{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T01:56:52Z","timestamp":1791338212441,"version":"4.3.1"},"reference-count":65,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,8,2]],"date-time":"2025-08-02T00:00:00Z","timestamp":1754092800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2025,8,2]],"date-time":"2025-08-02T00:00:00Z","timestamp":1754092800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"name":"Ankara University Scientific Research Projects","award":["20L0230010"],"award-info":[{"award-number":["20L0230010"]}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"DOI":"10.1038\/s41598-025-12819-5","type":"journal-article","created":{"date-parts":[[2025,8,2]],"date-time":"2025-08-02T03:18:46Z","timestamp":1754104726000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["The relationship between sperm secretion and the Hippo signaling pathway in the rat seminiferous tubules"],"prefix":"10.1038","volume":"15","author":[{"given":"\u0130rem","family":"\u0130nan\u00e7","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Onur","family":"Bender","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Arzu","family":"Atalay","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Esra","family":"Erdemli","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,8,2]]},"reference":[{"key":"12819_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, P. K., O\u2019Bryan, M. K., McLachlan, R. I. & Stanton, P. G. Spermiation: the process of sperm release. Spermatogenesis 1, 14\u201335 (2011).","journal-title":"Spermatogenesis"},{"key":"12819_CR2","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1080\/10409230903061207","volume":"44","author":"CY Cheng","year":"2009","unstructured":"Cheng, C. Y. & Mruk, D. D. An intracellular trafficking pathway in the seminiferous epithelium regulating spermatogenesis: a biochemical and molecular perspective. Crit. Rev. Biochem. Mol. Biol. 44, 245\u2013263 (2009).","journal-title":"Crit. Rev. Biochem. Mol. Biol."},{"key":"12819_CR3","doi-asserted-by":"publisher","first-page":"380","DOI":"10.1038\/nrendo.2010.71","volume":"6","author":"CY Cheng","year":"2010","unstructured":"Cheng, C. Y. & Mruk, D. D. A local autocrine axis in the testes that regulates spermatogenesis. Nat. Rev. Endocrinol. 6, 380\u2013395 (2010).","journal-title":"Nat. Reviews Endocrinol."},{"key":"12819_CR4","doi-asserted-by":"publisher","first-page":"692","DOI":"10.1016\/j.bbamem.2007.11.006","volume":"1778","author":"EW Wong","year":"2008","unstructured":"Wong, E. W., Mruk, D. D. & Cheng, C. Y. Biology and regulation of ectoplasmic specialization, an atypical adherens junction type, in the testis. Biochim. Et Biophys. Acta (BBA) Biomembranes. 1778, 692\u2013708 (2008).","journal-title":"Biochim. Et Biophys. Acta (BBA)-Biomembranes"},{"key":"12819_CR5","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1124\/pr.110.002790","volume":"64","author":"CY Cheng","year":"2012","unstructured":"Cheng, C. Y. & Mruk, D. D. The blood-testis barrier and its implications for male contraception. Pharmacol. Rev. 64, 16\u201364 (2012).","journal-title":"Pharmacol. Rev."},{"key":"12819_CR6","doi-asserted-by":"publisher","first-page":"4300","DOI":"10.1210\/en.2017-00660","volume":"158","author":"L Li","year":"2017","unstructured":"Li, L. et al. Sperm release at spermiation is regulated by changes in the organization of actin-and microtubule-based cytoskeletons at the apical ectoplasmic specialization\u2014a study using the adjudin model. Endocrinology 158, 4300\u20134316 (2017).","journal-title":"Endocrinology"},{"key":"12819_CR7","doi-asserted-by":"publisher","first-page":"1614","DOI":"10.1242\/dev.102376","volume":"141","author":"X Varelas","year":"2014","unstructured":"Varelas, X. The Hippo pathway effectors TAZ and YAP in development, homeostasis and disease. Development 141, 1614\u20131626 (2014).","journal-title":"Development"},{"key":"12819_CR8","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/2001-1326-3-22","volume":"3","author":"M Gomez","year":"2014","unstructured":"Gomez, M., Gomez, V. & Hergovich, A. The Hippo pathway in disease and therapy: cancer and beyond. Clin. Transl. Med. 3, 1\u201312 (2014).","journal-title":"Clin. Translational Med."},{"key":"12819_CR9","doi-asserted-by":"publisher","first-page":"1287","DOI":"10.1152\/physrev.00005.2014","volume":"94","author":"S Piccolo","year":"2014","unstructured":"Piccolo, S., Dupont, S. & Cordenonsi, M. The biology of YAP\/TAZ: Hippo signaling and beyond. Physiol. Rev. 94, 1287\u20131312 (2014).","journal-title":"Physiol. Rev."},{"key":"12819_CR10","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1042\/bse0530111","volume":"53","author":"C Cherrett","year":"2012","unstructured":"Cherrett, C., Furutani-Seiki, M. & Bagby, S. The Hippo pathway: key interaction and catalytic domains in organ growth control, stem cell self-renewal and tissue regeneration. Essays Biochem. 53, 111\u2013127 (2012).","journal-title":"Essays Biochem."},{"key":"12819_CR11","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1146\/annurev-biochem-013118-111829","volume":"88","author":"S Ma","year":"2019","unstructured":"Ma, S., Meng, Z., Chen, R. & Guan, K. L. The Hippo pathway: biology and pathophysiology. Annu. Rev. Biochem. 88, 577\u2013604 (2019).","journal-title":"Annu. Rev. Biochem."},{"key":"12819_CR12","doi-asserted-by":"publisher","first-page":"63","DOI":"10.2174\/1872214809666151029113043","volume":"9","author":"YH Cheng","year":"2015","unstructured":"Cheng, Y. H. et al. Adjudin-a male contraceptive with other biological activities. Recent Pat. Endocr. Metab. Immune Drug Discov.. 9, 63\u201373 (2015).","journal-title":"Recent Pat. Endocr., Metab. Immune Drug Discovery"},{"key":"12819_CR13","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1530\/REP-10-0464","volume":"141","author":"KW Mok","year":"2011","unstructured":"Mok, K. W., Mruk, D. D., Lie, P. P., Lui, W. Y. & Cheng, C. Y. Adjudin, a potential male contraceptive, exerts its effects locally in the seminiferous epithelium of mammalian testes. Reprod. (Cambridge England). 141, 571 (2011).","journal-title":"Reprod. (Cambridge England)"},{"key":"12819_CR14","doi-asserted-by":"publisher","first-page":"345","DOI":"10.1016\/j.bcp.2012.11.008","volume":"85","author":"QR Xie","year":"2013","unstructured":"Xie, Q. R. et al. Male contraceptive adjudin is a potential anti-cancer drug. Biochem. Pharmacol. 85, 345\u2013355 (2013).","journal-title":"Biochem. Pharmacol."},{"key":"12819_CR15","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1016\/j.jneuroim.2012.09.012","volume":"254","author":"J Shao","year":"2013","unstructured":"Shao, J. et al. Adjudin attenuates lipopolysaccharide (LPS)-and ischemia-induced microglial activation. J. Neuroimmunol. 254, 83\u201390 (2013).","journal-title":"J. Neuroimmunol."},{"key":"12819_CR16","doi-asserted-by":"crossref","unstructured":"Fran\u00e7a, L. R., Auharek, S. A., Hess, R. A., Dufour, J. M. & Hinton, B. T. Blood-tissue barriers: morphofunctional and immunological aspects of the blood-testis and blood-epididymal barriers. Biol. Regul. Blood-Tissue Barriers 237\u2013259 (2013).","DOI":"10.1007\/978-1-4614-4711-5_12"},{"key":"12819_CR17","unstructured":"Stanton, P. G. Seminars in Cell & Developmental Biology. 166\u2013173 (Elsevier)."},{"key":"12819_CR18","doi-asserted-by":"crossref","unstructured":"Vogl, A. W., Vaid, K. S. & Guttman, J. A. The sertoli cell cytoskeleton. Mol. Mech. Spermatogen. 186\u2013211 (2008).","DOI":"10.1007\/978-0-387-09597-4_11"},{"key":"12819_CR19","doi-asserted-by":"publisher","first-page":"86","DOI":"10.1111\/j.1365-2605.2011.01183.x","volume":"35","author":"KW Mok","year":"2012","unstructured":"Mok, K. W., Mruk, D. D., Lee, W. M. & Cheng, C. Y. Spermatogonial stem cells alone are not sufficient to re-initiate spermatogenesis in the rat testis following adjudin\u2010induced infertility. Int. J. Androl. 35, 86\u2013101 (2012).","journal-title":"Int. J. Androl."},{"key":"12819_CR20","doi-asserted-by":"publisher","first-page":"553","DOI":"10.1042\/BJ20102121","volume":"435","author":"CY Cheng","year":"2011","unstructured":"Cheng, C. Y. & Mruk, D. D. Regulation of spermiogenesis, spermiation and blood\u2013testis barrier dynamics: novel insights from studies on Eps8 and Arp3. Biochem. J. 435, 553\u2013562 (2011).","journal-title":"Biochem. J."},{"key":"12819_CR21","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1016\/j.reprotox.2019.07.002","volume":"89","author":"S Wu","year":"2019","unstructured":"Wu, S. et al. mTORC1\/rpS6 and spermatogenic function in the testis\u2014Insights from the adjudin model. Reprod. Toxicol. 89, 54\u201366 (2019).","journal-title":"Reprod. Toxicol."},{"key":"12819_CR22","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1016\/j.ceb.2017.10.006","volume":"51","author":"A Fulford","year":"2018","unstructured":"Fulford, A., Tapon, N. & Ribeiro, P. S. Upstairs, downstairs: Spatial regulation of Hippo signalling. Curr. Opin. Cell Biol. 51, 22\u201332 (2018).","journal-title":"Curr. Opin. Cell Biol."},{"key":"12819_CR23","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1016\/j.theriogenology.2018.12.029","volume":"126","author":"GM Zhang","year":"2019","unstructured":"Zhang, G. M. et al. Expression of Hippo signaling pathway components in Hu sheep male reproductive tract and spermatozoa. Theriogenology 126, 239\u2013248 (2019).","journal-title":"Theriogenology"},{"key":"12819_CR24","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1016\/j.mce.2017.04.017","volume":"450","author":"SS Sharma","year":"2017","unstructured":"Sharma, S. S. & Majumdar, S. S. Transcriptional co-activator YAP regulates cAMP signaling in Sertoli cells. Mol. Cell. Endocrinol. 450, 64\u201373 (2017).","journal-title":"Mol. Cell. Endocrinol."},{"key":"12819_CR25","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1016\/j.repbio.2019.01.003","volume":"19","author":"SS Sharma","year":"2019","unstructured":"Sharma, S. S., Vats, A. & Majumdar, S. Regulation of Hippo pathway components by FSH in testis. Reprod. Biol. 19, 61\u201366 (2019).","journal-title":"Reprod. Biol."},{"key":"12819_CR26","doi-asserted-by":"publisher","first-page":"1035","DOI":"10.1111\/acel.12631","volume":"16","author":"MG Jeong","year":"2017","unstructured":"Jeong, M. G. et al. Transcriptional coactivator with PDZ-binding motif is required to sustain testicular function on aging. Aging Cell. 16, 1035\u20131042 (2017).","journal-title":"Aging Cell."},{"key":"12819_CR27","doi-asserted-by":"publisher","first-page":"604","DOI":"10.1038\/s41419-021-03885-4","volume":"12","author":"X Chi","year":"2021","unstructured":"Chi, X. et al. Kindlin-2 in Sertoli cells is essential for testis development and male fertility in mice. Cell Death Dis. 12, 604 (2021).","journal-title":"Cell Death Dis."},{"key":"12819_CR28","doi-asserted-by":"publisher","first-page":"979","DOI":"10.1095\/biolreprod65.4.979","volume":"65","author":"B Selam","year":"2001","unstructured":"Selam, B., Kayisli, U. A., Mulayim, N. & Arici, A. Regulation of Fas ligand expression by estradiol and progesterone in human endometrium. Biol. Reprod. 65, 979\u2013985 (2001).","journal-title":"Biol. Reprod."},{"key":"12819_CR29","doi-asserted-by":"publisher","first-page":"e13660","DOI":"10.1111\/and.13660","volume":"52","author":"AH Hasani Fard","year":"2020","unstructured":"Hasani Fard, A. H., Kouchesfehani, M., Jalali, H. & H. & Investigation of cholestasis-related changes in characteristics of spermatogonial stem cells in testis tissue of male Wistar rats. Andrologia 52, e13660 (2020).","journal-title":"Andrologia"},{"key":"12819_CR30","doi-asserted-by":"publisher","first-page":"1685","DOI":"10.1007\/s43032-025-01793-y","volume":"32","author":"\u0130 \u0130nan\u00e7","year":"2025","unstructured":"\u0130nan\u00e7, \u0130., Bender, O., Atalay, A., K\u00f6se, S. K. & Erdemli, E. Alterations in the Hippo signaling pathway during adenogenesis impairment in postnatal mouse uterus. Reprod. Sci. 32, 1685\u20131698 (2025).","journal-title":"Reproductive Sci."},{"key":"12819_CR31","unstructured":"Alafnan, A. et al. Beta elemene induces cytotoxic effects in FLT3 ITD-mutated acute myeloid leukemia by modulating apoptosis. Eur. Rev. Med. Pharmacol. Sci. 27 (2023)."},{"key":"12819_CR32","doi-asserted-by":"publisher","first-page":"119671","DOI":"10.1016\/j.lfs.2021.119671","volume":"280","author":"D Yildirim","year":"2021","unstructured":"Yildirim, D. et al. Role of autophagy and evaluation the effects of MicroRNAs 214, 132, 34c and prorenin receptor in a rat model of focal segmental glomerulosclerosis. Life Sci. 280, 119671 (2021).","journal-title":"Life Sci."},{"key":"12819_CR33","doi-asserted-by":"publisher","first-page":"2200407","DOI":"10.1002\/ardp.202200407","volume":"356","author":"O Bender","year":"2023","unstructured":"Bender, O. et al. Discovery of oxindole-based FLT3 inhibitors as a promising therapeutic lead for acute myeloid leukemia carrying the oncogenic ITD mutation. Arch. Pharm. 356, 2200407 (2023).","journal-title":"Arch. Pharm."},{"key":"12819_CR34","doi-asserted-by":"publisher","first-page":"671","DOI":"10.1038\/nmeth.2089","volume":"9","author":"CA Schneider","year":"2012","unstructured":"Schneider, C. A., Rasband, W. S. & Eliceiri, K. W. NIH image to imageJ: 25 years of image analysis. Nat. Methods. 9, 671\u2013675 (2012).","journal-title":"Nat. Methods"},{"key":"12819_CR35","doi-asserted-by":"publisher","first-page":"124","DOI":"10.1038\/nrg2723","volume":"11","author":"MA Handel","year":"2010","unstructured":"Handel, M. A. & Schimenti, J. C. Genetics of mammalian meiosis: regulation, dynamics and impact on fertility. Nat. Rev. Genet. 11, 124\u2013136 (2010).","journal-title":"Nat. Rev. Genet."},{"key":"12819_CR36","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1146\/annurev.cellbio.24.110707.175355","volume":"24","author":"JM Oatley","year":"2008","unstructured":"Oatley, J. M. & Brinster, R. L. Regulation of spermatogonial stem cell self-renewal in mammals. Annu. Rev. Cell Dev. Biol. 24, 263\u2013286 (2008).","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"12819_CR37","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1152\/physrev.00025.2011","volume":"92","author":"JM Oatley","year":"2012","unstructured":"Oatley, J. M. & Brinster, R. L. The germline stem cell niche unit in mammalian testes. Physiol. Rev. 92, 577\u2013595 (2012).","journal-title":"Physiol. Rev."},{"key":"12819_CR38","doi-asserted-by":"publisher","first-page":"1653","DOI":"10.1098\/rstb.2009.0254","volume":"365","author":"DJ Wolgemuth","year":"2010","unstructured":"Wolgemuth, D. J. & Roberts, S. S. Regulating mitosis and meiosis in the male germ line: critical functions for cyclins. Philos. Trans. R. Soc. B Biol. Sci. 365, 1653\u20131662 (2010).","journal-title":"Philosophical Trans. Royal Soc. B: Biol. Sci."},{"key":"12819_CR39","doi-asserted-by":"crossref","unstructured":"Clekmont, Y. & Perey, B. Quantitative study of the cell population of the seminiferous tubules in immature rats. (1957).","DOI":"10.1002\/aja.1001000205"},{"key":"12819_CR40","doi-asserted-by":"publisher","first-page":"525","DOI":"10.1095\/biolreprod43.3.525","volume":"43","author":"RA Hess","year":"1990","unstructured":"Hess, R. A. Quantitative and qualitative characteristics of the stages and transitions in the cycle of the rat seminiferous epithelium: light microscopic observations of perfusion-fixed and plastic-embedded testes. Biol. Reprod. 43, 525\u2013542 (1990).","journal-title":"Biol. Reprod."},{"key":"12819_CR41","doi-asserted-by":"crossref","unstructured":"Hess, R. A. & De Franca, L. R. Spermatogenesis and cycle of the seminiferous epithelium. Mol. Mech. Spermatogen. 1\u201315 (2009).","DOI":"10.1007\/978-0-387-09597-4_1"},{"key":"12819_CR42","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1111\/andr.12165","volume":"4","author":"LR Fran\u00e7a","year":"2016","unstructured":"Fran\u00e7a, L. R., Hess, R. A., Dufour, J. M., Hofmann, M. C. & Griswold, M. D. The Sertoli cell: one hundred Fifty years of beauty and plasticity. Andrology 4, 189\u2013212 (2016).","journal-title":"Andrology"},{"key":"12819_CR43","doi-asserted-by":"publisher","first-page":"1520","DOI":"10.1210\/en.2013-1850","volume":"155","author":"MW Li","year":"2014","unstructured":"Li, M. W., Cheng, C. Y. & Mruk, D. D. Sertolin mediates blood-testis barrier restructuring. Endocrinology 155, 1520\u20131531 (2014).","journal-title":"Endocrinology"},{"key":"12819_CR44","doi-asserted-by":"publisher","first-page":"223","DOI":"10.1016\/B978-0-12-385859-7.00005-7","volume":"286","author":"PP Lie","year":"2011","unstructured":"Lie, P. P., Cheng, C. Y. & Mruk, D. D. The biology of the desmosome-like junction: a versatile anchoring junction and signal transducer in the seminiferous epithelium. Int. Rev. Cell. Mol. Biol. 286, 223\u2013269 (2011).","journal-title":"Int. Rev. Cell. Mol. Biology"},{"key":"12819_CR45","doi-asserted-by":"publisher","first-page":"162","DOI":"10.1095\/biolreprod.108.070623","volume":"80","author":"JNS Young","year":"2009","unstructured":"Young, J. N. S., Guttman, J. A., Vaid, K. S. & Vogl, A. W. Tubulobulbar complexes are intercellular podosome-like structures that internalize intact intercellular junctions during epithelial remodeling events in the rat testis. Biol. Reprod. 80, 162\u2013174 (2009).","journal-title":"Biol. Reprod."},{"key":"12819_CR46","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1002\/ar.1091940204","volume":"194","author":"LD Russell","year":"1979","unstructured":"Russell, L. D. Further observations on tubulobulbar complexes formed by late spermatids and Sertoli cells in the rat testis. Anat. Rec. 194, 213\u2013232 (1979).","journal-title":"Anat. Rec."},{"key":"12819_CR47","doi-asserted-by":"publisher","first-page":"2555","DOI":"10.1096\/fj.06-070573","volume":"23","author":"PP Lie","year":"2009","unstructured":"Lie, P. P., Mruk, D. D., Lee, W. M. & Cheng, C. Y. Epidermal growth factor receptor pathway substrate 8 (Eps8) is a novel regulator of cell adhesion and the blood-testis barrier integrity in the seminiferous epithelium. FASEB J. 23, 2555 (2009).","journal-title":"FASEB J."},{"key":"12819_CR48","doi-asserted-by":"publisher","first-page":"188341","DOI":"10.1016\/j.bbcan.2020.188341","volume":"1873","author":"F Reggiani","year":"2020","unstructured":"Reggiani, F., Gobbi, G., Ciarrocchi, A., Ambrosetti, D. C. & Sancisi, V. Multiple roles and context-specific mechanisms underlying YAP and TAZ-mediated resistance to anti-cancer therapy. Biochim. Biophys. Acta (BBA) Rev. Cancer. 1873, 188341 (2020).","journal-title":"Biochim. Et Biophys. Acta (BBA)-Reviews Cancer"},{"key":"12819_CR49","doi-asserted-by":"publisher","first-page":"781","DOI":"10.1101\/gad.315127.118","volume":"32","author":"J Cai","year":"2018","unstructured":"Cai, J. et al. A RhoA\u2013YAP\u2013c-Myc signaling axis promotes the development of polycystic kidney disease. Genes Dev. 32, 781\u2013793 (2018).","journal-title":"Genes Dev."},{"key":"12819_CR50","doi-asserted-by":"publisher","first-page":"454","DOI":"10.1161\/CIRCRESAHA.114.302810","volume":"114","author":"SN Chen","year":"2014","unstructured":"Chen, S. N. et al. The Hippo pathway is activated and is a causal mechanism for adipogenesis in arrhythmogenic cardiomyopathy. Circul. Res. 114, 454\u2013468 (2014).","journal-title":"Circul. Res."},{"key":"12819_CR51","doi-asserted-by":"publisher","first-page":"17474","DOI":"10.1073\/pnas.1312830110","volume":"110","author":"K Kawamura","year":"2013","unstructured":"Kawamura, K. et al. Hippo signaling disruption and Akt stimulation of ovarian follicles for infertility treatment. Proc. Natl. Acad. Sci. 110, 17474\u201317479 (2013).","journal-title":"Proc. Natl. Acad. Sci."},{"key":"12819_CR52","doi-asserted-by":"publisher","first-page":"162","DOI":"10.1093\/biolre\/iox057","volume":"97","author":"A Levasseur","year":"2017","unstructured":"Levasseur, A., Paquet, M., Boerboom, D. & Boyer, A. Yes-associated protein and WW-containing transcription regulator 1 regulate the expression of sex-determining genes in Sertoli cells, but their inactivation does not cause sex reversal. Biol. Reprod. 97, 162\u2013175 (2017).","journal-title":"Biol. Reprod."},{"key":"12819_CR53","doi-asserted-by":"publisher","first-page":"dev175950","DOI":"10.1242\/dev.175950","volume":"146","author":"VA Ruthig","year":"2019","unstructured":"Ruthig, V. A. et al. The RNA-binding protein DND1 acts sequentially as a negative regulator of pluripotency and a positive regulator of epigenetic modifiers required for germ cell reprogramming. Development 146, dev175950 (2019).","journal-title":"Development"},{"key":"12819_CR54","doi-asserted-by":"publisher","first-page":"bqab238","DOI":"10.1210\/endocr\/bqab238","volume":"163","author":"A Lalonde-Larue","year":"2022","unstructured":"Lalonde-Larue, A. et al. The Hippo pathway effectors YAP and TAZ regulate LH release by pituitary gonadotrope cells in mice. Endocrinology 163, bqab238 (2022).","journal-title":"Endocrinology"},{"key":"12819_CR55","doi-asserted-by":"publisher","first-page":"904","DOI":"10.1007\/s13238-013-3084-z","volume":"4","author":"Y Matsui","year":"2013","unstructured":"Matsui, Y. & Lai, Z. C. Mutual regulation between Hippo signaling and actin cytoskeleton. Protein Cell. 4, 904\u2013910 (2013).","journal-title":"Protein Cell."},{"key":"12819_CR56","doi-asserted-by":"publisher","first-page":"e73763","DOI":"10.1371\/journal.pone.0073763","volume":"8","author":"P Reddy","year":"2013","unstructured":"Reddy, P., Deguchi, M., Cheng, Y. & Hsueh, A. J. Actin cytoskeleton regulates Hippo signaling. PloS One. 8, e73763 (2013).","journal-title":"PloS One"},{"key":"12819_CR57","doi-asserted-by":"publisher","first-page":"151","DOI":"10.5483\/BMBRep.2018.51.3.012","volume":"51","author":"J Seo","year":"2018","unstructured":"Seo, J. & Kim, J. Regulation of Hippo signaling by actin remodeling. BMB Rep. 51, 151 (2018).","journal-title":"BMB Rep."},{"key":"12819_CR58","unstructured":"Bu, T. et al. Seminars Cell. & Dev. Biology, 40\u201352 (Elsevier)."},{"key":"12819_CR59","doi-asserted-by":"publisher","first-page":"449","DOI":"10.1095\/biolreprod65.2.449","volume":"65","author":"CY Cheng","year":"2001","unstructured":"Cheng, C. Y. et al. Two new male contraceptives exert their effects by depleting germ cells prematurely from the testis. Biol. Reprod. 65, 449\u2013461 (2001).","journal-title":"Biol. Reprod."},{"key":"12819_CR60","doi-asserted-by":"publisher","first-page":"747","DOI":"10.1210\/er.2003-0022","volume":"25","author":"DD Mruk","year":"2004","unstructured":"Mruk, D. D. & Cheng, C. Y. Sertoli-Sertoli and Sertoli-germ cell interactions and their significance in germ cell movement in the seminiferous epithelium during spermatogenesis. Endocr. Rev. 25, 747\u2013806 (2004).","journal-title":"Endocr. Rev."},{"key":"12819_CR61","doi-asserted-by":"publisher","first-page":"439","DOI":"10.1016\/j.tem.2004.09.009","volume":"15","author":"DD Mruk","year":"2004","unstructured":"Mruk, D. D. & Cheng, C. Y. Cell\u2013cell interactions at the ectoplasmic specialization in the testis. Trends Endocrinol. Metab. 15, 439\u2013447 (2004).","journal-title":"Trends Endocrinol. Metabolism"},{"key":"12819_CR62","doi-asserted-by":"publisher","first-page":"790","DOI":"10.2164\/jandrol.106.000422","volume":"27","author":"KM Wolski","year":"2006","unstructured":"Wolski, K. M., Mruk, D. D. & Cameron, D. F. The Sertoli-Spermatid junctional complex adhesion strength is affected in vitro by adjudin. J. Androl. 27, 790\u2013794 (2006).","journal-title":"J. Androl."},{"key":"12819_CR63","doi-asserted-by":"publisher","first-page":"36","DOI":"10.1002\/bies.20513","volume":"29","author":"HH Yan","year":"2007","unstructured":"Yan, H. H., Mruk, D. D., Lee, W. M. & Cheng, C. Y. Ectoplasmic specialization: a friend or a foe of spermatogenesis? Bioessays 29, 36\u201348 (2007).","journal-title":"Bioessays"},{"key":"12819_CR64","doi-asserted-by":"publisher","first-page":"4724","DOI":"10.1210\/en.2008-1779","volume":"150","author":"IA Kopera","year":"2009","unstructured":"Kopera, I. A., Su, L., Bilinska, B., Cheng, C. Y. & Mruk, D. D. An in vivo study on adjudin and blood-testis barrier dynamics. Endocrinology 150, 4724\u20134733 (2009).","journal-title":"Endocrinology"},{"key":"12819_CR65","doi-asserted-by":"publisher","first-page":"130544","DOI":"10.1016\/j.jhazmat.2022.130544","volume":"445","author":"T Zhao","year":"2023","unstructured":"Zhao, T. et al. Prenatal and postnatal exposure to polystyrene microplastics induces testis developmental disorder and affects male fertility in mice. J. Hazard. Mater. 445, 130544 (2023).","journal-title":"J. Hazard. Mater."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-12819-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-12819-5","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-12819-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,9,8]],"date-time":"2025-09-08T12:49:31Z","timestamp":1757335771000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-12819-5"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8,2]]},"references-count":65,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["12819"],"URL":"https:\/\/doi.org\/10.1038\/s41598-025-12819-5","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,8,2]]},"assertion":[{"value":"18 March 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"21 July 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 August 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"28204"}}