{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T16:20:39Z","timestamp":1791822039547,"version":"4.3.4"},"reference-count":47,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2020,6,1]],"date-time":"2020-06-01T00:00:00Z","timestamp":1590969600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2020,6,1]],"date-time":"2020-06-01T00:00:00Z","timestamp":1590969600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"}],"funder":[{"name":"IFCAH-ESPE"}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["The Journal of Steroid Biochemistry and Molecular Biology"],"published-print":{"date-parts":[[2020,6]]},"DOI":"10.1016\/j.jsbmb.2020.105684","type":"journal-article","created":{"date-parts":[[2020,4,28]],"date-time":"2020-04-28T02:44:34Z","timestamp":1588041874000},"page":"105684","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":9,"special_numbering":"C","title":["Metformin inhibits the activation of melanocortin receptors 2 and 3 in vitro: A possible mechanism for its anti-androgenic and weight balancing effects in vivo?"],"prefix":"10.1016","volume":"200","author":[{"given":"Shaheena","family":"Parween","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Silvia","family":"Rihs","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christa E.","family":"Fl\u00fcck","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.jsbmb.2020.105684_bib0005","doi-asserted-by":"crossref","first-page":"2532","DOI":"10.2174\/1381612822666161201152941","article-title":"Is metformin a perfect drug? Updates in pharmacokinetics and pharmacodynamics","volume":"23","author":"Markowicz-Piasecka","year":"2017","journal-title":"Curr. Pharm. Des."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0010","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1007\/s11892-017-0829-8","article-title":"Metformin: new preparations and nonglycemic benefits","volume":"17","author":"Fujita","year":"2017","journal-title":"Curr. Diab. Rep."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0015","doi-asserted-by":"crossref","first-page":"482","DOI":"10.5603\/EP.2017.0050","article-title":"Metformin - a new old drug","volume":"68","author":"Wrobel","year":"2017","journal-title":"Endokrynol. Pol."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0020","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1530\/EJE-09-0733","article-title":"Metformin: an old medication of new fashion: evolving new molecular mechanisms and clinical implications in polycystic ovary syndrome","volume":"162","author":"Diamanti-Kandarakis","year":"2010","journal-title":"Eur. J. Endocrinol."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0025","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1146\/annurev-med-062613-093128","article-title":"Metformin in cancer treatment and prevention","volume":"66","author":"Morales","year":"2015","journal-title":"Annu. Rev. Med."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0030","doi-asserted-by":"crossref","first-page":"657","DOI":"10.1016\/j.numecd.2017.04.009","article-title":"Metformin effects on the heart and the cardiovascular system: a review of experimental and clinical data","volume":"27","author":"Nesti","year":"2017","journal-title":"Nutr. Metab. Cardiovasc. Dis."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0035","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1007\/s11934-017-0693-8","article-title":"Metformin and prostate cancer: a new role for an old drug","volume":"18","author":"Whitburn","year":"2017","journal-title":"Curr. Urol. Rep."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0040","doi-asserted-by":"crossref","DOI":"10.1101\/cshperspect.a025932","article-title":"Metformin: a hopeful promise in aging research","volume":"6","author":"Novelle","year":"2016","journal-title":"Cold Spring Harb. Perspect. Med."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0045","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1016\/j.tem.2019.07.015","article-title":"Metformin as anti-aging therapy: is it for everyone?","volume":"30","author":"Soukas","year":"2019","journal-title":"Trends Endocrinol. Metab."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0050","doi-asserted-by":"crossref","first-page":"1577","DOI":"10.1007\/s00125-017-4342-z","article-title":"The mechanisms of action of metformin","volume":"60","author":"Rena","year":"2017","journal-title":"Diabetologia"},{"key":"10.1016\/j.jsbmb.2020.105684_bib0055","doi-asserted-by":"crossref","first-page":"156","DOI":"10.1007\/s13679-019-00335-3","article-title":"Metformin: mechanisms in human obesity and weight loss","volume":"8","author":"Yerevanian","year":"2019","journal-title":"Curr. Obes. Rep."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0060","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1038\/nature15766","article-title":"Disentangling type 2 diabetes and metformin treatment signatures in the human gut microbiota","volume":"528","author":"Forslund","year":"2015","journal-title":"Nature"},{"key":"10.1016\/j.jsbmb.2020.105684_bib0065","doi-asserted-by":"crossref","first-page":"850","DOI":"10.1038\/nm.4345","article-title":"Metformin alters the gut microbiome of individuals with treatment-naive type 2 diabetes, contributing to the therapeutic effects of the drug","volume":"23","author":"Wu","year":"2017","journal-title":"Nat. Med."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0070","doi-asserted-by":"crossref","first-page":"E510","DOI":"10.1210\/jc.2011-1754","article-title":"Metformin inhibits growth of thyroid carcinoma cells, suppresses self-renewal of derived cancer stem cells, and potentiates the effect of chemotherapeutic agents","volume":"97","author":"Chen","year":"2012","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0075","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1007\/s40199-018-0208-2","article-title":"Effects of metformin on the PI3K\/AKT\/FOXO1 pathway in anaplastic thyroid cancer cell lines","volume":"26","author":"Nozhat","year":"2018","journal-title":"Daru"},{"key":"10.1016\/j.jsbmb.2020.105684_bib0080","doi-asserted-by":"crossref","first-page":"4030","DOI":"10.1158\/1078-0432.CCR-17-3167","article-title":"Metformin targets mitochondrial glycerophosphate dehydrogenase to control rate of oxidative phosphorylation and growth of thyroid cancer in vitro and in vivo","volume":"24","author":"Thakur","year":"2018","journal-title":"Clin. Cancer Res."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0085","doi-asserted-by":"crossref","first-page":"428","DOI":"10.1080\/09513590.2017.1409714","article-title":"Metformin is associated with reduced cell proliferation in human endometrial cancer by inbibiting PI3K\/AKT\/mTOR signaling","volume":"34","author":"Zhao","year":"2018","journal-title":"Gynecol. Endocrinol."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0090","doi-asserted-by":"crossref","first-page":"8145","DOI":"10.1038\/srep08145","article-title":"Antihyperglycemic mechanism of metformin occurs via the AMPK\/LXR\u03b1\/POMC pathway","volume":"5","author":"Cho","year":"2015","journal-title":"Sci. Rep."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0095","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1210\/jc.2018-01455","article-title":"Type 2 diabetes in neuroendocrine tumors: are biguanides and statins part of the solution?","volume":"104","author":"Herrera-Martinez","year":"2019","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0100","doi-asserted-by":"crossref","first-page":"498","DOI":"10.14310\/horm.2002.1517","article-title":"Anticancer effects of metformin on neuroendocrine tumor cells in vitro","volume":"13","author":"Vlotides","year":"2014","journal-title":"Hormones (Athens)"},{"key":"10.1016\/j.jsbmb.2020.105684_bib0105","doi-asserted-by":"crossref","first-page":"3501","DOI":"10.1210\/jc.2019-00056","article-title":"Biguanides exert antitumoral actions in pituitary tumor cells through AMPK-dependent and -Independent mechanisms","volume":"104","author":"Vazquez-Borrego","year":"2019","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0110","doi-asserted-by":"crossref","first-page":"37538","DOI":"10.18632\/oncotarget.16556","article-title":"Metformin inhibits proliferation and growth hormone secretion of GH3 pituitary adenoma cells","volume":"8","author":"An","year":"2017","journal-title":"Oncotarget"},{"key":"10.1016\/j.jsbmb.2020.105684_bib0115","doi-asserted-by":"crossref","first-page":"985","DOI":"10.1016\/S0015-0282(99)00407-0","article-title":"Effects of metformin on adrenal steroidogenesis in women with polycystic ovary syndrome","volume":"72","author":"la Marca","year":"1999","journal-title":"Fertil. Steril."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0120","doi-asserted-by":"crossref","first-page":"1555","DOI":"10.1210\/jcem.87.4.8398","article-title":"Metformin therapy in obese adolescents with polycystic ovary syndrome and impaired glucose tolerance: amelioration of exaggerated adrenal response to adrenocorticotropin with reduction of insulinemia\/insulin resistance","volume":"87","author":"Arslanian","year":"2002","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0125","doi-asserted-by":"crossref","first-page":"889","DOI":"10.4158\/EP.14.7.889","article-title":"Metformin-responsive classic salt-losing congenital adrenal hyperplasia due to 21-hydroxylase deficiency: a case report","volume":"14","author":"Mapas-Dimaya","year":"2008","journal-title":"Endocr. Pract."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0130","article-title":"Treatment of congenital adrenal Hyperplasia by ReducingInsulin resistance and cysticercosis induced polycystic ovarian syndrome","author":"Sacerdote","year":"2015","journal-title":"Contemp. Gynecol. Pract."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0135","doi-asserted-by":"crossref","first-page":"4354","DOI":"10.1210\/en.2012-1145","article-title":"Metformin inhibits human androgen production by regulating steroidogenic enzymes HSD3B2 and CYP17A1 and complex I activity of the respiratory chain","volume":"153","author":"Hirsch","year":"2012","journal-title":"Endocrinology"},{"key":"10.1016\/j.jsbmb.2020.105684_bib0140","doi-asserted-by":"crossref","first-page":"619","DOI":"10.1530\/eje.0.1440619","article-title":"The effects of long-term metformin treatment on adrenal and ovarian steroidogenesis in women with polycystic ovary syndrome","volume":"144","author":"Vrbikova","year":"2001","journal-title":"Eur. J. Endocrinol."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0145","doi-asserted-by":"crossref","first-page":"8652","DOI":"10.1038\/s41598-017-09189-y","article-title":"Combined transcriptome and metabolome analyses of metformin effects reveal novel links between metabolic networks in steroidogenic systems","volume":"7","author":"Udhane","year":"2017","journal-title":"Sci. Rep."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0150","doi-asserted-by":"crossref","first-page":"164","DOI":"10.1002\/jcp.1041630119","article-title":"Adrenocorticotropin-resistant mutants of the Y1 adrenal cell line fail to express the adrenocorticotropin receptor","volume":"163","author":"Schimmer","year":"1995","journal-title":"J. Cell. Physiol."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0155","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1172\/JCI9238","article-title":"Melanocortin-4 receptor mutations are a frequent and heterogeneous cause of morbid obesity","volume":"106","author":"Vaisse","year":"2000","journal-title":"J. Clin. Invest."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0160","doi-asserted-by":"crossref","first-page":"4318","DOI":"10.1210\/jc.2002-020501","article-title":"Clinical, genetic, and functional characterization of adrenocorticotropin receptor mutations using a novel receptor assay","volume":"87","author":"Fluck","year":"2002","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0165","doi-asserted-by":"crossref","first-page":"E65","DOI":"10.1210\/jc.2010-1056","article-title":"Loss of the C terminus of melanocortin receptor 2 (MC2R) results in impaired cell surface expression and ACTH insensitivity","volume":"96","author":"Hirsch","year":"2011","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0170","doi-asserted-by":"crossref","first-page":"491","DOI":"10.3389\/fendo.2019.00491","article-title":"Stability and turnover of the ACTH receptor complex","volume":"10","author":"Clark","year":"2019","journal-title":"Front. Endocrinol. (Lausanne)"},{"key":"10.1016\/j.jsbmb.2020.105684_bib0175","doi-asserted-by":"crossref","first-page":"434","DOI":"10.1007\/PL00000868","article-title":"Melanocortin receptors: their functions and regulation by physiological agonists and antagonists","volume":"58","author":"Abdel-Malek","year":"2001","journal-title":"Cell. Mol. Life Sci."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0180","doi-asserted-by":"crossref","first-page":"2411","DOI":"10.1016\/j.bbadis.2017.08.001","article-title":"Melanocortin receptors","volume":"1863","author":"Tao","year":"2017","journal-title":"Biochim. Biophys. Acta Mol. Basis Dis."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0185","doi-asserted-by":"crossref","first-page":"T119","DOI":"10.1530\/JME-15-0292","article-title":"60 years of POMC: melanocortin receptors: evolution of ligand selectivity for melanocortin peptides","volume":"56","author":"Dores","year":"2016","journal-title":"J. Mol. Endocrinol."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0190","doi-asserted-by":"crossref","first-page":"3518","DOI":"10.1210\/endo.141.9.7791","article-title":"A unique metabolic syndrome causes obesity in the melanocortin-3 receptor-deficient mouse","volume":"141","author":"Butler","year":"2000","journal-title":"Endocrinology"},{"key":"10.1016\/j.jsbmb.2020.105684_bib0195","doi-asserted-by":"crossref","first-page":"34938","DOI":"10.1038\/srep34938","article-title":"Effects of melanocortin 3 and 4 receptor deficiency on energy homeostasis in rats","volume":"6","author":"You","year":"2016","journal-title":"Sci. Rep."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0200","doi-asserted-by":"crossref","DOI":"10.1126\/sciadv.aat0866","article-title":"Regulation of energy rheostasis by the melanocortin-3 receptor","volume":"4","author":"Ghamari-Langroudi","year":"2018","journal-title":"Sci. Adv."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0205","doi-asserted-by":"crossref","DOI":"10.3390\/ijms19102850","article-title":"Pleiotropic effects of metformin on cancer","volume":"19","author":"Schulten","year":"2018","journal-title":"Int. J. Mol. Sci."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0210","doi-asserted-by":"crossref","first-page":"867","DOI":"10.1016\/j.tips.2018.07.006","article-title":"Metformin as an anticancer agent","volume":"39","author":"Vancura","year":"2018","journal-title":"Trends Pharmacol. Sci."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0215","doi-asserted-by":"crossref","first-page":"R17","DOI":"10.1530\/JME-19-0083","article-title":"The role of an anti-diabetic drug metformin in the treatment of endocrine tumors","author":"Thakur","year":"2019","journal-title":"J. Mol. Endocrinol."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0220","doi-asserted-by":"crossref","DOI":"10.1177\/2036361317749645","article-title":"Efficacy of oral metformin in a patient with metastatic adrenocortical carcinoma: examination of mechanisms and therapeutic implications","volume":"10","author":"Peixoto","year":"2018","journal-title":"Rare Tumors"},{"key":"10.1016\/j.jsbmb.2020.105684_bib0225","doi-asserted-by":"crossref","first-page":"1956","DOI":"10.1124\/dmd.107.015495","article-title":"Metformin transport by a newly cloned proton-stimulated organic cation transporter (plasma membrane monoamine transporter) expressed in human intestine","volume":"35","author":"Zhou","year":"2007","journal-title":"Drug Metab. Dispos."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0230","doi-asserted-by":"crossref","first-page":"1422","DOI":"10.1172\/JCI30558","article-title":"Effect of genetic variation in the organic cation transporter 1 (OCT1) on metformin action","volume":"117","author":"Shu","year":"2007","journal-title":"J. Clin. Invest."},{"key":"10.1016\/j.jsbmb.2020.105684_bib0235","doi-asserted-by":"crossref","first-page":"1552","DOI":"10.1007\/s00125-008-1053-5","article-title":"Metformin and the intestine","volume":"51","author":"Bailey","year":"2008","journal-title":"Diabetologia"}],"container-title":["The Journal of Steroid Biochemistry and Molecular Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0960076019307265?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0960076019307265?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2025,10,18]],"date-time":"2025-10-18T00:07:44Z","timestamp":1760746064000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0960076019307265"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,6]]},"references-count":47,"alternative-id":["S0960076019307265"],"URL":"https:\/\/doi.org\/10.1016\/j.jsbmb.2020.105684","relation":{},"ISSN":["0960-0760"],"issn-type":[{"value":"0960-0760","type":"print"}],"subject":[],"published":{"date-parts":[[2020,6]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Metformin inhibits the activation of melanocortin receptors 2 and 3 in vitro: A possible mechanism for its anti-androgenic and weight balancing effects in vivo?","name":"articletitle","label":"Article Title"},{"value":"The Journal of Steroid Biochemistry and Molecular Biology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.jsbmb.2020.105684","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2020 Elsevier Ltd. All rights reserved.","name":"copyright","label":"Copyright"}],"article-number":"105684"}}