date-parts":[[2021,9]]},"DOI":"10.1016\/j.biocel.2021.106053","type":"journal-article","created":{"date-parts":[[2021,8,8]],"date-time":"2021-08-08T01:28:57Z","timestamp":1628386137000},"page":"106053","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":11,"special_numbering":"C","title":["Loss of lymphocyte cytosolic protein 1 (LCP1) induces browning in 3T3-L1 adipocytes via \u03b23-AR and the ERK-independent signaling pathway"],"prefix":"10.1016","volume":"138","author":[{"given":"Manigandan","family":"Subramani","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2453-2282","authenticated-orcid":false,"given":"Jong Won","family":"Yun","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"78","reference":[{"key":"10.1016\/j.biocel.2021.106053_bib0005","doi-asserted-by":"crossref","first-page":"590","DOI":"10.1002\/hep.26184","article-title":"Association between liver-specific gene polymorphisms and their expression levels with nonalcoholic fatty liver disease","volume":"57","author":"Adams","year":"2013","journal-title":"Hepatology"},{"key":"10.1016\/j.biocel.2021.106053_bib0010","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1016\/j.cmet.2016.08.002","article-title":"Beige adipocyte maintenance is regulated by autophagy-induced mitochondrial clearance","volume":"24","author":"Altshuler-Keylin","year":"2016","journal-title":"Cell Metab."},{"key":"10.1016\/j.biocel.2021.106053_bib0015","first-page":"1","article-title":"Increased mitochondrial and lipid metabolism is a conserved effect of Insulin\/PI3K pathway downregulation in adipose tissue","volume":"1","author":"Bettedi","year":"2020","journal-title":"Sci. Rep."},{"key":"10.1016\/j.biocel.2021.106053_bib0020","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1152\/physrev.00015.2003","article-title":"Brown adipose tissue: function and physiological significance","volume":"84","author":"Cannon","year":"2004","journal-title":"Physiol. Rev."},{"key":"10.1016\/j.biocel.2021.106053_bib0025","doi-asserted-by":"crossref","first-page":"e3060","DOI":"10.1038\/cddis.2017.437","article-title":"AP4 modulated by the PI3K\/AKT pathway promotes prostate cancer proliferation and metastasis of prostate cancer via upregulating L-plastin","volume":"8","author":"Chen","year":"2017","journal-title":"Cell Death Dis."},{"key":"10.1016\/j.biocel.2021.106053_bib0030","first-page":"102","article-title":"\u03b2-Adrenoceptor signaling networks in adipocytes for recruiting stored fat and energy expenditure","volume":"2","author":"Collins","year":"2012","journal-title":"Front. Endocrinol."},{"key":"10.1016\/j.biocel.2021.106053_bib0035","doi-asserted-by":"crossref","first-page":"1509","DOI":"10.1056\/NEJMoa0810780","article-title":"Identification and importance of brown adipose tissue in adult humans","volume":"360","author":"Cypess","year":"2009","journal-title":"N. Engl. J. Med."},{"key":"10.1016\/j.biocel.2021.106053_bib0040","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1186\/1756-0500-5-513","article-title":"A simple protocol for the subcellular fractionation of skeletal muscle cells and tissue","volume":"5","author":"Dimauro","year":"2012","journal-title":"BMC Res. Notes"},{"key":"10.1016\/j.biocel.2021.106053_bib0045","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1016\/j.neuroscience.2020.04.034","article-title":"Recent updates on obesity treatments: available drugs and future directions","volume":"437","author":"Dragano","year":"2020","journal-title":"Neuroscience"},{"key":"10.1016\/j.biocel.2021.106053_bib0050","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1016\/j.cmet.2008.12.014","article-title":"UCP1 ablation nduces obesity andabolishes diet-induced thermogenesis in mice exempt from thermal stress by living at thermoneutrality","volume":"9","author":"Feldmann","year":"2009","journal-title":"Cell Metab."},{"key":"10.1016\/j.biocel.2021.106053_bib0055","first-page":"9072","article-title":"Non-alcoholic fatty liver disease and type 2 diabetes mellitus: the liver disease of our age?","volume":"20","author":"Firneisz","year":"2014","journal-title":"World J. Gastroenterol."},{"key":"10.1016\/j.biocel.2021.106053_bib0060","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1016\/j.tips.2009.05.003","article-title":"BAT: a new target for human obesity?","volume":"30","author":"Fr\u00fchbeck","year":"2009","journal-title":"Trends Pharmacol. Sci."},{"key":"10.1016\/j.biocel.2021.106053_bib0065","doi-asserted-by":"crossref","first-page":"8243","DOI":"10.1074\/jbc.M115.637785","article-title":"Mitochondrial turnover: a phenotype distinguishing brown adipocytes from interscapular brown adipose tissue and white adipose tissue","volume":"290","author":"Gospodarska","year":"2015","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.biocel.2021.106053_bib0070","doi-asserted-by":"crossref","first-page":"4281","DOI":"10.1073\/pnas.1909814117","article-title":"Decision between mitophagy and apoptosis by Parkin via VDAC1 ubiquitination","volume":"117","author":"Ham","year":"2020","journal-title":"PNAS"},{"key":"10.1016\/j.biocel.2021.106053_bib0075","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1038\/nrm3311","article-title":"AMPK: a nutrient and energy sensor that maintains energy homeostasis","volume":"13","author":"Hardie","year":"2012","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"10.1016\/j.biocel.2021.106053_bib0080","doi-asserted-by":"crossref","first-page":"1252","DOI":"10.1038\/nm.3361","article-title":"Brown and beige fat: development, function and therapeutic potential","volume":"19","author":"Harms","year":"2013","journal-title":"Nat. Med."},{"key":"10.1016\/j.biocel.2021.106053_bib0085","doi-asserted-by":"crossref","first-page":"3997","DOI":"10.1096\/fj.201801429RR","article-title":"Stress hormone signaling through \u03b2-adrenergic receptors regulates macrophage mechanotype and function","volume":"33","author":"Kim","year":"2019","journal-title":"FASEB J."},{"key":"10.1016\/j.biocel.2021.106053_bib0090","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1080\/15548627.2015.1100356","article-title":"Guidelines for the use and interpretation of assays for monitoring autophagy","volume":"12","author":"Klionsky","year":"2016","journal-title":"Autophagy"},{"key":"10.1016\/j.biocel.2021.106053_bib0095","doi-asserted-by":"crossref","first-page":"286","DOI":"10.1002\/cm.20442","article-title":"Characterization of L\u2010plastin interaction with beta integrin and its regulation by micro\u2010calpain","volume":"67","author":"Le Goff","year":"2010","journal-title":"Cytoskeleton"},{"key":"10.1016\/j.biocel.2021.106053_bib0100","doi-asserted-by":"crossref","first-page":"2781","DOI":"10.1016\/S0021-9258(18)53842-4","article-title":"Human plastin genes. Comparative gene structure, chromosome location, and differential expression in normal and neoplastic cells","volume":"268","author":"Lin","year":"1993","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.biocel.2021.106053_bib0105","doi-asserted-by":"crossref","first-page":"711","DOI":"10.1042\/BSR20130046","article-title":"Turning WAT into BAT: a review on regulators controlling the browning of white adipocytes","volume":"33","author":"Lo","year":"2013","journal-title":"Biosci. Rep."},{"key":"10.1016\/j.biocel.2021.106053_bib0110","first-page":"2303","article-title":"Identification of polymorphisms associated with hypertriglyceridemia and prolonged survival induced by bexarotene in treating non-small cell lung cancer","volume":"31","author":"Luo","year":"2011","journal-title":"Anticancer Res."},{"key":"10.1016\/j.biocel.2021.106053_bib0115","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1002\/mc.23131","article-title":"LCP1 triggers mTORC2\/AKT activity and is pharmacologically targeted by enzastaurin in hypereosinophilia","volume":"59","author":"Ma","year":"2020","journal-title":"Mol. Carcinog."},{"key":"10.1016\/j.biocel.2021.106053_bib0120","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1007\/s12257-020-0149-8","article-title":"Urolithin A induces brown-like phenotype in 3T3-L1 white adipocytes via \u03b23-adrenergic receptor-p38 MAPK signaling pathway","volume":"25","author":"Manigandan","year":"2020","journal-title":"Biotechnol. Bioprocess Eng."},{"key":"10.1016\/j.biocel.2021.106053_bib0125","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1038\/srep14841","article-title":"Extensive weight loss reveals distinct gene expression changes in human subcutaneous and visceral adipose tissue","volume":"5","author":"Mardinoglu","year":"2015","journal-title":"Sci. Rep."},{"key":"10.1016\/j.biocel.2021.106053_bib0130","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1016\/0022-1759(83)90303-4","article-title":"Rapid colorimetric assay for cellular growth and survival: application to proliferation and cytotoxicity assays","volume":"65","author":"Mosmann","year":"1983","journal-title":"J. Immunol. Methods"},{"key":"10.1016\/j.biocel.2021.106053_bib0135","doi-asserted-by":"crossref","first-page":"1089","DOI":"10.1139\/apnm-2018-0814","article-title":"Novel regulatory roles of carboxylesterase 3 in lipid metabolism and browning in 3T3-L1 white adipocytes","volume":"44","author":"Mukherjee","year":"2019","journal-title":"Appl. Physiol. Nutr. Metab."},{"key":"10.1016\/j.biocel.2021.106053_bib0140","doi-asserted-by":"crossref","DOI":"10.1016\/j.mce.2020.110757","article-title":"Secreted protein acidic and rich in cysteine (SPARC) regulates thermogenesis in white and brown adipocytes","volume":"506","author":"Mukherjee","year":"2020","journal-title":"Mol. Cell. Endocrinol."},{"key":"10.1016\/j.biocel.2021.106053_bib0145","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1007\/s12257-020-0023-8","article-title":"Roles of macrophage colony stimulating factor in white and brown adipocytes","volume":"25","author":"Mukherjee","year":"2020","journal-title":"Biotechnol. Bioprocess Eng."},{"key":"10.1016\/j.biocel.2021.106053_bib0150","doi-asserted-by":"crossref","first-page":"R199","DOI":"10.1530\/JME-13-0277","article-title":"Dissecting adipose tissue lipolysis: molecular regulation and implications for metabolic disease","volume":"52","author":"Nielsen","year":"2014","journal-title":"J. Mol. Endocrinol."},{"key":"10.1016\/j.biocel.2021.106053_bib0155","doi-asserted-by":"crossref","first-page":"2086","DOI":"10.18632\/oncotarget.26747","article-title":"Dual inhibition of ABCE1 and LCP1 by microRNA-96 results in an additive effect in breast cancer mouse model","volume":"10","author":"Pillar","year":"2019","journal-title":"Oncotarget"},{"key":"10.1016\/j.biocel.2021.106053_bib0160","doi-asserted-by":"crossref","first-page":"873","DOI":"10.1074\/mcp.M116.065524","article-title":"Extended multiplexing of tandem mass tags (TMT) labeling reveals age and high fat diet specific proteome changes in mouse epididymal adipose tissue","volume":"16","author":"Plubell","year":"2017","journal-title":"Mol. Cell Proteomics"},{"key":"10.1016\/j.biocel.2021.106053_bib0165","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1038\/ncomms6864","article-title":"Dietary cholesterol directly induces acute inflammasome-dependent intestinal inflammation","volume":"5","author":"Progatzky","year":"2014","journal-title":"Nat. Commun."},{"key":"10.1016\/j.biocel.2021.106053_bib0170","article-title":"Active integrins regulate white adipose tissue insulin sensitivity and brown fat thermogenesis","volume":"30","author":"Ruiz-Ojeda","year":"2020","journal-title":"Mol. Metab."},{"key":"10.1016\/j.biocel.2021.106053_bib0175","first-page":"1","article-title":"Increased apoptosis and browning of TAK1-deficient adipocytes protects against obesity","volume":"19","author":"Sassmann-Schweda","year":"2016","journal-title":"JCI Insight"},{"key":"10.1016\/j.biocel.2021.106053_bib0180","doi-asserted-by":"crossref","first-page":"1537","DOI":"10.2337\/db06-0263","article-title":"Adipose tissue: from lipid storage compartment to endocrine organ","volume":"55","author":"Scherer","year":"2006","journal-title":"Diabetes"},{"key":"10.1016\/j.biocel.2021.106053_bib0185","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1172\/JCI44271","article-title":"Prdm16 determines the thermogenic program of subcutaneous white adipose tissue in mice","volume":"121","author":"Seale","year":"2011","journal-title":"J. Clin. Invest."},{"key":"10.1016\/j.biocel.2021.106053_bib0190","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.jnutbio.2018.09.028","article-title":"Dietary polyphenols and their roles in fat browning","volume":"64","author":"Silvester","year":"2019","journal-title":"J. Nutr. Biochem."},{"key":"10.1016\/j.biocel.2021.106053_bib0195","doi-asserted-by":"crossref","first-page":"531","DOI":"10.1016\/S0092-8674(01)00240-9","article-title":"Obesity and the regulation of energy balance","volume":"104","author":"Spiegelman","year":"2001","journal-title":"Cell"},{"key":"10.1016\/j.biocel.2021.106053_bib0200","doi-asserted-by":"crossref","first-page":"199","DOI":"10.3109\/10409231003667500","article-title":"Genetic control of de novo lipogenesis: role in diet-induced obesity","volume":"45","author":"Strable","year":"2010","journal-title":"Crit. Rev. Biochem. Mol. Biol."},{"key":"10.1016\/j.biocel.2021.106053_bib0205","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1089\/ars.2012.4536","article-title":"Increased mitochondrial activity in BMP7-treated brown adipocytes, due to increased CPT1-and CD36-mediated fatty acid uptake","volume":"19","author":"Townsend","year":"2013","journal-title":"Antioxid. Redox Signal."},{"key":"10.1016\/j.biocel.2021.106053_bib0210","doi-asserted-by":"crossref","first-page":"512","DOI":"10.1016\/j.cmet.2014.06.010","article-title":"The PPAR\u03b1-FGF21 hormone axis contributes to metabolic regulation by the hepatic JNK signaling pathway","volume":"20","author":"Vernia","year":"2014","journal-title":"Cell Metab."},{"key":"10.1016\/j.biocel.2021.106053_bib0215","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1038\/s41598-019-40992-x","article-title":"A comparison between the abdominal and femoral adipose tissue proteome of overweight and obese women","volume":"9","author":"Vogel","year":"2019","journal-title":"Sci. Rep."},{"key":"10.1016\/j.biocel.2021.106053_bib0220","doi-asserted-by":"crossref","first-page":"691","DOI":"10.1038\/nrm.2016.96","article-title":"Control of brown and beige fat development","volume":"17","author":"Wang","year":"2016","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"10.1016\/j.biocel.2021.106053_bib0225","doi-asserted-by":"crossref","first-page":"696","DOI":"10.1016\/j.bbrc.2015.08.049","article-title":"Sulforaphane induces apoptosis in adipocytes via Akt\/p70s6k1\/Bad inhibition and ERK activation","volume":"465","author":"Yao","year":"2015","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"10.1016\/j.biocel.2021.106053_bib0230","doi-asserted-by":"crossref","first-page":"706","DOI":"10.3390\/cells9030706","article-title":"Role of c-Jun N-terminal kinase (JNK) in obesity and type 2 diabetes","volume":"9","author":"Yung","year":"2020","journal-title":"Cells"},{"key":"10.1016\/j.biocel.2021.106053_bib0235","doi-asserted-by":"crossref","first-page":"514","DOI":"10.2337\/db13-1106","article-title":"Irisin stimulates browning of white adipocytes through mitogen-activated protein kinase p38 MAP kinase and ERK MAP kinase signaling","volume":"63","author":"Zhang","year":"2014","journal-title":"Diabetes"},{"key":"10.1016\/j.biocel.2021.106053_bib0240","doi-asserted-by":"crossref","first-page":"347","DOI":"10.33549\/physiolres.933650","article-title":"Transcription regulators and hormones involved in the development of brown fat and white fat browning: transcriptional and hormonal control of brown\/beige fat development","volume":"67","author":"Zhang","year":"2018","journal-title":"Physiol. Res."}],"container-title":["The International Journal of Biochemistry & Cell Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1357272521001333?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1357272521001333?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2025,10,23]],"date-time":"2025-10-23T07:19:14Z","timestamp":1761203954000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S1357272521001333"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,9]]},"references-count":48,"alternative-id":["S1357272521001333"],"URL":"https:\/\/doi.org\/10.1016\/j.biocel.2021.106053","relation":{},"ISSN":["1357-2725"],"issn-type":[{"value":"1357-2725","type":"print"}],"subject":[],"published":{"date-parts":[[2021,9]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Loss of lymphocyte cytosolic protein 1 (LCP1) induces browning in 3T3-L1 adipocytes via \u03b23-AR and the ERK-independent signaling pathway","name":"articletitle","label":"Article Title"},{"value":"The International Journal of Biochemistry & Cell Biology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.biocel.2021.106053","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2021 Elsevier Ltd. All rights reserved.","name":"copyright","label":"Copyright"}],"article-number":"106053"}}