{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,13]],"date-time":"2026-09-13T01:00:00Z","timestamp":1789261200120,"version":"build-2803163510"},"reference-count":39,"publisher":"The Endocrine Society","issue":"1","license":[{"start":{"date-parts":[[2023,7,19]],"date-time":"2023-07-19T00:00:00Z","timestamp":1689724800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"funder":[{"name":"Antidoping Denmark"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2023,12,21]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:sec>\n                    <jats:title>Context<\/jats:title>\n                    <jats:p>No information exists on the long-lasting effects of supraphysiological anabolic androgenic steroids (AASs) usage on the myocellular properties of human skeletal muscle in previous AAS users.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Objective<\/jats:title>\n                    <jats:p>We hypothesized that former AAS users would demonstrate smaller myonuclei domains (ie, higher myonuclei density) than matched controls.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>A community-based cross-sectional study in men aged 18-50 years engaged in recreational strength training. Muscle biopsies were obtained from the m. vastus lateralis. Immunofluorescence analyses were performed to quantify myonuclei density and myofiber size.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>Twenty-five males were included: 8 current and 7 previous AAS users and 10 controls. Median (25th-75th percentiles) accumulated duration of AAS use was 174 (101-206) and 140 (24-260) weeks in current and former AAS users, respectively (P = .482). Geometric mean (95% CI) elapsed duration since AAS cessation was 4.0 (1.2; 12.7) years among former AAS users. Type II muscle fibers in former AAS users displayed higher myonuclei density and DNA to cytoplasm ratio than controls, corresponding to smaller myonuclei domains (P = .013). Longer accumulated AAS use (weeks, log2) was associated with smaller myonuclei domains in previous AAS users: beta-coefficient (95% CI) \u221294 (\u2212169; \u221218), P = .024. Type I fibers in current AAS users exhibited a higher amount of satellite cells per myofiber (P = .031) than controls.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusion<\/jats:title>\n                    <jats:p>Muscle fibers in former AAS users demonstrated persistently higher myonuclei density and DNA to cytoplasm ratio 4 years after AAS cessation suggestive of enhanced retraining capacity.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.1210\/clinem\/dgad432","type":"journal-article","created":{"date-parts":[[2023,7,19]],"date-time":"2023-07-19T07:30:26Z","timestamp":1689751826000},"page":"e266-e273","source":"Crossref","is-referenced-by-count":17,"title":["Higher Myonuclei Density in Muscle Fibers Persists Among Former Users of Anabolic Androgenic Steroids"],"prefix":"10.1210","volume":"109","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0979-4647","authenticated-orcid":false,"given":"Jakob Lindberg","family":"Nielsen","sequence":"first","affiliation":[{"name":"Department of Sports Science and Clinical Biomechanics, University of Southern Denmark , Odense DK-5230 , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2898-1771","authenticated-orcid":false,"given":"Jon Jarl\u00f8v","family":"Rasmussen","sequence":"additional","affiliation":[{"name":"Department of Medical Endocrinology, Copenhagen University Hospital, Rigshospitalet , Copenhagen DK-2200 , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mikkel Nicklas","family":"Frandsen","sequence":"additional","affiliation":[{"name":"Department of Medical Endocrinology, Copenhagen University Hospital, Rigshospitalet , Copenhagen DK-2200 , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jeppe","family":"Fredberg","sequence":"additional","affiliation":[{"name":"Department of Medical Endocrinology, Copenhagen University Hospital, Rigshospitalet , Copenhagen DK-2200 , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0067-1413","authenticated-orcid":false,"given":"Niels H\u00f8egh","family":"Brandt-Jacobsen","sequence":"additional","affiliation":[{"name":"Department of Medical Endocrinology, Copenhagen University Hospital, Rigshospitalet , Copenhagen DK-2200 , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9773-7361","authenticated-orcid":false,"given":"Per","family":"Aagaard","sequence":"additional","affiliation":[{"name":"Department of Sports Science and Clinical Biomechanics, University of Southern Denmark , Odense DK-5230 , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3019-6775","authenticated-orcid":false,"given":"Caroline","family":"Kistorp","sequence":"additional","affiliation":[{"name":"Department of Medical Endocrinology, Copenhagen University Hospital, Rigshospitalet , Copenhagen DK-2200 , Denmark"},{"name":"Department of Clinical Medicine, Faculty of Health and Medical Sciences, University of Copenhagen , Copenhagen DK-2200 , Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"80","published-online":{"date-parts":[[2023,7,19]]},"reference":[{"issue":"4","key":"2023122122172851300_dgad432-B1","doi-asserted-by":"crossref","first-page":"371","DOI":"10.1111\/j.1521-0391.2013.12118.x","article-title":"The lifetime prevalence of anabolic-androgenic steroid use and dependence in Americans: current best estimates","volume":"23","author":"Pope","year":"2014","journal-title":"Am J Addict"},{"issue":"1","key":"2023122122172851300_dgad432-B2","doi-asserted-by":"crossref","first-page":"e000708","DOI":"10.1136\/bmjsem-2019-000708","article-title":"Is competitive body-building pathological? Survey of 984 male strength trainers","volume":"6","author":"Steele","year":"2020","journal-title":"BMJ Open Sport Exerc Med"},{"key":"2023122122172851300_dgad432-B3","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1016\/j.drugalcdep.2015.04.013","article-title":"Anabolic steroids and cardiovascular risk: a national population-based cohort study","volume":"152","author":"Thiblin","year":"2015","journal-title":"Drug Alcohol Depend"},{"issue":"7","key":"2023122122172851300_dgad432-B4","doi-asserted-by":"crossref","first-page":"644","DOI":"10.1136\/bjsm.2005.025668","article-title":"Homocysteine induced cardiovascular events: a consequence of long term anabolic-androgenic steroid (AAS) abuse","volume":"40","author":"Graham","year":"2006","journal-title":"Br J Sports Med"},{"issue":"3","key":"2023122122172851300_dgad432-B5","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1111\/joim.12850","article-title":"Health consequences of androgenic anabolic steroid use","volume":"285","author":"Horwitz","year":"2019","journal-title":"J Intern Med"},{"issue":"8","key":"2023122122172851300_dgad432-B6","doi-asserted-by":"crossref","first-page":"e0161208","DOI":"10.1371\/journal.pone.0161208","article-title":"Former abusers of anabolic androgenic steroids exhibit decreased testosterone levels and hypogonadal symptoms years after cessation: a case-control study","volume":"11","author":"Rasmussen","year":"2016","journal-title":"PLoS One"},{"issue":"7","key":"2023122122172851300_dgad432-B7","doi-asserted-by":"crossref","first-page":"e2664","DOI":"10.1210\/clinem\/dgab129","article-title":"Serum insulin-like factor 3 levels are reduced in former androgen users, suggesting impaired Leydig cell capacity","volume":"106","author":"Rasmussen","year":"2021","journal-title":"J Clin Endocrinol Metab"},{"issue":"2","key":"2023122122172851300_dgad432-B8","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1097\/HJH.0000000000001546","article-title":"Increased blood pressure and aortic stiffness among abusers of anabolic androgenic steroids: potential effect of suppressed natriuretic peptides in plasma?","volume":"36","author":"Rasmussen","year":"2018","journal-title":"J Hypertens"},{"key":"2023122122172851300_dgad432-B9","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1016\/j.ahj.2018.06.010","article-title":"Cardiac systolic dysfunction in past illicit users of anabolic androgenic steroids","volume":"203","author":"Rasmussen","year":"2018","journal-title":"Am Heart J"},{"issue":"2","key":"2023122122172851300_dgad432-B10","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1007\/s00418-005-0029-5","article-title":"Skeletal muscle morphology in power-lifters with and without anabolic steroids","volume":"124","author":"Eriksson","year":"2005","journal-title":"Histochem Cell Biol"},{"issue":"1","key":"2023122122172851300_dgad432-B11","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1007\/s004180050003","article-title":"The expression of androgen receptors in human neck and limb muscles: effects of training and self-administration of androgenic-anabolic steroids","volume":"113","author":"Kadi","year":"2000","journal-title":"Histochem Cell Biol"},{"issue":"11","key":"2023122122172851300_dgad432-B12","doi-asserted-by":"crossref","first-page":"1528","DOI":"10.1097\/00005768-199911000-00006","article-title":"Effects of anabolic steroids on the muscle cells of strength-trained athletes","volume":"31","author":"Kadi","year":"1999","journal-title":"Med Sci Sports Exerc"},{"issue":"9","key":"2023122122172851300_dgad432-B13","doi-asserted-by":"crossref","first-page":"e105330","DOI":"10.1371\/journal.pone.0105330","article-title":"Effects of long term supplementation of anabolic androgen steroids on human skeletal muscle","volume":"9","author":"Yu","year":"2014","journal-title":"PLoS One"},{"issue":"8","key":"2023122122172851300_dgad432-B14","doi-asserted-by":"crossref","first-page":"3024","DOI":"10.1210\/jc.2006-0357","article-title":"Effects of testosterone supplementation on skeletal muscle fiber hypertrophy and satellite cells in community-dwelling older men","volume":"91","author":"Sinha-Hikim","year":"2006","journal-title":"J Clin Endocrinol Metab"},{"issue":"1","key":"2023122122172851300_dgad432-B15","doi-asserted-by":"crossref","first-page":"E197","DOI":"10.1152\/ajpendo.00370.2002","article-title":"Testosterone-induced muscle hypertrophy is associated with an increase in satellite cell number in healthy, young men","volume":"285","author":"Sinha-Hikim","year":"2003","journal-title":"Am J Physiol Endocrinol Metab"},{"issue":"1","key":"2023122122172851300_dgad432-B16","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1056\/NEJM199607043350101","article-title":"The effects of supraphysiologic doses of testosterone on muscle size and strength in normal men","volume":"335","author":"Bhasin","year":"1996","journal-title":"N Engl J Med"},{"issue":"6","key":"2023122122172851300_dgad432-B17","doi-asserted-by":"crossref","first-page":"E1172","DOI":"10.1152\/ajpendo.2001.281.6.E1172","article-title":"Testosterone dose-response relationships in healthy young men","volume":"281","author":"Bhasin","year":"2001","journal-title":"Am J Physiol Endocrinol Metabol"},{"issue":"2","key":"2023122122172851300_dgad432-B18","doi-asserted-by":"crossref","first-page":"R337","DOI":"10.1152\/ajpregu.00469.2004","article-title":"Mechanisms of disuse muscle atrophy: role of oxidative stress","volume":"288","author":"Powers","year":"2005","journal-title":"Am J Physiol Regul Integr Comp Physiol"},{"issue":"2","key":"2023122122172851300_dgad432-B19","doi-asserted-by":"crossref","first-page":"290","DOI":"10.1152\/japplphysiol.00436.2012","article-title":"No change in myonuclear number during muscle unloading and reloading","volume":"113","author":"Bruusgaard","year":"2012","journal-title":"J Appl Physiol (1985)"},{"issue":"4","key":"2023122122172851300_dgad432-B20","doi-asserted-by":"crossref","first-page":"1450","DOI":"10.1172\/JCI34022","article-title":"In vivo time-lapse microscopy reveals no loss of murine myonuclei during weeks of muscle atrophy","volume":"118","author":"Bruusgaard","year":"2008","journal-title":"J Clin Invest"},{"issue":"34","key":"2023122122172851300_dgad432-B21","doi-asserted-by":"crossref","first-page":"15111","DOI":"10.1073\/pnas.0913935107","article-title":"Myonuclei acquired by overload exercise precede hypertrophy and are not lost on detraining","volume":"107","author":"Bruusgaard","year":"2010","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"3","key":"2023122122172851300_dgad432-B22","doi-asserted-by":"crossref","first-page":"e13204","DOI":"10.1111\/apha.13204","article-title":"Cachexia does not induce loss of myonuclei or muscle fibres during xenografted prostate cancer in mice","volume":"225","author":"Winje","year":"2019","journal-title":"Acta Physiol (Oxf)"},{"issue":"24","key":"2023122122172851300_dgad432-B23","doi-asserted-by":"crossref","first-page":"6221","DOI":"10.1113\/jphysiol.2013.264457","article-title":"A cellular memory mechanism aids overload hypertrophy in muscle long after an episodic exposure to anabolic steroids","volume":"591","author":"Egner","year":"2013","journal-title":"J Physiol"},{"issue":"2","key":"2023122122172851300_dgad432-B24","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1242\/jeb.124495","article-title":"Muscle memory and a new cellular model for muscle atrophy and hypertrophy","volume":"219","author":"Gundersen","year":"2016","journal-title":"J Exp Biol"},{"issue":"6","key":"2023122122172851300_dgad432-B25","doi-asserted-by":"crossref","first-page":"dgz324","DOI":"10.1210\/clinem\/dgz324","article-title":"Rate and extent of recovery from reproductive and cardiac dysfunction due to androgen abuse in men","volume":"105","author":"Shankara-Narayana","year":"2020","journal-title":"J Clin Endocrinol Metab"},{"issue":"17","key":"2023122122172851300_dgad432-B26","doi-asserted-by":"crossref","first-page":"4351","DOI":"10.1113\/jphysiol.2012.237008","article-title":"Proliferation of myogenic stem cells in human skeletal muscle in response to low-load resistance training with blood flow restriction","volume":"590","author":"Nielsen","year":"2012","journal-title":"J Physiol"},{"issue":"10-11","key":"2023122122172851300_dgad432-B27","doi-asserted-by":"crossref","first-page":"911","DOI":"10.1177\/0269215508090092","article-title":"Does functional strength training of the leg in subacute stroke improve physical performance? A pilot randomized controlled trial","volume":"22","author":"Bale","year":"2008","journal-title":"Clin Rehabil"},{"issue":"4","key":"2023122122172851300_dgad432-B28","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1007\/s00418-006-0176-3","article-title":"Hypertrophic muscle fibers with fissures in power-lifters; fiber splitting or defect regeneration?","volume":"126","author":"Eriksson","year":"2006","journal-title":"Histochem Cell Biol"},{"issue":"1","key":"2023122122172851300_dgad432-B29","doi-asserted-by":"crossref","first-page":"E154","DOI":"10.1152\/ajpendo.00502.2001","article-title":"Testosterone-induced increase in muscle size in healthy young men is associated with muscle fiber hypertrophy","volume":"283","author":"Sinha-Hikim","year":"2002","journal-title":"Am J Physiol Endocrinol Metab"},{"issue":"10","key":"2023122122172851300_dgad432-B30","doi-asserted-by":"crossref","first-page":"5245","DOI":"10.1210\/jc.2004-0084","article-title":"Androgen receptor in human skeletal muscle and cultured muscle satellite cells: up-regulation by androgen treatment","volume":"89","author":"Sinha-Hikim","year":"2004","journal-title":"J Clin Endocrinol Metab"},{"issue":"5","key":"2023122122172851300_dgad432-B31","doi-asserted-by":"crossref","first-page":"1240","DOI":"10.1152\/japplphysiol.00893.2019","article-title":"Fiber type-specific hypertrophy and increased capillarization in skeletal muscle following testosterone administration in young women","volume":"128","author":"Horwath","year":"2020","journal-title":"J Appl Physiol (1985)"},{"issue":"2","key":"2023122122172851300_dgad432-B32","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1016\/0378-5122(80)90044-4","article-title":"Interrelationships between sex hormone-binding globulin and testosterone, 5 alpha-dihydrotestosterone and oestradiol-17 beta in blood of normal men","volume":"2","author":"Bartsch","year":"1980","journal-title":"Maturitas"},{"issue":"5","key":"2023122122172851300_dgad432-B33","doi-asserted-by":"crossref","first-page":"C649","DOI":"10.1152\/ajpcell.00050.2019","article-title":"Elevated myonuclear density during skeletal muscle hypertrophy in response to training is reversed during detraining","volume":"316","author":"Dungan","year":"2019","journal-title":"Am J Physiol Cell Physiol"},{"issue":"6","key":"2023122122172851300_dgad432-B34","doi-asserted-by":"crossref","first-page":"1705","DOI":"10.1002\/jcsm.12617","article-title":"Muscle memory: myonuclear accretion, maintenance, morphology, and miRNA levels with training and detraining in adult mice","volume":"11","author":"Murach","year":"2020","journal-title":"J Cachexia Sarcopenia Muscle"},{"key":"2023122122172851300_dgad432-B35","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1016\/j.exger.2019.04.002","article-title":"Muscle mass and strength gains following 6 months of resistance type exercise training are only partly preserved within one year with autonomous exercise continuation in older adults","volume":"121","author":"Snijders","year":"2019","journal-title":"Exp Gerontol"},{"issue":"1","key":"2023122122172851300_dgad432-B36","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/j.mce.2011.11.019","article-title":"Testosterone inhibits transforming growth factor-\u03b2 signaling during myogenic differentiation and proliferation of mouse satellite cells: potential role of follistatin in mediating testosterone action","volume":"350","author":"Braga","year":"2012","journal-title":"Mol Cell Endocrinol"},{"key":"2023122122172851300_dgad432-B37","doi-asserted-by":"crossref","first-page":"283","DOI":"10.3389\/fphys.2015.00283","article-title":"Satellite cells in human skeletal muscle plasticity","volume":"6","author":"Snijders","year":"2015","journal-title":"Front Physiol"},{"issue":"6","key":"2023122122172851300_dgad432-B38","doi-asserted-by":"crossref","first-page":"957","DOI":"10.1111\/jgs.12279","article-title":"Mechanical muscle function and lean body mass during supervised strength training and testosterone therapy in aging men with low-normal testosterone levels","volume":"61","author":"Kvorning","year":"2013","journal-title":"J Am Geriatr Soc"},{"issue":"9","key":"2023122122172851300_dgad432-B39","doi-asserted-by":"crossref","first-page":"e3790","DOI":"10.1210\/clinem\/dgac348","article-title":"Reliability of drug history to verify androgen abuse in men","volume":"107","author":"Shankara-Narayana","year":"2022","journal-title":"J Clin Endocrinol Metab"}],"container-title":["The Journal of Clinical Endocrinology &amp; Metabolism"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/jcem\/advance-article-pdf\/doi\/10.1210\/clinem\/dgad432\/51037469\/dgad432.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jcem\/article-pdf\/109\/1\/e266\/54731249\/dgad432.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jcem\/article-pdf\/109\/1\/e266\/54731249\/dgad432.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,12,21]],"date-time":"2023-12-21T17:17:59Z","timestamp":1703179079000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/jcem\/article\/109\/1\/e266\/7226351"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,7,19]]},"references-count":39,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2023,7,19]]},"published-print":{"date-parts":[[2023,12,21]]}},"URL":"https:\/\/doi.org\/10.1210\/clinem\/dgad432","relation":{},"ISSN":["0021-972X","1945-7197"],"issn-type":[{"value":"0021-972X","type":"print"},{"value":"1945-7197","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2024,1]]},"published":{"date-parts":[[2023,7,19]]}}}