{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T03:31:23Z","timestamp":1788233483425,"version":"build-2803163510"},"reference-count":90,"publisher":"MDPI AG","issue":"1","license":[{"start":{"date-parts":[[2024,12,24]],"date-time":"2024-12-24T00:00:00Z","timestamp":1734998400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32102578"],"award-info":[{"award-number":["32102578"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["U22A20515"],"award-info":[{"award-number":["U22A20515"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["2023YFD1302300"],"award-info":[{"award-number":["2023YFD1302300"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"National Key R&amp;D Program of China","doi-asserted-by":"publisher","award":["32102578"],"award-info":[{"award-number":["32102578"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"National Key R&amp;D Program of China","doi-asserted-by":"publisher","award":["U22A20515"],"award-info":[{"award-number":["U22A20515"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"National Key R&amp;D Program of China","doi-asserted-by":"publisher","award":["2023YFD1302300"],"award-info":[{"award-number":["2023YFD1302300"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Antioxidants"],"abstract":"<jats:p>This review systematically examines the influence of oxidative stress on the reproductive function of male livestock, with a particular focus on the modulation of autophagy. Spermatogenesis, a highly precise biological process, is vulnerable to a range of internal and external factors, among which oxidative stress notably disrupts autophagic processes within the testes. This disruption results in diminished sperm quality, impaired testosterone synthesis, and compromised integrity of the blood\u2013testis barrier. Furthermore, this review elucidates the molecular mechanisms by which oxidative stress-induced autophagy dysfunction impairs spermatogenesis and mitochondrial function, consequently reducing sperm motility. These findings aim to provide a theoretical foundation and serve as a reference for improving reproductive performance and sperm quality in livestock.<\/jats:p>","DOI":"10.3390\/antiox14010002","type":"journal-article","created":{"date-parts":[[2024,12,24]],"date-time":"2024-12-24T09:00:57Z","timestamp":1735030857000},"page":"2","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":7,"title":["Oxidative Stress and Autophagy: Unraveling the Hidden Threat to Boars\u2019 Fertility"],"prefix":"10.3390","volume":"14","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5256-927X","authenticated-orcid":false,"given":"Ruizhi","family":"Hu","sequence":"first","affiliation":[{"name":"National Engineering Laboratory for Pollution Control and Waste Utilization in Livestock and Poultry Production, Institute of Subtropical Agriculture, The Chinese Academy of Sciences, Changsha 410125, China"},{"name":"Hunan Collaborative Innovation Center for Utilization of Botanical Functional Ingredients, College of Animal Science and Technology, Hunan Agricultural University, Changsha 410128, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xizi","family":"Yang","sequence":"additional","affiliation":[{"name":"Hunan Collaborative Innovation Center for Utilization of Botanical Functional Ingredients, College of Animal Science and Technology, Hunan Agricultural University, Changsha 410128, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9289-4405","authenticated-orcid":false,"given":"Jianhua","family":"He","sequence":"additional","affiliation":[{"name":"Hunan Collaborative Innovation Center for Utilization of Botanical Functional Ingredients, College of Animal Science and Technology, Hunan Agricultural University, Changsha 410128, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3532-1165","authenticated-orcid":false,"given":"Shusong","family":"Wu","sequence":"additional","affiliation":[{"name":"Hunan Collaborative Innovation Center for Utilization of Botanical Functional Ingredients, College of Animal Science and Technology, Hunan Agricultural University, Changsha 410128, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2024,12,24]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"420","DOI":"10.1002\/mrd.23666","article-title":"One Health challenges for pig reproduction","volume":"90","author":"Peltoniemi","year":"2023","journal-title":"Mol. Reprod. Dev."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"114505","DOI":"10.1016\/j.envpol.2020.114505","article-title":"Effects of endocrine disrupting chemicals in pigs","volume":"263","author":"Yang","year":"2020","journal-title":"Environ. Pollut."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"969","DOI":"10.1002\/mrd.22841","article-title":"Scrotal insulation and sperm production in the boar","volume":"84","author":"Parrish","year":"2017","journal-title":"Mol. Reprod. Dev."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1016\/j.theriogenology.2017.01.038","article-title":"Linear model analysis of the influencing factors of boar longevity in Southern China","volume":"93","author":"Wang","year":"2017","journal-title":"Theriogenology"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"133375","DOI":"10.1016\/j.jhazmat.2023.133375","article-title":"A meta-analysis-based adverse outcome pathway for the male reproductive toxicity induced by microplastics and nanoplastics in mammals","volume":"465","author":"Hu","year":"2024","journal-title":"J. Hazard. Mater."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1566","DOI":"10.1016\/j.bbagen.2014.10.020","article-title":"ROS, thiols and thiol-regulating systems in male gametogenesis","volume":"1850","author":"Conrad","year":"2015","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1111\/rda.14249","article-title":"Linseed oil in boar\u2019s diet improved in vivo fertility and antioxidant status","volume":"58","author":"Singh","year":"2023","journal-title":"Reprod. Domest. Anim."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1016\/j.anireprosci.2016.04.002","article-title":"Antioxidant effects of cultured wild ginseng root extracts on the male reproductive function of boars and guinea pigs","volume":"170","author":"Yun","year":"2016","journal-title":"Anim. Reprod. Sci."},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Zhou, Y., Wei, Z., Tan, J., Sun, H., Jiang, H., Gao, Y., Zhang, H., and Schroyen, M. (2023). Alginate oligosaccharide extends the service lifespan by improving the sperm metabolome and gut microbiota in an aging Duroc boars model. Front. Cell. Infect. Microbiol., 13.","DOI":"10.3389\/fcimb.2023.1308484"},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Guo, H., Ouyang, Y., Yin, H., Cui, H., Deng, H., Liu, H., Jian, Z., Fang, J., Zuo, Z., and Wang, X. (2022). Induction of autophagy via the ROS-dependent AMPK-mTOR pathway protects copper-induced spermatogenesis disorder. Redox Biol., 49.","DOI":"10.1016\/j.redox.2021.102227"},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Sharma, P., Kaushal, N., Saleth, L.R., Ghavami, S., Dhingra, S., and Kaur, P. (2023). Oxidative stress-induced apoptosis and autophagy: Balancing the contrary forces in spermatogenesis. Biochim. Biophys. Acta Mol. Basis Dis., 1869.","DOI":"10.1016\/j.bbadis.2023.166742"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"2313","DOI":"10.1016\/j.devcel.2021.07.015","article-title":"Autophagic elimination of ribosomes during spermiogenesis provides energy for flagellar motility","volume":"56","author":"Lei","year":"2021","journal-title":"Dev. Cell"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"881","DOI":"10.1038\/aja.2011.85","article-title":"Autophagic deficiency is related to steroidogenic decline in aged rat Leydig cells","volume":"13","author":"Li","year":"2011","journal-title":"Asian J. Androl."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1083\/jcb.3.3.349","article-title":"Cellular differentiation in the kidneys of newborn mice studies with the electron microscope","volume":"3","author":"Clark","year":"1957","journal-title":"J. Biophys. Biochem. Cytol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1542","DOI":"10.1038\/sj.cdd.4401765","article-title":"Autophagy: Molecular machinery for self-eating","volume":"12","author":"Yorimitsu","year":"2005","journal-title":"Cell Death Differ."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"814","DOI":"10.1038\/ncb0910-814","article-title":"Eaten alive: A history of macroautophagy","volume":"12","author":"Yang","year":"2010","journal-title":"Nat. Cell Biol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1007\/BF02912068","article-title":"Autophagy, heterophagy, microautophagy and crinophagy as the means for intracellular degradation","volume":"36","author":"Marzella","year":"1981","journal-title":"Virchows Arch. B Cell Pathol. Incl. Mol. Pathol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1242\/jcs.073874","article-title":"Chaperone-mediated autophagy at a glance","volume":"124","author":"Kaushik","year":"2011","journal-title":"J. Cell Sci."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1016\/j.lfs.2017.08.029","article-title":"Autophagy: The spotlight for cellular stress responses","volume":"188","author":"Ravanan","year":"2017","journal-title":"Life Sci."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1038\/s41580-019-0199-y","article-title":"mTOR at the nexus of nutrition, growth, ageing and disease","volume":"21","author":"Liu","year":"2020","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"585","DOI":"10.1042\/EBC20170021","article-title":"The mammalian ULK1 complex and autophagy initiation","volume":"61","author":"Zachari","year":"2017","journal-title":"Essays Biochem."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1038\/s41418-022-01057-1","article-title":"Bud31-mediated alternative splicing is required for spermatogonial stem cell self-renewal and differentiation","volume":"30","author":"Qin","year":"2023","journal-title":"Cell Death Differ."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1753","DOI":"10.1080\/15548627.2020.1783822","article-title":"Autophagy core protein ATG5 is required for elongating spermatid development, sperm individualization and normal fertility in male mice","volume":"17","author":"Huang","year":"2021","journal-title":"Autophagy"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"4967","DOI":"10.1093\/jas\/sky376","article-title":"Macro-environmental sensitivity for growth rate in Danish Duroc pigs is under genetic control","volume":"96","author":"Madsen","year":"2018","journal-title":"J. Anim. Sci."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"120434","DOI":"10.1016\/j.lfs.2022.120434","article-title":"Vitamin E rescues valproic acid-induced testicular injury in rats: Role of autophagy","volume":"296","author":"Alsemeh","year":"2022","journal-title":"Life Sci."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Zhou, Y., Chen, L., Han, H., Xiong, B., Zhong, R., Jiang, Y., Liu, L., Sun, H., Tan, J., and Cheng, X. (2022). Taxifolin increased semen quality of Duroc boars by improving gut microbes and blood metabolites. Front. Microbiol., 13.","DOI":"10.3389\/fmicb.2022.1020628"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1093\/tas\/txaa003","article-title":"The effect of varicocele on semen quality in boars exposed to heat stress","volume":"4","author":"Gruhot","year":"2020","journal-title":"Transl. Anim. Sci."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"170","DOI":"10.1111\/j.1439-0531.2012.02072.x","article-title":"The effects of increased testicular temperature on testis-specific isoform of Na+\/K+ -ATPase in sperm and its role in spermatogenesis and sperm function","volume":"47","author":"Thundathil","year":"2012","journal-title":"Reprod. Domest. Anim."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"169","DOI":"10.4103\/aja.aja_14_19","article-title":"Mechanistic target of rapamycin kinase (Mtor) is required for spermatogonial proliferation and differentiation in mice","volume":"22","author":"Cao","year":"2020","journal-title":"Asian J. Androl."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"4029","DOI":"10.3892\/mmr.2017.7120","article-title":"Rapamycin inhibits spermatogenesis by changing the autophagy status through suppressing mechanistic target of rapamycin-p70S6 kinase in male rats","volume":"16","author":"Liu","year":"2017","journal-title":"Mol. Med. Rep."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"2215","DOI":"10.1089\/ars.2010.3554","article-title":"Autophagy signaling through reactive oxygen species","volume":"14","author":"Huang","year":"2011","journal-title":"Antioxid. Redox Signal."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"5486","DOI":"10.1038\/s41467-019-13114-4","article-title":"Global redox proteome and phosphoproteome analysis reveals redox switch in Akt","volume":"10","author":"Su","year":"2019","journal-title":"Nat. Commun."},{"key":"ref_33","first-page":"1749","article-title":"Reactive oxygen species are essential for autophagy and specifically regulate the activity of Atg4","volume":"38","author":"Shvets","year":"2019","journal-title":"EMBO J."},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"S\u00e1nchez-\u00c1lvarez, M., Strippoli, R., Donadelli, M., Bazhin, A.V., and Cordani, M. (2019). Sestrins as a Therapeutic Bridge between ROS and Autophagy in Cancer. Cancers, 11.","DOI":"10.3390\/cancers11101415"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1016\/j.canlet.2008.09.042","article-title":"Polygonatum cyrtonema lectin induces apoptosis and autophagy in human melanoma A375 cells through a mitochondria-mediated ROS-p38-p53 pathway","volume":"275","author":"Liu","year":"2009","journal-title":"Cancer Lett."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1016\/S0014-5793(04)00112-7","article-title":"Redox regulation of PTEN and protein tyrosine phosphatases in H2O2 mediated cell signaling","volume":"560","author":"Cho","year":"2004","journal-title":"FEBS Lett."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1007\/s13238-020-00771-1","article-title":"Sirt1 regulates testosterone biosynthesis in Leydig cells via modulating autophagy","volume":"12","author":"Khawar","year":"2021","journal-title":"Protein Cell"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1080\/19396368.2019.1606361","article-title":"Autophagy in male reproduction","volume":"65","author":"Zhu","year":"2019","journal-title":"Syst. Biol. Reprod. Med."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Qiao, N., Chen, H., Du, P., Kang, Z., Pang, C., Liu, B., Zeng, Q., Pan, J., Zhang, H., and Mehmood, K. (2021). Acetyl-L-Carnitine Induces Autophagy to Promote Mouse Spermatogonia Cell Recovery after Heat Stress Damage. Biomed. Res. Int., 2021.","DOI":"10.1155\/2021\/8871328"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1673","DOI":"10.1210\/endo-119-4-1673","article-title":"The phagocytic function of Sertoli cells: A morphological, biochemical, and endocrinological study of lysosomes and acid phosphatase localization in the rat testis","volume":"119","author":"Chemes","year":"1986","journal-title":"Endocrinology"},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Zhang, M., Jiang, M., Bi, Y., Zhu, H., Zhou, Z., and Sha, J. (2012). Autophagy and apoptosis act as partners to induce germ cell death after heat stress in mice. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0041412"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"107220","DOI":"10.1016\/j.envint.2022.107220","article-title":"Repression of autophagy leads to acrosome biogenesis disruption caused by a sub-chronic oral administration of polystyrene nanoparticles","volume":"163","author":"Zhou","year":"2022","journal-title":"Environ. Int."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"56791","DOI":"10.18632\/oncotarget.18221","article-title":"In vivo autophagy and biogenesis of autophagosomes within male haploid cells during spermiogenesis","volume":"8","author":"Yang","year":"2017","journal-title":"Oncotarget"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"441","DOI":"10.1242\/dev.147074","article-title":"Sirt1 regulates acrosome biogenesis by modulating autophagic flux during spermiogenesis in mice","volume":"144","author":"Liu","year":"2017","journal-title":"Development"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"653","DOI":"10.4161\/auto.23839","article-title":"A chimerical phagocytosis model reveals the recruitment by Sertoli cells of autophagy for the degradation of ingested illegitimate substrates","volume":"9","author":"Yefimova","year":"2013","journal-title":"Autophagy"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"814","DOI":"10.1080\/15548627.2016.1159377","article-title":"Autophagy is required for ectoplasmic specialization assembly in sertoli cells","volume":"12","author":"Liu","year":"2016","journal-title":"Autophagy"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"11100","DOI":"10.1074\/jbc.275.15.11100","article-title":"CLP-36 PDZ-LIM protein associates with nonmuscle alpha-actinin-1 and alpha-actinin-4","volume":"275","author":"Vallenius","year":"2000","journal-title":"J. Biol. Chem."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"3993","DOI":"10.1007\/s11033-010-0517-0","article-title":"The role of actin and myosin during spermatogenesis","volume":"38","author":"Sun","year":"2011","journal-title":"Mol. Biol. Rep."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1016\/j.semcdb.2014.04.007","article-title":"The roles and regulation of Sertoli cells in fate determinations of spermatogonial stem cells and spermatogenesis","volume":"29","author":"Hai","year":"2014","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Ma, Y., Yang, H.Z., Xu, L.M., Huang, Y.R., Dai, H.L., and Kang, X.N. (2015). Testosterone regulates the autophagic clearance of androgen binding protein in rat Sertoli cells. Sci. Rep., 5.","DOI":"10.1038\/srep08894"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1007\/s10735-011-9372-0","article-title":"Enhanced mitophagy in Sertoli cells of ethanol-treated rats: Morphological evidence and clinical relevance","volume":"43","author":"Eid","year":"2012","journal-title":"J. Mol. Histol."},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Yang, Y., Yao, M., Zeng, J., Zheng, D., Li, Q., Ni, Y., and Xiao, X. (2022). FYN regulates cell adhesion at the blood-testis barrier and the apical ectoplasmic specialization via its effect on Arp3 in the mouse testis. Front. Immunol., 13.","DOI":"10.3389\/fimmu.2022.915274"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1708251","DOI":"10.1155\/2023\/1708251","article-title":"Curcumin Ameliorates Age-Induced Tight Junction Impaired in Porcine Sertoli Cells by Inactivating the NLRP3 Inflammasome through the AMPK\/SIRT3\/SOD2\/mtROS Signaling Pathway","volume":"2023","author":"Xiao","year":"2023","journal-title":"Oxidative Med. Cell. Longev."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1016\/j.ecoenv.2018.09.073","article-title":"Sodium Selenite inhibits mitophagy, downregulation and mislocalization of blood-testis barrier proteins of bovine Sertoli cell exposed to microcystin-leucine arginine (MC-LR) via TLR4\/NF-kB and mitochondrial signaling pathways blockage","volume":"166","author":"Adegoke","year":"2018","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"134126","DOI":"10.1016\/j.jhazmat.2024.134126","article-title":"The FAK\/occludin\/ZO-1 complex is critical for cadmium-induced testicular damage by disruption of the integrity of the blood-testis barrier in chickens","volume":"470","author":"Li","year":"2024","journal-title":"J. Hazard. Mater."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"116865","DOI":"10.1016\/j.jep.2023.116865","article-title":"Wuzi Yanzong administration alleviates Sertoli cell injury by recovering AKT\/mTOR-mediated autophagy and the mTORC1-mTROC2 balance in aging-induced testicular dysfunction","volume":"318","author":"Chang","year":"2024","journal-title":"J. Ethnopharmacol."},{"key":"ref_57","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_58","doi-asserted-by":"crossref","unstructured":"Lin, Y., Wu, D., Che, L., Fang, Z., Xu, S., Feng, B., Zhuo, Y., Li, J., Wu, C., and Zhang, J. (2022). Dietary Fibre Supplementation Improves Semen Production by Increasing Leydig Cells and Testosterone Synthesis in a Growing Boar Model. Front. Vet. Sci., 9.","DOI":"10.3389\/fvets.2022.850685"},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Kirat, D., Alahwany, A.M., Arisha, A.H., Abdelkhalek, A., and Miyasho, T. (2023). Role of Macroautophagy in Mammalian Male Reproductive Physiology. Cells, 12.","DOI":"10.3390\/cells12091322"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1002\/j.1939-4640.2005.tb02868.x","article-title":"Temporal relationships among testosterone production, steroidogenic acute regulatory protein (StAR), and P450 side-chain cleavage enzyme (P450scc) during Leydig cell aging","volume":"26","author":"Luo","year":"2005","journal-title":"J. Androl."},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Hu, R., Yang, X., Gong, J., Lv, J., Yuan, X., Shi, M., Fu, C., Tan, B., Fan, Z., and Chen, L. (2024). Patterns of alteration in boar semen quality from 9 to 37 months old and improvement by protocatechuic acid. J. Anim. Sci. Biotechnol., 15.","DOI":"10.1186\/s40104-024-01031-6"},{"key":"ref_62","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-AMPK-mTOR axis","volume":"10","author":"Li","year":"2019","journal-title":"Cell Death Dis."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"2632","DOI":"10.1111\/jcmm.14157","article-title":"Fibroblast growth factor 16 stimulates proliferation but blocks differentiation of rat stem Leydig cells during regeneration","volume":"23","author":"Duan","year":"2019","journal-title":"J. Cell. Mol. Med."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.bbrc.2018.04.078","article-title":"Hyperhomocysteinemia and hyperandrogenemia share PCSK9-LDLR pathway to disrupt lipid homeostasis in PCOS","volume":"503","author":"Mondal","year":"2018","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1023\/A:1007178532236","article-title":"Rat Leydig cells use apolipoprotein E depleted high density lipoprotein to regulate testosterone production","volume":"213","author":"Travert","year":"2000","journal-title":"Mol. Cell. Biochem."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"2103","DOI":"10.1083\/jcb.201710078","article-title":"Autophagy regulates testosterone synthesis by facilitating cholesterol uptake in Leydig cells","volume":"217","author":"Gao","year":"2018","journal-title":"J. Cell Biol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1016\/j.yexcr.2015.11.024","article-title":"Lipid droplets and steroidogenic cells","volume":"340","author":"Shen","year":"2016","journal-title":"Exp. Cell Res."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1119","DOI":"10.1210\/en.2017-03020","article-title":"Lipophagy Contributes to Testosterone Biosynthesis in Male Rat Leydig Cells","volume":"159","author":"Ma","year":"2018","journal-title":"Endocrinology"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1016\/j.theriogenology.2024.09.010","article-title":"Sirtuin 1-mediated autophagy regulates testosterone synthesis in Leydig cells of piglets","volume":"230","author":"Zhang","year":"2024","journal-title":"Theriogenology"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1625","DOI":"10.1038\/s41556-023-01264-z","article-title":"Mitopherogenesis, a form of mitochondria-specific ectocytosis, regulates sperm mitochondrial quantity and fertility","volume":"25","author":"Liu","year":"2023","journal-title":"Nat. Cell Biol."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1093\/humupd\/dmp016","article-title":"Mitochondrial functionality in reproduction: From gonads and gametes to embryos and embryonic stem cells","volume":"15","author":"Varum","year":"2009","journal-title":"Hum. Reprod. Update"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"R163","DOI":"10.1530\/REP-13-0178","article-title":"Mitochondria functionality and sperm quality","volume":"146","author":"Amaral","year":"2013","journal-title":"Reproduction"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1016\/j.mito.2017.01.001","article-title":"Mitochondria in teleost spermatozoa","volume":"34","author":"Figueroa","year":"2017","journal-title":"Mitochondrion"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"697","DOI":"10.1093\/humupd\/dmab001","article-title":"Mitochondria: Their role in spermatozoa and in male infertility","volume":"27","author":"Boguenet","year":"2021","journal-title":"Hum. Reprod. Update"},{"key":"ref_75","doi-asserted-by":"crossref","unstructured":"Ma, K., Chen, G., Li, W., Kepp, O., Zhu, Y., and Chen, Q. (2020). Mitophagy, Mitochondrial Homeostasis, and Cell Fate. Front. Cell Dev. Biol., 8.","DOI":"10.3389\/fcell.2020.00467"},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Yogo, K. (2022). Molecular basis of the morphogenesis of sperm head and tail in mice. Reprod. Med. Biol., 21.","DOI":"10.1002\/rmb2.12466"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1016\/j.freeradbiomed.2019.06.018","article-title":"Negative effects of ROS generated during linear sperm motility on gene expression and ATP generation in boar sperm mitochondria","volume":"141","author":"Zhu","year":"2019","journal-title":"Free. Radic. Biol. Med."},{"key":"ref_78","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_79","doi-asserted-by":"crossref","first-page":"587","DOI":"10.1071\/RD09078","article-title":"Antiperoxidative and anti-apoptotic effects of lycopene and ellagic acid on cyclophosphamide-induced testicular lipid peroxidation and apoptosis","volume":"22","author":"Sakin","year":"2010","journal-title":"Reprod. Fertil. Dev."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"26","DOI":"10.1016\/j.mito.2020.06.003","article-title":"Mitochondria, spermatogenesis, and male infertility\u2014An update","volume":"54","author":"Vertika","year":"2020","journal-title":"Mitochondrion"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1801","DOI":"10.4161\/auto.25884","article-title":"The TOMM machinery is a molecular switch in PINK1 and PARK2\/PARKIN-dependent mitochondrial clearance","volume":"9","author":"Bertolin","year":"2013","journal-title":"Autophagy"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"1188","DOI":"10.1021\/cb5010417","article-title":"Chemogenomic profiling of endogenous PARK2 expression using a genome-edited coincidence reporter","volume":"10","author":"Hasson","year":"2015","journal-title":"ACS Chem. Biol."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"737","DOI":"10.1016\/j.cell.2013.07.030","article-title":"A neo-substrate that amplifies catalytic activity of parkinson\u2019s-disease-related kinase PINK1","volume":"154","author":"Hertz","year":"2013","journal-title":"Cell"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"e104705","DOI":"10.15252\/embj.2020104705","article-title":"Molecular mechanisms and physiological functions of mitophagy","volume":"40","author":"Onishi","year":"2021","journal-title":"EMBO J."},{"key":"ref_85","doi-asserted-by":"crossref","unstructured":"Poole, L.P., Bock-Hughes, A., Berardi, D.E., and Macleod, K.F. (2021). ULK1 promotes mitophagy via phosphorylation and stabilization of BNIP3. Sci. Rep., 11.","DOI":"10.1038\/s41598-021-00170-4"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"17590914231159226","DOI":"10.1177\/17590914231159226","article-title":"Thioredoxin-1 Promotes Mitochondrial Biogenesis Through Regulating AMPK\/Sirt1\/PGC1\u03b1 Pathway in Alzheimer\u2019s Disease","volume":"15","author":"Jia","year":"2023","journal-title":"ASN Neuro"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"928","DOI":"10.1038\/s41419-022-05364-w","article-title":"Cadmium-induced apoptosis of Leydig cells is mediated by excessive mitochondrial fission and inhibition of mitophagy","volume":"13","author":"Yi","year":"2022","journal-title":"Cell Death Dis."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"362","DOI":"10.1016\/j.theriogenology.2017.12.017","article-title":"Influences of different dietary energy level on sheep testicular development associated with AMPK\/ULK1\/autophagy pathway","volume":"108","author":"Pang","year":"2018","journal-title":"Theriogenology"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.theriogenology.2021.12.032","article-title":"Dietary folic acid supplementation improves semen quality and spermatogenesis through altering autophagy and histone methylation in the testis of aged broiler breeder roosters","volume":"181","author":"Ye","year":"2022","journal-title":"Theriogenology"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1016\/j.theriogenology.2022.05.030","article-title":"Dietary supplementation with metformin improves testis function and semen quality and increases antioxidants and autophagy capacity in goats","volume":"188","author":"Zhao","year":"2022","journal-title":"Theriogenology"}],"container-title":["Antioxidants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-3921\/14\/1\/2\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T16:59:25Z","timestamp":1760115565000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-3921\/14\/1\/2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,12,24]]},"references-count":90,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,1]]}},"alternative-id":["antiox14010002"],"URL":"https:\/\/doi.org\/10.3390\/antiox14010002","relation":{},"ISSN":["2076-3921"],"issn-type":[{"value":"2076-3921","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,12,24]]}}}