{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T04:45:02Z","timestamp":1791434702123,"version":"4.3.3"},"reference-count":129,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,4,29]],"date-time":"2026-04-29T00:00:00Z","timestamp":1777420800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100003725","name":"National Research Foundation of Korea","doi-asserted-by":"publisher","award":["RS-2020-NR049559"],"award-info":[{"award-number":["RS-2020-NR049559"]}],"id":[{"id":"10.13039\/501100003725","id-type":"DOI","asserted-by":"publisher"}]},{"award":["RS-2020-NR049559"],"award-info":[{"award-number":["RS-2020-NR049559"]}],"id":[{"id":"https:\/\/ror.org\/013aysd81","id-type":"ROR","asserted-by":"publisher"}]},{"name":"Basic Science Research Program","award":["RS-2020-NR054734"],"award-info":[{"award-number":["RS-2020-NR054734"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Cells"],"abstract":"<jats:p>Male infertility is an escalating global health issue, frequently associated with metabolic problems like obesity, diabetes, and age. Recent evidence designates AMP-activated protein kinase (AMPK) as a pivotal regulator linking energy balance to male reproductive function. AMPK regulates essential activities such as spermatogenesis, metabolic support of Sertoli cells, and steroidogenesis in Leydig cells, as well as sperm motility, capacitation, and the acrosome reaction. At the molecular level, AMPK coordinates signaling networks that include mTOR, SIRT1, PGC-1\u03b1, and FOXO to modulate mitochondrial function, oxidative stress, and autophagy-related quality control. Dysregulation of AMPK during metabolic and environmental stress results in compromised spermatogenesis, diminished sperm quality, mitochondrial malfunction, and reduced testosterone synthesis. Targeting AMPK signaling constitutes a possible therapeutic approach for enhancing male reproductive health. Pharmacological agents like metformin and AICAR, together with natural bioactive substances, lifestyle modifications, and exercise mimetics, have shown promise in reestablishing metabolic equilibrium and improving reproductive results. Moreover, combinatorial strategies that integrate antioxidants and autophagy modulators may yield synergistic advantages. Nonetheless, obstacles concerning tissue selectivity, optimum dose, and clinical translation persist. Future perspectives highlight precision medicine, biomarker-directed therapies, and the incorporation of metabolic health into fertility treatment. AMPK-targeted treatments collectively provide a novel and mechanistically sound method for addressing male infertility.<\/jats:p>","DOI":"10.3390\/cells15090808","type":"journal-article","created":{"date-parts":[[2026,4,29]],"date-time":"2026-04-29T13:32:22Z","timestamp":1777469542000},"page":"808","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Targeting AMPK Networks for Male Reproductive Health: Mechanisms and Emerging Therapies"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-6649-3694","authenticated-orcid":false,"given":"Md Ataur","family":"Rahman","sequence":"first","affiliation":[{"name":"Department of Oncology, Karmanos Cancer Institute, Wayne State University, Detroit, MI 48201, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2170-1303","authenticated-orcid":false,"given":"Abdel Halim","family":"Harrath","sequence":"additional","affiliation":[{"name":"Zoology Department, College of Science, King Saud University, Riyadh 11451, Saudi Arabia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maroua","family":"Jalouli","sequence":"additional","affiliation":[{"name":"Department of Biology, College of Science, Imam Mohammad Ibn Saud Islamic University (IMSIU), Riyadh 11623, Saudi Arabia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-7569-6480","authenticated-orcid":false,"given":"Jinwon","family":"Choi","sequence":"additional","affiliation":[{"name":"Department of Pathology, College of Korean Medicine, Kyung Hee University, Seoul 02447, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Min","family":"Choi","sequence":"additional","affiliation":[{"name":"Department of Pathology, College of Korean Medicine, Kyung Hee University, Seoul 02447, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-2470-4613","authenticated-orcid":false,"given":"Sohyun","family":"Park","sequence":"additional","affiliation":[{"name":"Department of Pathology, College of Korean Medicine, Kyung Hee University, Seoul 02447, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hyo Jeong","family":"Kim","sequence":"additional","affiliation":[{"name":"Department of Pathology, College of Korean Medicine, Kyung Hee University, Seoul 02447, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-7916-5743","authenticated-orcid":false,"given":"Amama","family":"Rani","sequence":"additional","affiliation":[{"name":"Department of Pathology, College of Korean Medicine, Kyung Hee University, Seoul 02447, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1769-5879","authenticated-orcid":false,"given":"Salima","family":"Akter","sequence":"additional","affiliation":[{"name":"Department of Pathology, College of Korean Medicine, Kyung Hee University, Seoul 02447, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9276-3894","authenticated-orcid":false,"given":"Moon Nyeo","family":"Park","sequence":"additional","affiliation":[{"name":"Department of Pathology, College of Korean Medicine, Kyung Hee University, Seoul 02447, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8678-156X","authenticated-orcid":false,"given":"Bonglee","family":"Kim","sequence":"additional","affiliation":[{"name":"Department of Pathology, College of Korean Medicine, Kyung Hee University, Seoul 02447, Republic of Korea"},{"name":"Korean Medicine-Based Drug Repositioning Cancer Research Center, College of Korean Medicine, Kyung Hee University, Seoul 02447, Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,4,29]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Huang, B., Wang, Z., Kong, Y., Jin, M., and Ma, L. (2023). Global, regional and national burden of male infertility in 204 countries and territories between 1990 and 2019: An analysis of global burden of disease study. BMC Public Health, 23.","DOI":"10.1186\/s12889-023-16793-3"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1038\/s41572-023-00459-w","article-title":"Male infertility","volume":"9","author":"Eisenberg","year":"2023","journal-title":"Nat. Rev. Dis. Primers"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1007\/978-3-030-89340-8_11","article-title":"Oxidative stress in men with obesity, metabolic syndrome and type 2 diabetes mellitus: Mechanisms and management of reproductive dysfunction","volume":"Volume 1","author":"Leisegang","year":"2022","journal-title":"Oxidative Stress and Toxicity in Reproductive Biology and Medicine: A Comprehensive Update on Male Infertility"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"E160","DOI":"10.1152\/ajpendo.00074.2025","article-title":"Metabolic pathways and male fertility: Exploring the role of Sertoli cells in energy homeostasis and spermatogenesis","volume":"329","author":"Chen","year":"2025","journal-title":"Am. J. Physiol.-Endocrinol. Metab."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1539","DOI":"10.1007\/s00109-021-02125-8","article-title":"AMPK: A key regulator of energy stress and calcium-induced autophagy","volume":"99","author":"Saikia","year":"2021","journal-title":"J. Mol. Med."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"910","DOI":"10.1210\/endrev\/bnad012","article-title":"AMPK and the Endocrine Control of Metabolism","volume":"44","author":"Townsend","year":"2023","journal-title":"Endocr. Rev."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1038\/s41580-022-00547-x","article-title":"New insights into activation and function of the AMPK","volume":"24","author":"Steinberg","year":"2023","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Yang, W., Wang, L., Wang, F., and Yuan, S. (2020). Roles of AMP-activated protein kinase (AMPK) in mammalian reproduction. Front. Cell Dev. Biol., 8.","DOI":"10.3389\/fcell.2020.593005"},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Cris\u00f3stomo, L., Alves, M.G., Gorga, A., Sousa, M., Riera, M.F., Galardo, M.N., Meroni, S.B., and Oliveira, P.F. (2018). Molecular mechanisms and signaling pathways involved in the nutritional support of spermatogenesis by Sertoli cells. Sertoli Cells: Methods and Protocols, Springer.","DOI":"10.1007\/978-1-4939-7698-0_11"},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Li, Y., Zhang, G., Wen, F., Xian, M., Guo, S., Zhang, X., Feng, X., Hu, Z., and Hu, J. (2023). Glucose starvation inhibits ferroptosis by activating the LKB1\/AMPK signaling pathway and promotes the high speed linear motility of dairy goat sperm. Animals, 13.","DOI":"10.3390\/ani13091442"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"7365","DOI":"10.18632\/oncotarget.3629","article-title":"Targeting AMPK for cancer prevention and treatment","volume":"6","author":"Li","year":"2015","journal-title":"Oncotarget"},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Rakoubian, A., Khinchin, J., Yarbro, J., Kobayashi, S., and Liang, Q. (2025). Isoform-specific roles of AMP-activated protein kinase in cardiac physiology and pathophysiology. Front. Cardiovasc. Med., 12.","DOI":"10.3389\/fcvm.2025.1638515"},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Santos, D., and Payan-Carreira, R. (2019). Retinoic acid signaling in spermatogenesis and male (in) fertility. Molecular Signaling in Spermatogenesis and Male Infertility, CRC Press.","DOI":"10.1201\/9780429244216-7"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1016\/j.cmet.2008.11.008","article-title":"The glycogen-binding domain on the AMPK \u03b2 subunit allows the kinase to act as a glycogen sensor","volume":"9","author":"McBride","year":"2009","journal-title":"Cell Metab."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"813","DOI":"10.1016\/j.chembiol.2017.05.021","article-title":"Mechanisms of paradoxical activation of AMPK by the kinase inhibitors SU6656 and sorafenib","volume":"24","author":"Ross","year":"2017","journal-title":"Cell Chem. Biol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1016\/j.cmet.2017.10.009","article-title":"AMPK: Sensing glucose as well as cellular energy status","volume":"27","author":"Lin","year":"2018","journal-title":"Cell Metab."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"36391","DOI":"10.31083\/RCM36391","article-title":"The Role of AMPK\u03b1 in the Mechanism of Development and Treatment of Heart Failure","volume":"26","author":"Feng","year":"2025","journal-title":"Rev. Cardiovasc. Med."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"e70054","DOI":"10.1002\/mnfr.70054","article-title":"Obesity, white adipose tissue, and adipokines signaling in male reproduction","volume":"69","author":"Martins","year":"2025","journal-title":"Mol. Nutr. Food Res."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Bertoldo, M.J., Faure, M., Dupont, J., and Froment, P. (2015). AMPK: A master energy regulator for gonadal function. Front. Neurosci., 9.","DOI":"10.3389\/fnins.2015.00235"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1016\/j.coemr.2019.03.001","article-title":"Regulation of Leydig cell steroidogenesis: Intriguing network of signaling pathways and mitochondrial signalosome","volume":"6","author":"Andric","year":"2019","journal-title":"Curr. Opin. Endocr. Metab. Res."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1038\/s41419-019-1782-z","article-title":"Multiple signaling pathways in Sertoli cells: Recent findings in spermatogenesis","volume":"10","author":"Ni","year":"2019","journal-title":"Cell Death Dis."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Kim, S., Baek, J., and Kim, M.S. (2025). The Link Between Dietary Timing and Exercise Performance Through Adipocyte AMPK\u03b12 Signaling. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26136061"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1038\/s41392-023-01608-z","article-title":"Multifaceted role of mTOR (mammalian target of rapamycin) signaling pathway in human health and disease","volume":"8","author":"Panwar","year":"2023","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1007\/s13105-017-0576-y","article-title":"Multi-regulatory network of ROS: The interconnection of ROS, PGC-1 alpha, and AMPK-SIRT1 during exercise","volume":"73","author":"Thirupathi","year":"2017","journal-title":"J. Physiol. Biochem."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Jiang, Y., Yan, F., Feng, Z., Lazarovici, P., and Zheng, W. (2019). Signaling network of forkhead family of transcription factors (FOXO) in dietary restriction. Cells, 9.","DOI":"10.3390\/cells9010100"},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Martin-Hidalgo, D., Hurtado de Llera, A., Calle-Guisado, V., Gonzalez-Fernandez, L., Garcia-Marin, L., and Bragado, M.J. (2018). AMPK function in mammalian spermatozoa. Int. J. Mol. Sci., 19.","DOI":"10.3390\/ijms19113293"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1080\/19396368.2025.2593339","article-title":"A concise overview of mammalian spermatogenesis","volume":"72","author":"Sultana","year":"2026","journal-title":"Syst. Biol. Reprod. Med."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"3198","DOI":"10.1007\/s43032-025-01970-z","article-title":"Regulatory Interplay of p53, AMPK, and mTOR in Decidualization: Implications for Reproductive Competence and Cancer Biology","volume":"32","author":"Kobayashi","year":"2025","journal-title":"Reprod. Sci."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Ma, R., Cui, Y., Yu, S.-J., Pan, Y.-Y., He, J.-f., Wang, Y.-y., Wang, J.-l., Wang, X.-y., Bai, X.-f., and Zhang, H. (2025). The glucose metabolism reprogramming of yak Sertoli cells under hypoxia is regulated by autophagy. BMC Genom., 26.","DOI":"10.1186\/s12864-025-11497-x"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1124\/pr.110.002790","article-title":"The blood-testis barrier and its implications for male contraception","volume":"64","author":"Cheng","year":"2012","journal-title":"Pharmacol. Rev."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"2681","DOI":"10.1007\/s00018-019-03101-9","article-title":"The roles and mechanisms of Leydig cells and myoid cells in regulating spermatogenesis","volume":"76","author":"Zhou","year":"2019","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"489","DOI":"10.1038\/s41419-019-1728-5","article-title":"Adrenomedullin alleviates the pyroptosis of Leydig cells by promoting autophagy via the ROS\u2013AMPK\u2013mTOR axis","volume":"10","author":"Li","year":"2019","journal-title":"Cell Death Dis."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"e1569","DOI":"10.1002\/wsbm.1569","article-title":"Energy metabolism in mammalian sperm motility","volume":"14","author":"Amaral","year":"2022","journal-title":"WIREs Mech. Dis."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"789","DOI":"10.1016\/j.molcel.2017.05.032","article-title":"AMPK: Mechanisms of cellular energy sensing and restoration of metabolic balance","volume":"66","author":"Garcia","year":"2017","journal-title":"Mol. Cell"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"816","DOI":"10.1038\/aja.2012.81","article-title":"Mechanism of sperm capacitation and the acrosome reaction: Role of protein kinases","volume":"14","author":"Ickowicz","year":"2012","journal-title":"Asian J. Androl."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Park, Y.-J., and Pang, M.-G. (2021). Mitochondrial functionality in male fertility: From spermatogenesis to fertilization. Antioxidants, 10.","DOI":"10.3390\/antiox10010098"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"102227","DOI":"10.1016\/j.redox.2021.102227","article-title":"Induction of autophagy via the ROS-dependent AMPK-mTOR pathway protects copper-induced spermatogenesis disorder","volume":"49","author":"Guo","year":"2022","journal-title":"Redox Biol."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Wang, Y., Fu, X., and Li, H. (2025). Mechanisms of oxidative stress-induced sperm dysfunction. Front. Endocrinol., 16.","DOI":"10.3389\/fendo.2025.1520835"},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Mohan, C., and Sikka, S.C. (2018). Role of Environmental Toxicants in Sperm Autophagy, Mitophagy, and Apoptosis. Bioenvironmental Issues Affecting Men\u2019s Reproductive and Sexual Health, Elsevier.","DOI":"10.1016\/B978-0-12-801299-4.00020-7"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"680","DOI":"10.1093\/humrep\/deac021","article-title":"Autophagy is activated in human spermatozoa subjected to oxidative stress and its inhibition impairs sperm quality and promotes cell death","volume":"37","author":"Uribe","year":"2022","journal-title":"Hum. Reprod."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1186\/s43043-022-00099-2","article-title":"Obesity and male infertility: Multifaceted reproductive disruption","volume":"27","author":"Chaudhuri","year":"2022","journal-title":"Middle East Fertil. Soc. J."},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Hasan, H., Bhushan, S., Fijak, M., and Meinhardt, A. (2022). Mechanism of inflammatory associated impairment of sperm function, spermatogenesis and steroidogenesis. Front. Endocrinol., 13.","DOI":"10.3389\/fendo.2022.897029"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Zhou, J., Hu, S., Ouyang, Y., Kong, Y., and Liu, M. (2025). Immunometabolism and male reproductive function: Linking inflammation, oxidative stress, and declining fertility. Front. Immunol., 16.","DOI":"10.3389\/fimmu.2025.1736492"},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Chianese, R., and Pierantoni, R. (2021). Mitochondrial reactive oxygen species (ROS) production alters sperm quality. Antioxidants, 10.","DOI":"10.3390\/antiox10010092"},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Dutta, S., Sengupta, P., Slama, P., and Roychoudhury, S. (2021). Oxidative stress, testicular inflammatory pathways, and male reproduction. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms221810043"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"e13178","DOI":"10.1111\/aji.13178","article-title":"Obesity and metabolic syndrome associated with systemic inflammation and the impact on the male reproductive system","volume":"82","author":"Leisegang","year":"2019","journal-title":"Am. J. Reprod. Immunol."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Tesarik, J. (2025). Lifestyle and environmental factors affecting male fertility, individual predisposition, prevention, and intervention. Int. J. Mol. Sci., 26.","DOI":"10.20944\/preprints202503.1302.v1"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"5563759","DOI":"10.1155\/2022\/5563759","article-title":"A common feature of pesticides: Oxidative stress\u2014The role of oxidative stress in pesticide-induced toxicity","volume":"2022","author":"Sule","year":"2022","journal-title":"Oxidative Med. Cell. Longev."},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Lyons, C.L., and Roche, H.M. (2018). Nutritional modulation of AMPK-impact upon metabolic-inflammation. Int. J. Mol. Sci., 19.","DOI":"10.3390\/ijms19103092"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"1207","DOI":"10.1093\/humrep\/deac067","article-title":"At the crossroads of fertility and metabolism: The importance of AMPK-dependent signaling in female infertility associated with hyperandrogenism","volume":"37","author":"Froment","year":"2022","journal-title":"Hum. Reprod."},{"key":"ref_51","unstructured":"Teklu, A.A. (2025). Deciphering Environmental Influences in Human Aging and Health. [Ph.D. Thesis, University of Copenhagen]."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"163","DOI":"10.3390\/jox11040012","article-title":"The impact of Endocrine-Disrupting chemicals in male fertility: Focus on the action of obesogens","volume":"11","author":"Rato","year":"2021","journal-title":"J. Xenobiotics"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1093\/toxsci\/kfy008","article-title":"Mitochondrial toxicity","volume":"162","author":"Meyer","year":"2018","journal-title":"Toxicol. Sci."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"902","DOI":"10.2174\/0113816128286655240304070740","article-title":"Revisiting the mitochondrial function and communication in neurodegenerative diseases","volume":"30","author":"Wankhede","year":"2024","journal-title":"Curr. Pharm. Des."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"667","DOI":"10.1007\/s00109-011-0748-0","article-title":"AMP-activated protein kinase inhibits NF-\u03baB signaling and inflammation: Impact on healthspan and lifespan","volume":"89","author":"Salminen","year":"2011","journal-title":"J. Mol. Med."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"132819","DOI":"10.1016\/j.chemosphere.2021.132819","article-title":"AMPK pathway is implicated in low level lead-induced pubertal testicular damage via disordered glycolysis","volume":"291","author":"Xie","year":"2022","journal-title":"Chemosphere"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"545","DOI":"10.1016\/j.tem.2017.05.004","article-title":"AMPK as a therapeutic target for treating metabolic diseases","volume":"28","author":"Day","year":"2017","journal-title":"Trends Endocrinol. Metab."},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Kim, H.I., Han, Y., and Park, J. (2025). AMP-Activated Protein Kinases in Health and Disease. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26168075"},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Agius, L., Ford, B.E., and Chachra, S.S. (2020). The metformin mechanism on gluconeogenesis and AMPK activation: The metabolite perspective. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21093240"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"17","DOI":"10.3390\/antiox10010017","article-title":"Chitosan-stabilized selenium nanoparticles and metformin synergistically rescue testicular oxidative damage and steroidogenesis-related genes dysregulation in high-fat diet\/streptozotocin-induced diabetic rats","volume":"10","author":"Khater","year":"2020","journal-title":"Antioxidants"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"3387","DOI":"10.21037\/tau-2025-440","article-title":"Antidiabetic agents and male fertility: Unraveling the impact of glucose-lowering therapies on sperm quality in diabetic males\u2014A narrative review","volume":"14","author":"Luo","year":"2025","journal-title":"Transl. Androl. Urol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"bio033852","DOI":"10.1242\/bio.033852","article-title":"The effects of AICAR and rapamycin on mitochondrial function in immortalized mitochondrial DNA mutator murine embryonic fibroblasts","volume":"7","author":"Delic","year":"2018","journal-title":"Biol. Open"},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Samsonov, M.V., Podkuychenko, N.V., Khapchaev, A.Y., Efremov, E.E., Yanushevskaya, E.V., Vlasik, T.N., Lankin, V.Z., Stafeev, I.S., Skulachev, M.V., and Shestakova, M.V. (2021). AICAR protects vascular endothelial cells from oxidative injury induced by the long-term palmitate excess. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23010211"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"562","DOI":"10.1177\/0394632016673369","article-title":"A concise review on advances in development of small molecule anti-inflammatory therapeutics emphasising AMPK: An emerging target","volume":"29","year":"2016","journal-title":"Int. J. Immunopathol. Pharmacol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"280","DOI":"10.1210\/endrev\/bnaf041","article-title":"Mechanisms of Testosterone\u2019s Anabolic Effects on Muscle and Function: Controversies and New Insights","volume":"47","author":"Bhasin","year":"2025","journal-title":"Endocr. Rev."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Simon, L., and Mariotti-Celis, M.S. (2024). Bioactive compounds as potential alternative treatments to prevent cancer therapy-induced male infertility. Front. Endocrinol., 14.","DOI":"10.3389\/fendo.2023.1293780"},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Anchimowicz, J., Zielonka, P., and Jakiela, S. (2025). Plant secondary metabolites as modulators of mitochondrial health: An overview of their anti-oxidant, anti-apoptotic, and mitophagic mechanisms. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26010380"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"20220210","DOI":"10.1098\/rstb.2022.0210","article-title":"Reactive oxygen species: Role in obesity and mitochondrial energy efficiency","volume":"378","author":"Corkey","year":"2023","journal-title":"Philos. Trans. R. Soc. B Biol. Sci."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1016\/j.rbmo.2015.06.010","article-title":"Spermatogenesis, DNA damage and DNA repair mechanisms in male infertility","volume":"31","author":"Gunes","year":"2015","journal-title":"Reprod. Biomed. Online"},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Capozzi, A., Saucier, C., Bisbal, C., and Lambert, K. (2022). Grape polyphenols in the treatment of human skeletal muscle damage due to inflammation and oxidative stress during obesity and aging: Early outcomes and promises. Molecules, 27.","DOI":"10.3390\/molecules27196594"},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Xue, C., Yan, Z., Cheng, W., Zhang, D., Zhang, R., Duan, H., Zhang, L., Ma, X., Hu, J., and Kang, J. (2025). Curcumin ameliorates aging-induced blood-testis barrier disruption by regulating AMPK\/mTOR mediated autophagy. PLoS ONE, 20.","DOI":"10.1371\/journal.pone.0321752"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"e23768","DOI":"10.1002\/mrd.23768","article-title":"Effect of berberine combined with metformin on autophagy in polycystic ovary syndrome by regulating AMPK\/AKT\/mTOR pathway","volume":"91","author":"Jin","year":"2024","journal-title":"Mol. Reprod. Dev."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"10333","DOI":"10.3390\/ijms262110333","article-title":"Impact of Bariatric Surgery on the Expression of Fertility-Related Genes in Obese Women: A Systematic Review of LEP, LEPR, MC4R, FTO, and POMC","volume":"26","author":"Voros","year":"2025","journal-title":"Int. J. Mol. Sci."},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Mishra, M.K., Sharma, S., Sharma, S., and Pamu, S. (2025). Mechanisms and modalities of herbal anti-fertility: Toward integrative reproductive health solutions. Adv. Tradit. Med., 1\u201325.","DOI":"10.1007\/s13596-025-00870-6"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"862","DOI":"10.1038\/s41573-021-00217-1","article-title":"Exercise mimetics: Harnessing the therapeutic effects of physical activity","volume":"20","author":"Gubert","year":"2021","journal-title":"Nat. Rev. Drug Discov."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Zineldeen, D.H., Tahoon, N.M., and Sarhan, N.I. (2023). AICAR ameliorates non-alcoholic fatty liver disease via modulation of the HGF\/NF-\u03baB\/SNARK signaling pathway and restores mitochondrial and endoplasmic reticular impairments in high-fat diet-fed rats. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24043367"},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Xu, X., Luo, D., Xuan, Q., Lu, P., Yu, C., and Guan, Q. (2022). Atlas of metabolism reveals palmitic acid results in mitochondrial dysfunction and cell apoptosis by inhibiting fatty acid \u03b2-oxidation in Sertoli cells. Front. Endocrinol., 13.","DOI":"10.3389\/fendo.2022.1021263"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"814","DOI":"10.1016\/j.fertnstert.2008.10.068","article-title":"Sperm viability, apoptosis, and intracellular reactive oxygen species levels in human spermatozoa before and after induction of oxidative stress","volume":"93","author":"Mahfouz","year":"2010","journal-title":"Fertil. Steril."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"242","DOI":"10.1016\/j.cmet.2016.10.022","article-title":"Exercise mimetics: Impact on health and performance","volume":"25","author":"Fan","year":"2017","journal-title":"Cell Metab."},{"key":"ref_80","doi-asserted-by":"crossref","unstructured":"Saleh, R., Sallam, H., Elsuity, M.A., Dutta, S., Sengupta, P., and Nasr, A. (2025). Antioxidant therapy for infertile couples: A comprehensive review of the current status and consideration of future prospects. Front. Endocrinol., 15.","DOI":"10.3389\/fendo.2024.1503905"},{"key":"ref_81","doi-asserted-by":"crossref","unstructured":"Nowicka-Bauer, K., and Nixon, B. (2020). Molecular changes induced by oxidative stress that impair human sperm motility. Antioxidants, 9.","DOI":"10.3390\/antiox9020134"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"1387","DOI":"10.1007\/s43032-021-00795-w","article-title":"The role of the natural antioxidant mechanism in sperm cells","volume":"29","author":"Kowalczyk","year":"2022","journal-title":"Reprod. Sci."},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Wang, S., Li, H., Yuan, M., Fan, H., and Cai, Z. (2022). Role of AMPK in autophagy. Front. Physiol., 13.","DOI":"10.3389\/fphys.2022.1015500"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"648","DOI":"10.1038\/s41419-023-06154-8","article-title":"Autophagy: Regulator of cell death","volume":"14","author":"Liu","year":"2023","journal-title":"Cell Death Dis."},{"key":"ref_85","doi-asserted-by":"crossref","unstructured":"Lei, X., and Rezaei, M.J. (2025). Synergistic effects of polyphenols and exercise on obesity: Targeting metabolism, muscle function, and adipose tissue remodeling. Front. Nutr., 12.","DOI":"10.3389\/fnut.2025.1679381"},{"key":"ref_86","doi-asserted-by":"crossref","unstructured":"Zhang, K., Ma, Y., Luo, Y., Song, Y., Xiong, G., Ma, Y., Sun, X., and Kan, C. (2023). Metabolic diseases and healthy aging: Identifying environmental and behavioral risk factors and promoting public health. Front. Public Health, 11.","DOI":"10.3389\/fpubh.2023.1253506"},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Nguyen, T.M.D. (2017). Impact of 5\u2032-amp-activated protein kinase on male gonad and spermatozoa functions. Front. Cell Dev. Biol., 5.","DOI":"10.3389\/fcell.2017.00025"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1080\/17425247.2016.1213238","article-title":"Surface functionalization of polymeric nanoparticles for tumor drug delivery: Approaches and challenges","volume":"14","author":"Abouelmagd","year":"2017","journal-title":"Expert Opin. Drug Deliv."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1002\/mrd.1120170106","article-title":"Intratesticular injection as a method to assess the potential toxicity of various agents and to study mechanisms of normal spermatogenesis","volume":"17","author":"Russell","year":"1987","journal-title":"Gamete Res."},{"key":"ref_90","doi-asserted-by":"crossref","unstructured":"Rodriguez-Cortes, B., Gomez-Mart\u00ednez, R., Spenle, R., Schwaninger, M., Nogueiras, R., Rasika, S., and Prevot, V. (Trends Endocrinol. Metab., 2025). Tanycytes: Bloodhounds of the metabolic brain, Trends Endocrinol. Metab., Epub ahead of printing.","DOI":"10.1016\/j.tem.2025.10.005"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"351","DOI":"10.1007\/s13346-011-0039-x","article-title":"Drug delivery to the testis: Current status and potential pathways for the development of novel therapeutics","volume":"1","author":"Samplaski","year":"2011","journal-title":"Drug Deliv. Transl. Res."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"e13617","DOI":"10.1111\/and.13617","article-title":"Obesity and male infertility: Mechanisms and management","volume":"53","author":"Leisegang","year":"2021","journal-title":"Andrologia"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"e224","DOI":"10.1038\/emm.2016.16","article-title":"AMPK activators: Mechanisms of action and physiological activities","volume":"48","author":"Kim","year":"2016","journal-title":"Exp. Mol. Med."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"350","DOI":"10.4103\/1008-682X.178857","article-title":"Recent advances in the genetics of testicular failure","volume":"18","author":"Song","year":"2016","journal-title":"Asian J. Androl."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1038\/nrm.2017.95","article-title":"AMPK: Guardian of metabolism and mitochondrial homeostasis","volume":"19","author":"Herzig","year":"2018","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1186\/s12958-022-01034-w","article-title":"How much obesity and diabetes do impair male fertility?","volume":"21","author":"AbbasiHormozi","year":"2023","journal-title":"Reprod. Biol. Endocrinol."},{"key":"ref_97","doi-asserted-by":"crossref","unstructured":"Tseng, C.-H. (2022). Metformin\u2019s effects on varicocele, erectile dysfunction, infertility and prostate-related diseases: A retrospective cohort study. Front. Pharmacol., 13.","DOI":"10.3389\/fphar.2022.799290"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"1149","DOI":"10.1016\/S0015-0282(00)00501-X","article-title":"Metformin therapy decreases hyperandrogenism and hyperinsulinemia in women with polycystic ovary syndrome","volume":"73","author":"Kolodziejczyk","year":"2000","journal-title":"Fertil. Steril."},{"key":"ref_99","unstructured":"Stoica, R.A., \u0218tefan, D.S., Rizzo, M., Suceveanu, A.I., Suceveanu, A.P., Serafinceanu, C., and Pantea-Stoian, A. (2019). Metformin indications, dosage, adverse reactions, and contraindications. Metformin, IntechOpen."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"560","DOI":"10.4103\/aja.aja_141_19","article-title":"Metformin treatment of high-fat diet-fed obese male mice restores sperm function and fetal growth, without requiring weight loss","volume":"22","author":"McPherson","year":"2020","journal-title":"Asian J. Androl."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"11","DOI":"10.5534\/wjmh.210001","article-title":"The effect of metformin on male reproductive function and prostate: An updated review","volume":"40","author":"Tseng","year":"2021","journal-title":"World J. Men\u2019s Health"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"449","DOI":"10.1210\/clinem\/dgad523","article-title":"Improvements in sperm motility following low-or high-intensity dietary interventions in men with obesity","volume":"109","author":"Sharma","year":"2024","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"4475","DOI":"10.2174\/1381612823666170823103830","article-title":"Diabetes mellitus and male aging: Pharmacotherapeutics and clinical implications","volume":"23","author":"Putta","year":"2017","journal-title":"Curr. Pharm. Des."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"e14524","DOI":"10.1111\/and.14524","article-title":"Relationship between obesity-related markers, biochemical metabolic parameters, hormonal profiles and sperm parameters among men attending an infertility clinic","volume":"54","author":"Esmaeili","year":"2022","journal-title":"Andrologia"},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1093\/biolre\/ioaf266","article-title":"Metabolic insights into male fertility: A narrative review of the role of adiponectin on the hypothalamic\u2013pituitary\u2013testicular axis","volume":"114","author":"Gabriel","year":"2025","journal-title":"Biol. Reprod."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"117642","DOI":"10.1016\/j.bcp.2025.117642","article-title":"Therapeutic potential of testosterone in hypogonadal males with MAFLD via AMPK pathways: Evidence from in vitro and in vivo high-fat diet models","volume":"245","author":"Hsieh","year":"2026","journal-title":"Biochem. Pharmacol."},{"key":"ref_107","doi-asserted-by":"crossref","unstructured":"Jin, Y.-H., Gao, D.-W., Li, C.-Y., Sun, D.-L., Mo, H., and Jin, B.-F. (2026). A narrative review on the role of autophagy in male infertility-associated sperm abnormalities. Asian J. Androl.","DOI":"10.4103\/aja202589"},{"key":"ref_108","doi-asserted-by":"crossref","unstructured":"Goel, S., Singh, R., Singh, V., Singh, H., Kumari, P., Chopra, H., Sharma, R., Nepovimova, E., Valis, M., and Kuca, K. (2022). Metformin: Activation of 5\u2032AMP-activated protein kinase and its emerging potential beyond anti-hyperglycemic action. Front. Genet., 13.","DOI":"10.3389\/fgene.2022.1022739"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"1702","DOI":"10.1194\/jlr.M015354","article-title":"Chronic AMP-kinase activation with AICAR reduces adiposity by remodeling adipocyte metabolism and increasing leptin sensitivity","volume":"52","author":"Gaidhu","year":"2011","journal-title":"J. Lipid Res."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"18293","DOI":"10.18632\/oncotarget.4715","article-title":"Exercise-mimetic AICAR transiently benefits brain function","volume":"6","author":"Guerrieri","year":"2015","journal-title":"Oncotarget"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"707","DOI":"10.4103\/1008-682X.185848","article-title":"AMP-activated kinase in human spermatozoa: Identification, intracellular localization, and key function in the regulation of sperm motility","volume":"19","author":"Mijares","year":"2017","journal-title":"Asian J. Androl."},{"key":"ref_112","doi-asserted-by":"crossref","unstructured":"Liu, C.Y., Chang, T.C., Lin, S.H., Wu, S.T., Cha, T.L., and Tsao, C.W. (2020). Metformin Ameliorates Testicular Function and Spermatogenesis in Male Mice with High-Fat and High-Cholesterol Diet-Induced Obesity. Nutrients, 12.","DOI":"10.3390\/nu12071932"},{"key":"ref_113","doi-asserted-by":"crossref","unstructured":"Kamar Bashah, N.A., Hamid, A.A., Adam, S.H., Jaffar, F.H.F., Abd Rahman, I.Z., and Mokhtar, M.H. (2025). Role of AMP-Activated Protein Kinase (AMPK) in Female Reproduction: A Review. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26146833"},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"4257","DOI":"10.1128\/MCB.00734-14","article-title":"A cell-autonomous molecular cascade initiated by AMP-activated protein kinase represses steroidogenesis","volume":"34","author":"Abdou","year":"2014","journal-title":"Mol. Cell. Biol."},{"key":"ref_115","doi-asserted-by":"crossref","unstructured":"M\u0103nescu, D.C., Pl\u0103stoi, C.D., Petre, R.L., M\u0103rg\u0103rit, I.R., M\u0103nescu, A.M., and P\u00eervan, A. (2026). Metabolic Overdrive in Elite Sport: A Systems Model of AMPK-mTOR Oscillation, NAD(+) Economy, and Epigenetic Drift. Int. J. Mol. Sci., 27.","DOI":"10.3390\/ijms27041817"},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"1211","DOI":"10.1016\/j.advnut.2023.07.006","article-title":"The Beneficial and Adverse Effects of Autophagic Response to Caloric Restriction and Fasting","volume":"14","author":"Shabkhizan","year":"2023","journal-title":"Adv. Nutr."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"5054","DOI":"10.2174\/1381612826666200523172334","article-title":"AMPK and its Activator Berberine in the Treatment of Neurodegenerative Diseases","volume":"26","author":"Qin","year":"2020","journal-title":"Curr. Pharm. Des."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"276","DOI":"10.3109\/10409238.2010.488215","article-title":"AMPK inhibition in health and disease","volume":"45","author":"Viollet","year":"2010","journal-title":"Crit. Rev. Biochem. Mol. Biol."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"198","DOI":"10.1007\/s13311-015-0399-x","article-title":"Chronic Treatment with the AMPK Agonist AICAR Prevents Skeletal Muscle Pathology but Fails to Improve Clinical Outcome in a Mouse Model of Severe Spinal Muscular Atrophy","volume":"13","author":"Montull","year":"2016","journal-title":"Neurotherapeutics"},{"key":"ref_120","doi-asserted-by":"crossref","unstructured":"do Val Lima, P.R., Ronconi, K.S., Morra, E.A., Rodrigues, P.L., \u00c1vila, R.A., Merlo, E., Graceli, J.B., Sim\u00f5es, M.R., Stefanon, I., and Ribeiro J\u00fanior, R.F. (2023). Testosterone deficiency impairs cardiac interfibrillar mitochondrial function and myocardial contractility while inducing oxidative stress. Front. Endocrinol., 14.","DOI":"10.3389\/fendo.2023.1206387"},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1186\/s12958-018-0406-2","article-title":"Reactive oxygen species and male reproductive hormones","volume":"16","author":"Darbandi","year":"2018","journal-title":"Reprod. Biol. Endocrinol."},{"key":"ref_122","doi-asserted-by":"crossref","unstructured":"Hurtado de Llera, A., Martin-Hidalgo, D., Gil, M.C., Garcia-Marin, L.J., and Bragado, M.J. (2012). AMP-activated kinase AMPK is expressed in boar spermatozoa and regulates motility. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0038840"},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"232","DOI":"10.1016\/j.bbcan.2016.09.006","article-title":"Functional characterization of AMP-activated protein kinase signaling in tumorigenesis","volume":"1866","author":"Cheng","year":"2016","journal-title":"Biochim. et Biophys. Acta (BBA)-Rev. Cancer"},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"3361","DOI":"10.1016\/j.febslet.2014.07.025","article-title":"Oxidative stress activates AMPK in cultured cells primarily by increasing cellular AMP and\/or ADP","volume":"588","author":"Auciello","year":"2014","journal-title":"FEBS Lett."},{"key":"ref_125","doi-asserted-by":"crossref","unstructured":"Fox, R., Akinboro, S., K\u0119dzia, A., and Niechcia\u0142, E. (2025). The Effects of Maternal Endocrinopathies and Exposure to Endocrine Disruptors During Pregnancy on the Fetus and Newborn. Biomedicines, 13.","DOI":"10.3390\/biomedicines13081965"},{"key":"ref_126","doi-asserted-by":"crossref","unstructured":"Hirsch, A., Hahn, D., Kempn\u00e1, P., Hofer, G., Mullis, P.E., Nuoffer, J.M., and Fl\u00fcck, C.E. (2012). Role of AMP-activated protein kinase on steroid hormone biosynthesis in adrenal NCI-H295R cells. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0030956"},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"2837","DOI":"10.1128\/MCB.05853-11","article-title":"The AMPK \u03b22 Subunit Is Required for Energy Homeostasis during Metabolic Stress","volume":"32","author":"Dasgupta","year":"2012","journal-title":"Mol. Cell. Biol."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"2143","DOI":"10.1016\/j.bbamem.2013.05.026","article-title":"AMP-activated kinase, AMPK, is involved in the maintenance of plasma membrane organization in boar spermatozoa","volume":"1828","author":"Gil","year":"2013","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"1057","DOI":"10.1007\/s11154-021-09659-9","article-title":"Metabolic syndrome and male fertility disorders: Is there a causal link?","volume":"22","author":"Cohen","year":"2021","journal-title":"Rev. Endocr. Metab. Disord."}],"container-title":["Cells"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4409\/15\/9\/808\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,4,29]],"date-time":"2026-04-29T13:46:22Z","timestamp":1777470382000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4409\/15\/9\/808"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,29]]},"references-count":129,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,5]]}},"alternative-id":["cells15090808"],"URL":"https:\/\/doi.org\/10.3390\/cells15090808","relation":{},"ISSN":["2073-4409"],"issn-type":[{"value":"2073-4409","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,4,29]]}}}