{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T15:58:29Z","timestamp":1791388709300,"version":"4.3.3"},"reference-count":41,"publisher":"Springer Science and Business Media LLC","issue":"10","license":[{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"funder":[{"DOI":"10.13039\/501100009997","name":"Earmarked Fund for Modern Agro-industry Technology Research System","doi-asserted-by":"publisher","award":["CARS-43"],"award-info":[{"award-number":["CARS-43"]}],"id":[{"id":"10.13039\/501100009997","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Reprod. Sci."],"published-print":{"date-parts":[[2026,10]]},"DOI":"10.1007\/s43032-026-02202-8","type":"journal-article","created":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T19:42:27Z","timestamp":1790192547000},"page":"2068-2080","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Beclin1 Regulates Sertoli Cell Function to Maintain Mouse Spermatogenesis"],"prefix":"10.1007","volume":"33","author":[{"given":"Yuqing","family":"Cai","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wanzhen","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yifeilong","family":"He","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhifeng","family":"Zhao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinyi","family":"Lin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jing","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuchuan","family":"Luo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yinghe","family":"Qin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2675-2625","authenticated-orcid":false,"given":"Yingjie","family":"Wu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,9,23]]},"reference":[{"issue":"1","key":"2202_CR1","doi-asserted-by":"publisher","first-page":"14","DOI":"10.4161\/spmg.1.1.14525","volume":"1","author":"L O\u2019Donnell","year":"2011","unstructured":"O\u2019Donnell L, Nicholls PK, O\u2019Bryan MK, et al. Spermiation: the process of sperm release. Spermatogenesis. 2011;1(1):14\u201335.","journal-title":"Spermatogenesis"},{"issue":"1546","key":"2202_CR2","doi-asserted-by":"publisher","first-page":"1501","DOI":"10.1098\/rstb.2009.0124","volume":"365","author":"C Shaha","year":"2010","unstructured":"Shaha C, Tripathi R, Mishra DP. Male germ cell apoptosis: regulation and biology. Philos Trans R Soc Lond B Biol Sci. 2010;365(1546):1501\u201315.","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"issue":"4","key":"2202_CR3","doi-asserted-by":"publisher","first-page":"411","DOI":"10.1006\/scdb.1998.0203","volume":"9","author":"MD Griswold","year":"1998","unstructured":"Griswold MD. The central role of Sertoli cells in spermatogenesis. Semin Cell Dev Biol. 1998;9(4):411\u20136.","journal-title":"Semin Cell Dev Biol"},{"issue":"1","key":"2202_CR4","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1093\/biolre\/ioy027","volume":"99","author":"MD Griswold","year":"2018","unstructured":"Griswold MD. 50 years of spermatogenesis: Sertoli cells and their interactions with germ cells. Biol Reprod. 2018;99(1):87\u2013100.","journal-title":"Biol Reprod"},{"issue":"7","key":"2202_CR5","doi-asserted-by":"publisher","first-page":"345","DOI":"10.1016\/j.tem.2004.07.003","volume":"15","author":"F Boussouar","year":"2004","unstructured":"Boussouar F, Benahmed M. Lactate and energy metabolism in male germ cells. Trends Endocrinol Metab. 2004;15(7):345\u201350.","journal-title":"Trends Endocrinol Metab"},{"issue":"2","key":"2202_CR6","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1002\/aja.1001670202","volume":"167","author":"V Wong","year":"1983","unstructured":"Wong V, Russell LD. Three-dimensional reconstruction of a rat stage V Sertoli cell: I. Methods, basic configuration, and dimensions. Am J Anat. 1983;167(2):143\u201361.","journal-title":"Am J Anat"},{"issue":"5","key":"2202_CR7","doi-asserted-by":"publisher","first-page":"564","DOI":"10.1210\/er.2014-1101","volume":"36","author":"DD Mruk","year":"2015","unstructured":"Mruk DD, Cheng CY. The mammalian blood\u2013testis barrier: its biology and regulation. Endocr Rev. 2015;36(5):564\u201391.","journal-title":"Endocr Rev"},{"issue":"2","key":"2202_CR8","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1152\/physrev.00025.2011","volume":"92","author":"JM Oatley","year":"2012","unstructured":"Oatley JM, Brinster RL. The germline stem cell niche unit in mammalian testes. Physiol Rev. 2012;92(2):577\u201395.","journal-title":"Physiol Rev"},{"key":"2202_CR9","first-page":"69","volume-title":"Male Reproductive Health","author":"F Wang","year":"2019","unstructured":"Wang F, Han D. Sertoli cell phagocytosis: an essential event for spermatogenesis. In: Wu W, Ziglioli F, Maestroni U, editors. Male reproductive health. London: IntechOpen; 2019. pp. 69\u201384."},{"issue":"5","key":"2202_CR10","doi-asserted-by":"publisher","first-page":"795","DOI":"10.1083\/jcb.200809125","volume":"183","author":"D Narendra","year":"2008","unstructured":"Narendra D, Tanaka A, Suen DF, Youle RJ. Parkin is recruited selectively to impaired mitochondria and promotes their autophagy. J Cell Biol. 2008;183(5):795\u2013803.","journal-title":"J Cell Biol"},{"issue":"6","key":"2202_CR11","doi-asserted-by":"publisher","first-page":"1149","DOI":"10.1016\/j.cell.2007.10.035","volume":"131","author":"M Komatsu","year":"2007","unstructured":"Komatsu M, Waguri S, Koike M, Sou YS, Ueno T, Hara T, et al. Homeostatic levels of p62 control cytoplasmic inclusion body formation in autophagy-deficient mice. Cell. 2007;131(6):1149\u201363.","journal-title":"Cell"},{"issue":"3","key":"2202_CR12","doi-asserted-by":"publisher","first-page":"1101","DOI":"10.1091\/mbc.e03-09-0704","volume":"15","author":"N Mizushima","year":"2004","unstructured":"Mizushima N, Yamamoto A, Matsui M, Yoshimori T, Ohsumi Y. In vivo analysis of autophagy in response to nutrient starvation using transgenic mice expressing a fluorescent autophagosome marker. Mol Biol Cell. 2004;15(3):1101\u201311.","journal-title":"Mol Biol Cell"},{"issue":"7","key":"2202_CR13","doi-asserted-by":"publisher","first-page":"651","DOI":"10.1056\/NEJMra1205406","volume":"368","author":"AM Choi","year":"2013","unstructured":"Choi AM, Ryter SW, Levine B. Autophagy in human health and disease. N Engl J Med. 2013;368(7):651\u201362.","journal-title":"N Engl J Med"},{"issue":"21","key":"2202_CR14","doi-asserted-by":"publisher","first-page":"5720","DOI":"10.1093\/emboj\/19.21.5720","volume":"19","author":"Y Kabeya","year":"2000","unstructured":"Kabeya Y, Mizushima N, Ueno T, Yamamoto A, Kirisako T, Noda T, et al. LC3, a mammalian homologue of yeast Apg8p, is localized in autophagosome membranes after processing. EMBO J. 2000;19(21):5720\u20138.","journal-title":"EMBO J"},{"key":"2202_CR15","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1146\/annurev-cellbio-092910-154005","volume":"27","author":"N Mizushima","year":"2011","unstructured":"Mizushima N, Yoshimori T, Ohsumi Y. The role of Atg proteins in autophagosome formation. Annu Rev Cell Dev Biol. 2011;27:107\u201332.","journal-title":"Annu Rev Cell Dev Biol"},{"issue":"2","key":"2202_CR16","doi-asserted-by":"publisher","first-page":"200","DOI":"10.1093\/humupd\/dmab043","volume":"28","author":"M Wang","year":"2022","unstructured":"Wang M, Zeng L, Su P, et al. Autophagy: a multifaceted player in the fate of sperm. Hum Reprod Update. 2022;28(2):200\u201331.","journal-title":"Hum Reprod Update"},{"issue":"4","key":"2202_CR17","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1080\/19396368.2019.1606361","volume":"65","author":"Y Zhu","year":"2019","unstructured":"Zhu Y, Yin Q, Wei D, et al. Autophagy in male reproduction. Syst Biol Reprod Med. 2019;65(4):265\u201372.","journal-title":"Syst Biol Reprod Med"},{"issue":"7","key":"2202_CR18","doi-asserted-by":"publisher","first-page":"852","DOI":"10.1038\/cr.2014.70","volume":"24","author":"H Wang","year":"2014","unstructured":"Wang H, Wan H, Li X, et al. Atg7 is required for acrosome biogenesis during spermatogenesis in mice. Cell Res. 2014;24(7):852\u201369.","journal-title":"Cell Res"},{"issue":"5","key":"2202_CR19","doi-asserted-by":"publisher","first-page":"814","DOI":"10.1080\/15548627.2016.1159377","volume":"12","author":"C Liu","year":"2016","unstructured":"Liu C, Wang H, Shang Y, et al. Autophagy is required for ectoplasmic specialization assembly in Sertoli cells. Autophagy. 2016;12(5):814\u201332.","journal-title":"Autophagy"},{"key":"2202_CR20","doi-asserted-by":"publisher","first-page":"1575","DOI":"10.1080\/15548627.2016.1192750","volume":"12","author":"Y Shang","year":"2016","unstructured":"Shang Y, Wang H, Jia P, et al. Autophagy regulates spermatid differentiation via degradation of PDLIM1. Autophagy. 2016;12:1575\u201392.","journal-title":"Autophagy"},{"issue":"11","key":"2202_CR21","doi-asserted-by":"publisher","first-page":"2934","DOI":"10.1080\/15548627.2023.2235195","volume":"19","author":"K Wang","year":"2023","unstructured":"Wang K, Kong F, Qiu Y, et al. Autophagy regulation and protein kinase activity of PIK3C3 controls Sertoli cell polarity through its negative regulation on SCIN (scinderin). Autophagy. 2023;19(11):2934\u201357.","journal-title":"Autophagy"},{"issue":"8","key":"2202_CR22","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1009688","volume":"17","author":"T Yamamuro","year":"2021","unstructured":"Yamamuro T, Nakamura S, Yamano Y, et al. Rubicon prevents autophagic degradation of GATA4 to promote Sertoli cell function. PLoS Genet. 2021;17(8):e1009688.","journal-title":"PLoS Genet"},{"issue":"8","key":"2202_CR23","doi-asserted-by":"publisher","first-page":"604","DOI":"10.1007\/s11626-019-00371-2","volume":"55","author":"J Pang","year":"2019","unstructured":"Pang J, Han L, Liu Z, Zheng J, Zhao J, Deng K, et al. ULK1 affects cell viability of goat Sertoli cell by modulating both autophagy and apoptosis. In Vitro Cell Dev Biol Anim. 2019;55(8):604\u201313.","journal-title":"In Vitro Cell Dev Biol Anim"},{"key":"2202_CR24","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1038\/cdd.2010.191","volume":"18","author":"R Kang","year":"2011","unstructured":"Kang R, Zeh HJ, Lotze MT, Tang D. The Beclin 1 network regulates autophagy and apoptosis. Cell Death Differ. 2011;18:571\u201380.","journal-title":"Cell Death Differ"},{"issue":"6762","key":"2202_CR25","doi-asserted-by":"publisher","first-page":"672","DOI":"10.1038\/45257","volume":"402","author":"XH Liang","year":"1999","unstructured":"Liang XH, Jackson S, Seaman M, Brown K, Kempkes B, Hibshoosh H, et al. Induction of autophagy and inhibition of tumorigenesis by beclin 1. Nature. 1999;402(6762):672\u20136.","journal-title":"Nature"},{"issue":"25","key":"2202_CR26","doi-asserted-by":"publisher","first-page":"15077","DOI":"10.1073\/pnas.2436255100","volume":"100","author":"Z Yue","year":"2003","unstructured":"Yue Z, Jin S, Yang C, Levine AJ, Heintz N. Beclin 1, an autophagy gene essential for early embryonic development, is a haploinsufficient tumor suppressor. Proc Natl Acad Sci U S A. 2003;100(25):15077\u201382.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"6","key":"2202_CR27","doi-asserted-by":"publisher","first-page":"1062","DOI":"10.1016\/j.cell.2009.03.048","volume":"137","author":"R Mathew","year":"2009","unstructured":"Mathew R, Karp CM, Beaudoin B, Vuong N, Chen G, Chen HY, et al. Autophagy suppresses tumorigenesis through elimination of p62. Cell. 2009;137(6):1062\u201375.","journal-title":"Cell"},{"issue":"3","key":"2202_CR28","doi-asserted-by":"publisher","first-page":"599","DOI":"10.1093\/biolre\/ioad160","volume":"110","author":"L Ke","year":"2024","unstructured":"Ke L, Lin X, Luo Y, et al. Autophagy core protein Beclin1 is vital for spermatogenesis and male fertility in mice. Biol Reprod. 2024;110(3):599\u2013614.","journal-title":"Biol Reprod"},{"issue":"7","key":"2202_CR29","doi-asserted-by":"publisher","first-page":"1753","DOI":"10.1080\/15548627.2020.1783822","volume":"17","author":"Q Huang","year":"2021","unstructured":"Huang Q, Liu Y, Zhang S, et al. Autophagy core protein ATG5 is required for elongating spermatid development, sperm individualization and normal fertility in male mice. Autophagy. 2021;17(7):1753\u201367.","journal-title":"Autophagy"},{"key":"2202_CR30","doi-asserted-by":"publisher","first-page":"1811","DOI":"10.15252\/embj.201796697","volume":"36","author":"L Galluzzi","year":"2017","unstructured":"Galluzzi L, Baehrecke EH, Ballabio A, et al. Molecular definitions of autophagy and related processes. EMBO J. 2017;36:1811\u201336.","journal-title":"EMBO J"},{"issue":"2","key":"2202_CR31","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1111\/andr.12165","volume":"4","author":"LR Fran\u00e7a","year":"2016","unstructured":"Fran\u00e7a LR, Hess RA, Dufour JM, et al. The Sertoli cell: one hundred fifty years of beauty and plasticity. Andrology. 2016;4(2):189\u2013212.","journal-title":"Andrology"},{"issue":"5","key":"2202_CR32","doi-asserted-by":"publisher","first-page":"381","DOI":"10.1002\/j.1939-4640.1994.tb00468.x","volume":"15","author":"SR Sylvester","year":"1994","unstructured":"Sylvester SR, Griswold MD. The testicular iron shuttle: a \u201cnurse\u201d function of the Sertoli cells. J Androl. 1994;15(5):381\u20135.","journal-title":"J Androl"},{"issue":"6009","key":"2202_CR33","doi-asserted-by":"publisher","first-page":"1344","DOI":"10.1126\/science.1193497","volume":"330","author":"JD Rabinowitz","year":"2010","unstructured":"Rabinowitz JD, White E. Autophagy and metabolism. Science. 2010;330(6009):1344\u20138.","journal-title":"Science"},{"key":"2202_CR34","doi-asserted-by":"publisher","DOI":"10.1016\/j.theriogenology.2026.117838","volume":"255","author":"X Qi","year":"2026","unstructured":"Qi X, Wu Y, Li Q, Wang H, Ma Y. BECN1-mediated autophagy activates the glycolytic pathway to promote lactate synthesis in Sertoli cells of Tibetan sheep. Theriogenology. 2026;255:117838.","journal-title":"Theriogenology"},{"key":"2202_CR35","doi-asserted-by":"publisher","first-page":"1424","DOI":"10.1002\/stem.464","volume":"28","author":"CJ Payne","year":"2010","unstructured":"Payne CJ, Gallagher SJ, Foreman O, et al. Sin3a is required by Sertoli cells to establish a niche for undifferentiated spermatogonia, germ cell tumors, and spermatid elongation. Stem Cells. 2010;28:1424\u201334.","journal-title":"Stem Cells"},{"issue":"6","key":"2202_CR36","doi-asserted-by":"publisher","first-page":"1565","DOI":"10.1093\/biolre\/ioac166","volume":"107","author":"F Chen","year":"2022","unstructured":"Chen F, Wu Y, Ke L, Lin X, Wang F, Qin Y. ARHGEF15 in Sertoli cells contributes to germ cell development and testicular immune privilege. Biol Reprod. 2022;107(6):1565\u201379.","journal-title":"Biol Reprod"},{"issue":"1546","key":"2202_CR37","doi-asserted-by":"publisher","first-page":"1621","DOI":"10.1098\/rstb.2010.0010","volume":"365","author":"DD Mruk","year":"2010","unstructured":"Mruk DD, Cheng CY. Tight junctions in the testis: new perspectives. Philos Trans R Soc Lond B Biol Sci. 2010;365(1546):1621\u201335.","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"issue":"7","key":"2202_CR38","doi-asserted-by":"publisher","first-page":"300","DOI":"10.1016\/j.tem.2005.07.001","volume":"16","author":"G Pointis","year":"2005","unstructured":"Pointis G, Segretain D. Role of connexin-based gap junction channels in testis. Trends Endocrinol Metab. 2005;16(7):300\u20136.","journal-title":"Trends Endocrinol Metab"},{"issue":"1546","key":"2202_CR39","doi-asserted-by":"publisher","first-page":"1581","DOI":"10.1098\/rstb.2009.0261","volume":"365","author":"PPY Lie","year":"2010","unstructured":"Lie PPY, Mruk DD, Lee WM, et al. Cytoskeletal dynamics and spermatogenesis. Philos Trans R Soc Lond B Biol Sci. 2010;365(1546):1581\u201392.","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"2202_CR40","doi-asserted-by":"crossref","unstructured":"Sakai Y, Koyama-Honda I, Tachikawa M, et al. Modeling membrane morphological change during autophagosome formation. iScience. 2020.","DOI":"10.1016\/j.isci.2020.101466"},{"key":"2202_CR41","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1007\/978-981-16-2830-6_2","volume":"1208","author":"Y Yang","year":"2021","unstructured":"Yang Y, Zheng L, Zheng X, et al. Autophagosomal membrane origin and formation. Adv Exp Med Biol. 2021;1208:17\u201342.","journal-title":"Adv Exp Med Biol."}],"container-title":["Reproductive Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s43032-026-02202-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s43032-026-02202-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s43032-026-02202-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T15:45:24Z","timestamp":1791387924000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s43032-026-02202-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,23]]},"references-count":41,"journal-issue":{"issue":"10","published-print":{"date-parts":[[2026,10]]}},"alternative-id":["2202"],"URL":"https:\/\/doi.org\/10.1007\/s43032-026-02202-8","relation":{},"ISSN":["1933-7191","1933-7205"],"issn-type":[{"value":"1933-7191","type":"print"},{"value":"1933-7205","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,23]]},"assertion":[{"value":"22 March 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"31 August 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"23 September 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"All the animal experimental procedures were approved by the Institutional Animal Care and Use Committee of China Agricultural University (AW22705202-1-03).","order":1,"name":"Ethics","label":"Ethical Approval","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors have declared that no conflict of interest exists.","order":2,"name":"Ethics","label":"Conflict of Interest","group":{"name":"EthicsHeading","label":"Declarations"}}]}}