{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T19:39:54Z","timestamp":1789501194493,"version":"build-2803163510"},"reference-count":49,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2024,10,1]],"date-time":"2024-10-01T00:00:00Z","timestamp":1727740800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2024,10,1]],"date-time":"2024-10-01T00:00:00Z","timestamp":1727740800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2024,6,7]],"date-time":"2024-06-07T00:00:00Z","timestamp":1717718400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100018234","name":"Inselspital Universitatsspital Bern","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100018234","id-type":"DOI","asserted-by":"publisher"}]}],"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":[[2024,10]]},"DOI":"10.1016\/j.jsbmb.2024.106561","type":"journal-article","created":{"date-parts":[[2024,6,12]],"date-time":"2024-06-12T21:40:37Z","timestamp":1718228437000},"page":"106561","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":2,"special_numbering":"C","title":["Mitochondrial dysfunction results in enhanced adrenal androgen production in H295R cells"],"prefix":"10.1016","volume":"243","author":[{"given":"D\u00e9borah","family":"Mathis","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Therina","family":"du Toit","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Emre Murat","family":"Altinkilic","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Darko","family":"Stojkov","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christian","family":"Urz\u00ec","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Clarissa D.","family":"Voegel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vincen","family":"Wu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nicola","family":"Zamboni","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hans-Uwe","family":"Simon","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jean-Marc","family":"Nuoffer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christa E.","family":"Fl\u00fcck","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrea","family":"Felser","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.jsbmb.2024.106561_bib1","doi-asserted-by":"crossref","DOI":"10.1016\/j.beem.2022.101665","article-title":"Basics of androgen synthesis and action","author":"Naamneh Elzenaty","year":"2022","journal-title":"Best. Pr. Res Clin. Endocrinol. Metab."},{"key":"10.1016\/j.jsbmb.2024.106561_bib2","doi-asserted-by":"crossref","first-page":"7247","DOI":"10.1073\/pnas.91.15.7247","article-title":"The mitochondrial environment is required for activity of the cholesterol side-chain cleavage enzyme, cytochrome P450scc","volume":"91","author":"Black","year":"1994","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.jsbmb.2024.106561_bib3","doi-asserted-by":"crossref","first-page":"771","DOI":"10.1016\/j.beem.2012.05.002","article-title":"Role of mitochondria in steroidogenesis","volume":"26","author":"Papadopoulos","year":"2012","journal-title":"Best. Pr. Res Clin. Endocrinol. Metab."},{"key":"10.1016\/j.jsbmb.2024.106561_bib4","doi-asserted-by":"crossref","first-page":"771","DOI":"10.1016\/j.tem.2017.09.002","article-title":"Steroidogenesis: unanswered questions","volume":"28","author":"Miller","year":"2017","journal-title":"Trends Endocrinol. Metab."},{"key":"10.1016\/j.jsbmb.2024.106561_bib5","doi-asserted-by":"crossref","first-page":"124","DOI":"10.1016\/j.mce.2014.12.015","article-title":"Regulation of androgen biosynthesis - a short review and preliminary results from the hyperandrogenic starvation NCI-H295R cell model","volume":"408","author":"Kempna","year":"2015","journal-title":"Mol. Cell Endocrinol."},{"key":"10.1016\/j.jsbmb.2024.106561_bib6","doi-asserted-by":"crossref","DOI":"10.1016\/j.molmet.2020.01.001","article-title":"Metabolic dysfunction in polycystic ovary syndrome: Pathogenic role of androgen excess and potential therapeutic strategies","volume":"35","author":"Sanchez-Garrido","year":"2020","journal-title":"Mol. Metab."},{"key":"10.1016\/j.jsbmb.2024.106561_bib7","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1196\/annals.1365.013","article-title":"Premature adrenarche leads to polycystic ovary syndrome? Long-term consequences","volume":"1092","author":"Kousta","year":"2006","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"10.1016\/j.jsbmb.2024.106561_bib8","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1007\/s11154-008-9102-4","article-title":"Androgen synthesis in adrenarche","volume":"10","author":"Miller","year":"2009","journal-title":"Rev. Endocr. Metab. Disord."},{"key":"10.1016\/j.jsbmb.2024.106561_bib9","doi-asserted-by":"crossref","first-page":"739","DOI":"10.1046\/j.1365-2265.2000.01144.x","article-title":"Developmental changes in steroidogenic enzymes in human postnatal adrenal cortex: immunohistochemical studies","volume":"53","author":"Suzuki","year":"2000","journal-title":"Clin. Endocrinol. (Oxf.)"},{"key":"10.1016\/j.jsbmb.2024.106561_bib10","doi-asserted-by":"crossref","DOI":"10.1016\/j.gene.2022.147126","article-title":"Molecular characterization of variants in mitochondrial DNA encoded genes using next generation sequencing analysis and mitochondrial dysfunction in women with PCOS","volume":"855","author":"Shukla","year":"2023","journal-title":"Gene"},{"key":"10.1016\/j.jsbmb.2024.106561_bib11","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/j.mito.2020.02.006","article-title":"Mitochondrial dysfunction: an emerging link in the pathophysiology of polycystic ovary syndrome","volume":"52","author":"Shukla","year":"2020","journal-title":"Mitochondrion"},{"key":"10.1016\/j.jsbmb.2024.106561_bib12","doi-asserted-by":"crossref","first-page":"1401","DOI":"10.1089\/dna.2019.5172","article-title":"Mitochondrial dysfunction in polycystic ovary syndrome","volume":"39","author":"Zeng","year":"2020","journal-title":"DNA Cell Biol."},{"key":"10.1016\/j.jsbmb.2024.106561_bib13","doi-asserted-by":"crossref","DOI":"10.3390\/ijms22083923","article-title":"Mitochondrial dysfunction and chronic inflammation in polycystic ovary syndrome","volume":"22","author":"Dabravolski","year":"2021","journal-title":"Int. J. Mol. Sci."},{"key":"10.1016\/j.jsbmb.2024.106561_bib14","doi-asserted-by":"crossref","DOI":"10.1210\/endocr\/bqaa158","article-title":"Mitochondrial dysfunction in obesity and reproduction","volume":"162","author":"Das","year":"2021","journal-title":"Endocrinology"},{"key":"10.1016\/j.jsbmb.2024.106561_bib15","doi-asserted-by":"crossref","first-page":"532","DOI":"10.3389\/fphys.2019.00532","article-title":"Mitochondrial (dys)function and insulin resistance: from pathophysiological molecular mechanisms to the impact of diet","volume":"10","author":"Sergi","year":"2019","journal-title":"Front Physiol."},{"key":"10.1016\/j.jsbmb.2024.106561_bib16","doi-asserted-by":"crossref","first-page":"981","DOI":"10.1210\/er.2011-1034","article-title":"Insulin resistance and the polycystic ovary syndrome revisited: an update on mechanisms and implications","volume":"33","author":"Diamanti-Kandarakis","year":"2012","journal-title":"Endocr. Rev."},{"key":"10.1016\/j.jsbmb.2024.106561_bib17","doi-asserted-by":"crossref","first-page":"7461","DOI":"10.1007\/s00216-022-03881-3","article-title":"Parallel targeted and non-targeted quantitative analysis of steroids in human serum and peritoneal fluid by liquid chromatography high-resolution mass spectrometry","volume":"414","author":"Andrieu","year":"2022","journal-title":"Anal. Bioanal. Chem."},{"key":"10.1016\/j.jsbmb.2024.106561_bib18","doi-asserted-by":"crossref","first-page":"7074","DOI":"10.1021\/ac201267k","article-title":"High-throughput, accurate mass metabolome profiling of cellular extracts by flow injection-time-of-flight mass spectrometry","volume":"83","author":"Fuhrer","year":"2011","journal-title":"Anal. Chem."},{"key":"10.1016\/j.jsbmb.2024.106561_bib19","doi-asserted-by":"crossref","DOI":"10.3390\/metabo10030103","volume":"10","author":"Rathod","year":"2020","journal-title":"Metabolites"},{"key":"10.1016\/j.jsbmb.2024.106561_bib20","doi-asserted-by":"crossref","DOI":"10.15252\/msb.202211033","article-title":"A functional analysis of 180 cancer cell lines reveals conserved intrinsic metabolic programs","volume":"18","author":"Cherkaoui","year":"2022","journal-title":"Mol. Syst. Biol."},{"key":"10.1016\/j.jsbmb.2024.106561_bib21","doi-asserted-by":"crossref","DOI":"10.3390\/ijms21144853","article-title":"Metabolomic insight into polycystic ovary syndrome-an overview","volume":"21","author":"Rajska","year":"2020","journal-title":"Int J. Mol. Sci."},{"key":"10.1016\/j.jsbmb.2024.106561_bib22","doi-asserted-by":"crossref","first-page":"e1987","DOI":"10.1210\/clinem\/dgab931","article-title":"Novel lipidomic signature associated with metabolic risk in women with and without polycystic ovary syndrome","volume":"107","author":"Mousa","year":"2022","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"10.1016\/j.jsbmb.2024.106561_bib23","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1007\/978-1-61779-382-0_3","article-title":"High-resolution respirometry: OXPHOS protocols for human cells and permeabilized fibers from small biopsies of human muscle","volume":"810","author":"Pesta","year":"2012","journal-title":"Methods Mol. Biol."},{"key":"10.1016\/j.jsbmb.2024.106561_bib24","doi-asserted-by":"crossref","first-page":"414","DOI":"10.1016\/j.molcel.2014.09.025","article-title":"Glutamine oxidation maintains the TCA cycle and cell survival during impaired mitochondrial pyruvate transport","volume":"56","author":"Yang","year":"2014","journal-title":"Mol. Cell"},{"key":"10.1016\/j.jsbmb.2024.106561_bib25","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1186\/1741-7015-10-153","article-title":"Metabolic profiles characterizing different phenotypes of polycystic ovary syndrome: plasma metabolomics analysis","volume":"10","author":"Zhao","year":"2012","journal-title":"BMC Med"},{"key":"10.1016\/j.jsbmb.2024.106561_bib26","doi-asserted-by":"crossref","first-page":"2937","DOI":"10.1021\/pr3000317","article-title":"Metabonomics reveals plasma metabolic changes and inflammatory marker in polycystic ovary syndrome patients","volume":"11","author":"Sun","year":"2012","journal-title":"J. Proteome Res"},{"key":"10.1016\/j.jsbmb.2024.106561_bib27","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1002\/prot.25204","article-title":"Crystal structure of heart 6-phosphofructo-2-kinase\/fructose-2,6-bisphosphatase (PFKFB2) and the inhibitory influence of citrate on substrate binding","volume":"85","author":"Crochet","year":"2017","journal-title":"Proteins"},{"key":"10.1016\/j.jsbmb.2024.106561_bib28","doi-asserted-by":"crossref","DOI":"10.3390\/ijms22126587","article-title":"Understanding the Central Role of Citrate in the Metabolism of Cancer Cells and Tumors: An Update","volume":"22","author":"Icard","year":"2021","journal-title":"Int J. Mol. Sci."},{"key":"10.1016\/j.jsbmb.2024.106561_bib29","doi-asserted-by":"crossref","first-page":"E518","DOI":"10.1210\/jc.2013-3198","article-title":"Transcriptome profiling reveals differentially expressed transcripts between the human adrenal zona fasciculata and zona reticularis","volume":"99","author":"Rege","year":"2014","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"10.1016\/j.jsbmb.2024.106561_bib30","doi-asserted-by":"crossref","DOI":"10.1016\/j.mce.2021.111296","article-title":"Targeted RNA sequencing of adrenal zones using immunohistochemistry-guided capture of formalin-fixed paraffin-embedded tissue","volume":"530","author":"Baker","year":"2021","journal-title":"Mol. Cell Endocrinol."},{"key":"10.1016\/j.jsbmb.2024.106561_bib31","doi-asserted-by":"crossref","DOI":"10.1016\/j.jbc.2022.101652","article-title":"Mitochondrial respiratory chain dysfunction alters ER sterol sensing and mevalonate pathway activity","volume":"298","author":"Wall","year":"2022","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.jsbmb.2024.106561_bib32","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.30952","article-title":"Transcriptomic and proteomic landscape of mitochondrial dysfunction reveals secondary coenzyme Q deficiency in mammals","volume":"6","author":"Kuhl","year":"2017","journal-title":"Elife"},{"key":"10.1016\/j.jsbmb.2024.106561_bib33","doi-asserted-by":"crossref","first-page":"2749","DOI":"10.1016\/j.biocel.2013.09.011","article-title":"Rotenone induces reductive stress and triacylglycerol deposition in C2C12 cells","volume":"45","author":"He","year":"2013","journal-title":"Int J. Biochem Cell Biol."},{"key":"10.1016\/j.jsbmb.2024.106561_bib34","doi-asserted-by":"crossref","first-page":"1133","DOI":"10.1194\/jlr.M400464-JLR200","article-title":"Mitochondrial dysfunction induces triglyceride accumulation in 3T3-L1 cells: role of fatty acid beta-oxidation and glucose","volume":"46","author":"Vankoningsloo","year":"2005","journal-title":"J. Lipid Res"},{"key":"10.1016\/j.jsbmb.2024.106561_bib35","doi-asserted-by":"crossref","first-page":"856","DOI":"10.1016\/j.cmet.2018.12.023","article-title":"Polyunsaturated fatty acid desaturation is a mechanism for glycolytic NAD(+) recycling","volume":"29","author":"Kim","year":"2019","journal-title":"Cell Metab."},{"key":"10.1016\/j.jsbmb.2024.106561_bib36","doi-asserted-by":"crossref","first-page":"507","DOI":"10.1016\/0960-0760(95)00199-9","article-title":"Organization of 3 beta-hydroxysteroid dehydrogenase\/isomerase and cytochrome P450scc into a catalytically active molecular complex in bovine adrenocortical mitochondria","volume":"55","author":"Cherradi","year":"1995","journal-title":"J. Steroid Biochem. Mol. Biol."},{"key":"10.1016\/j.jsbmb.2024.106561_bib37","article-title":"The adrenal lipid droplet is a new site for steroid hormone metabolism","volume":"18","author":"Yu","year":"2018","journal-title":"Proteomics"},{"key":"10.1016\/j.jsbmb.2024.106561_bib38","doi-asserted-by":"crossref","first-page":"1099","DOI":"10.1016\/j.mcna.2021.06.008","article-title":"Practical Approach to Hyperandrogenism in Women","volume":"105","author":"Sharma","year":"2021","journal-title":"Med Clin. North Am."},{"key":"10.1016\/j.jsbmb.2024.106561_bib39","doi-asserted-by":"crossref","first-page":"444","DOI":"10.1111\/j.0013-9580.2005.38404.x","article-title":"Effects of anticonvulsants on human p450c17 (17alpha-hydroxylase\/17,20 lyase) and 3beta-hydroxysteroid dehydrogenase type 2","volume":"46","author":"Fluck","year":"2005","journal-title":"Epilepsia"},{"key":"10.1016\/j.jsbmb.2024.106561_bib40","doi-asserted-by":"crossref","first-page":"773","DOI":"10.1016\/j.jhep.2010.11.006","article-title":"Drug-induced toxicity on mitochondria and lipid metabolism: mechanistic diversity and deleterious consequences for the liver","volume":"54","author":"Begriche","year":"2011","journal-title":"J. Hepatol."},{"key":"10.1016\/j.jsbmb.2024.106561_bib41","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1210\/endrev\/bnaa023","article-title":"Cellular and molecular mechanisms of metformin action","volume":"42","author":"LaMoia","year":"2021","journal-title":"Endocr. Rev."},{"key":"10.1016\/j.jsbmb.2024.106561_bib42","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.2024.106561_bib43","doi-asserted-by":"crossref","first-page":"1267170","DOI":"10.3389\/fendo.2023.1267170","article-title":"Advances in mitochondria-centered mechanism behind the roles of androgens and androgen receptor in the regulation of glucose and lipid metabolism","volume":"14","author":"Yin","year":"2023","journal-title":"Front Endocrinol. (Lausanne)"},{"key":"10.1016\/j.jsbmb.2024.106561_bib44","doi-asserted-by":"crossref","first-page":"C835","DOI":"10.1152\/ajpcell.00205.2022","article-title":"Role of androgens and androgen receptor in control of mitochondrial function","volume":"323","author":"Ahmad","year":"2022","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"10.1016\/j.jsbmb.2024.106561_bib45","doi-asserted-by":"crossref","first-page":"6621","DOI":"10.1074\/jbc.RA118.006727","article-title":"Mitochondrial localization, import, and mitochondrial function of the androgen receptor","volume":"294","author":"Bajpai","year":"2019","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.jsbmb.2024.106561_bib46","doi-asserted-by":"crossref","DOI":"10.3390\/ijms22010243","article-title":"Androgen reduces mitochondrial respiration in mouse brown adipocytes: a model for disordered energy balance in polycystic ovary syndrome","volume":"22","author":"Lerner","year":"2020","journal-title":"Int. J. Mol. Sci."},{"key":"10.1016\/j.jsbmb.2024.106561_bib47","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1038\/s42255-018-0002-y","article-title":"Mitochondrial pyruvate import is a metabolic vulnerability in androgen receptor-driven prostate cancer","volume":"1","author":"Bader","year":"2019","journal-title":"Nat. Metab."},{"key":"10.1016\/j.jsbmb.2024.106561_bib48","first-page":"5488","article-title":"Establishment and characterization of a human adrenocortical carcinoma cell line that expresses multiple pathways of steroid biosynthesis","volume":"50","author":"Gazdar","year":"1990","journal-title":"Cancer Res."},{"key":"10.1016\/j.jsbmb.2024.106561_bib49","doi-asserted-by":"crossref","DOI":"10.1210\/endocr\/bqac076","article-title":"Cholesterol deprivation drives DHEA biosynthesis in human adrenals","volume":"163","author":"Pignatti","year":"2022","journal-title":"Endocrinology"}],"container-title":["The Journal of Steroid Biochemistry and Molecular Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0960076024001092?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0960076024001092?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2024,12,5]],"date-time":"2024-12-05T04:18:38Z","timestamp":1733372318000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0960076024001092"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,10]]},"references-count":49,"alternative-id":["S0960076024001092"],"URL":"https:\/\/doi.org\/10.1016\/j.jsbmb.2024.106561","relation":{},"ISSN":["0960-0760"],"issn-type":[{"value":"0960-0760","type":"print"}],"subject":[],"published":{"date-parts":[[2024,10]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Mitochondrial dysfunction results in enhanced adrenal androgen production in H295R cells","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.2024.106561","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2024 The Author(s). Published by Elsevier Ltd.","name":"copyright","label":"Copyright"}],"article-number":"106561"}}