{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T23:36:41Z","timestamp":1790984201162,"version":"4.1.0"},"reference-count":88,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,4,1]],"date-time":"2026-04-01T00:00:00Z","timestamp":1775001600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2026,4,1]],"date-time":"2026-04-01T00:00:00Z","timestamp":1775001600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Mamm Genome"],"published-print":{"date-parts":[[2026,12]]},"DOI":"10.1007\/s00335-026-10217-4","type":"journal-article","created":{"date-parts":[[2026,4,1]],"date-time":"2026-04-01T10:36:20Z","timestamp":1775039780000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Unravelling male infertility through epigenetics: present understanding and future challenges"],"prefix":"10.1007","volume":"37","author":[{"given":"S. Amrita","family":"Malini","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Almitra Abhijit","family":"Dravid","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ritwika","family":"Saha","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0780-0492","authenticated-orcid":false,"given":"Abilash Valsala","family":"Gopalakrishnan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sandra","family":"Kannampuzha","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,4,1]]},"reference":[{"issue":"3","key":"10217_CR1","doi-asserted-by":"publisher","first-page":"325","DOI":"10.1097\/01.gco.0000193003.58158.4e","volume":"18","author":"A Agarwal","year":"2006","unstructured":"Agarwal A, Gupta S, Sikka S (2006) The role of free radicals and antioxidants in reproduction. Curr Opin Obstet Gynecol 18(3):325\u2013332","journal-title":"Curr Opin Obstet Gynecol"},{"key":"10217_CR2","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1007\/978-3-030-32300-4_6","volume-title":"Male infertility: contemporary clinical approaches, andrology, ART and antioxidants","author":"A Agarwal","year":"2020","unstructured":"Agarwal A, Sengupta P (2020) Oxidative stress and its association with male infertility. Male infertility: contemporary clinical approaches, andrology, ART and antioxidants. pp 57\u201368"},{"issue":"6","key":"10217_CR3","doi-asserted-by":"publisher","first-page":"727","DOI":"10.1016\/S1472-6483(10)60676-1","volume":"14","author":"RJ Aitken","year":"2007","unstructured":"Aitken RJ, De Iuliis GN (2007) Origins and consequences of DNA damage in male germ cells. Reprod Biomed Online 14(6):727\u2013733","journal-title":"Reprod Biomed Online"},{"issue":"5727","key":"10217_CR4","doi-asserted-by":"publisher","first-page":"1466","DOI":"10.1126\/science.1108190","volume":"308","author":"MD Anway","year":"2005","unstructured":"Anway MD et al (2005) Epigenetic transgenerational actions of endocrine disruptors and male fertility. Science 308(5727):1466\u20131469","journal-title":"Science"},{"issue":"10","key":"10217_CR5","doi-asserted-by":"publisher","first-page":"1711","DOI":"10.3390\/cells11101711","volume":"11","author":"MJC Assidi","year":"2022","unstructured":"Assidi MJC (2022) Infertility in men: advances towards a comprehensive and integrative strategy for precision theranostics. Cells 11(10):1711","journal-title":"Cells"},{"issue":"1","key":"10217_CR6","first-page":"e1","volume":"2","author":"F Azizi","year":"2018","unstructured":"Azizi F et al (2018) The genetic causes of male infertility in Iranian population; a systematic review. Men\u2019s Health J 2(1):e1\u2013e1","journal-title":"Men\u2019s Health J"},{"issue":"1","key":"10217_CR7","doi-asserted-by":"publisher","first-page":"5452","DOI":"10.1038\/s41598-022-09336-0","volume":"12","author":"D Beck","year":"2022","unstructured":"Beck D et al (2022) Environmental induced transgenerational inheritance impacts systems epigenetics in disease etiology. Sci Rep 12(1):5452","journal-title":"Sci Rep"},{"issue":"8","key":"10217_CR8","doi-asserted-by":"publisher","first-page":"7224","DOI":"10.3390\/ijms24087224","volume":"24","author":"R Cannarella","year":"2023","unstructured":"Cannarella R et al (2023) H19 sperm methylation in male infertility: a systematic review and meta-analysis. Int J Mol Sci 24(8):7224","journal-title":"Int J Mol Sci"},{"issue":"7","key":"10217_CR9","doi-asserted-by":"publisher","first-page":"3363","DOI":"10.1210\/en.2009-1340","volume":"151","author":"D Chan","year":"2010","unstructured":"Chan D et al (2010) Strain-specific defects in testicular development and sperm epigenetic patterns in 5,10-methylenetetrahydrofolate reductase-deficient mice. Endocrinology 151(7):3363\u20133373","journal-title":"Endocrinology"},{"issue":"12","key":"10217_CR10","doi-asserted-by":"publisher","first-page":"733","DOI":"10.1038\/nrg.2016.106","volume":"17","author":"Q Chen","year":"2016","unstructured":"Chen Q, Yan W, Duan E (2016) Epigenetic inheritance of acquired traits through sperm RNAs and sperm RNA modifications. Nat Rev Genet 17(12):733\u2013743","journal-title":"Nat Rev Genet"},{"issue":"7","key":"10217_CR11","first-page":"4142","volume":"45","author":"J Chen","year":"2017","unstructured":"Chen J et al (2017) MicroRNA-202 maintains spermatogonial stem cells by inhibiting cell cycle regulators and RNA binding proteins. Nucleic Acids Res 45(7):4142\u20134157","journal-title":"Nucleic Acids Res"},{"issue":"2","key":"10217_CR12","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1530\/REP-20-0415","volume":"161","author":"X Chen","year":"2021","unstructured":"Chen X et al (2021) Human sperm tsRNA as potential biomarker and therapy target for male fertility. Reproduction 161(2):111\u2013122","journal-title":"Reproduction"},{"issue":"3","key":"10217_CR13","first-page":"e0214275","volume":"14","author":"S Cheung","year":"2019","unstructured":"Cheung S et al (2019) Genetic epigenetic profiling infertile male 14(3):e0214275","journal-title":"Genetic epigenetic profiling infertile male"},{"key":"10217_CR14","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1007\/978-1-4939-7698-0_11","volume-title":"Sertoli Cells: Methods and Protocols","author":"L Cris\u00f3stomo","year":"2018","unstructured":"Cris\u00f3stomo L (2018) Molecular mechanisms and signaling pathways involved in the nutritional support of spermatogenesis by Sertoli cells. Sertoli Cells: Methods and Protocols. pp 129\u2013155"},{"issue":"4","key":"10217_CR15","doi-asserted-by":"publisher","first-page":"1973","DOI":"10.3892\/etm.2016.3569","volume":"12","author":"X Cui","year":"2016","unstructured":"Cui X et al (2016) DNA methylation in spermatogenesis and male infertility. Experimental therapeutic Med 12(4):1973\u20131979","journal-title":"Experimental therapeutic Med"},{"issue":"52","key":"10217_CR16","doi-asserted-by":"publisher","first-page":"44306","DOI":"10.1074\/jbc.M111.282756","volume":"286","author":"L Dai","year":"2011","unstructured":"Dai L et al (2011) Testis-specific miRNA-469 up-regulated in gonadotropin-regulated testicular RNA helicase (GRTH\/DDX25)-null mice silences transition protein 2 and protamine 2 messages at sites within coding region: implications of its role in germ cell development. J Biol Chem 286(52):44306\u201344318","journal-title":"J Biol Chem"},{"issue":"52","key":"10217_CR17","doi-asserted-by":"publisher","first-page":"44306","DOI":"10.1074\/jbc.M111.282756","volume":"286","author":"L Dai","year":"2011","unstructured":"Dai L et al (2011) Testis-specific miRNA-469 up-regulated in gonadotropin-regulated testicular RNA helicase (GRTH\/DDX25)-null mice silences transition protein 2 and protamine 2 messages at sites within coding region: implications of its role in germ cell development. Journal of Biological Chemistry 286(52):44306\u201344318","journal-title":"Journal of Biological Chemistry"},{"issue":"1","key":"10217_CR18","doi-asserted-by":"publisher","first-page":"e12881","DOI":"10.1111\/apha.12881","volume":"222","author":"J Denham","year":"2018","unstructured":"Denham J (2018) Exercise and epigenetic inheritance of disease risk. Acta Physiol 222(1):e12881","journal-title":"Acta Physiol"},{"issue":"1","key":"10217_CR19","doi-asserted-by":"publisher","first-page":"416","DOI":"10.1080\/19396368.2025.2548492","volume":"71","author":"O Dhikhirullahi","year":"2025","unstructured":"Dhikhirullahi O, Zhang Z (2025) Male infertility. Syst Biol Reprod Med 71(1):416\u2013438","journal-title":"Syst Biol Reprod Med"},{"issue":"2","key":"10217_CR20","doi-asserted-by":"publisher","first-page":"369","DOI":"10.1016\/j.cmet.2015.11.004","volume":"23","author":"I Donkin","year":"2016","unstructured":"Donkin I et al (2016) Obesity and Bariatric Surgery Drive Epigenetic Variation of Spermatozoa in Humans. Cell Metab 23(2):369\u2013378","journal-title":"Cell Metab"},{"issue":"8","key":"10217_CR21","first-page":"1776","volume":"28","author":"D Doroshow","year":"2017","unstructured":"Doroshow D, Eder J, LoRusso PJAoO (2017) BET inhibitors: novel epigenetic approach 28(8):1776\u20131787","journal-title":"BET inhibitors: novel epigenetic approach"},{"issue":"5","key":"10217_CR22","first-page":"1","volume":"3","author":"K Elango","year":"2022","unstructured":"Elango K et al (2022) Impact of sperm protamine on semen quality and fertility. J Reproductive Healthc Med 3(5):1","journal-title":"J Reproductive Healthc Med"},{"key":"10217_CR23","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1016\/j.toxlet.2017.10.004","volume":"282","author":"G Elmhiri","year":"2018","unstructured":"Elmhiri G et al (2018) DNA methylation and potential multigenerational epigenetic effects linked to uranium chronic low-dose exposure in gonads of males and females rats. Toxicol Lett 282:64\u201370","journal-title":"Toxicol Lett"},{"issue":"6422","key":"10217_CR24","doi-asserted-by":"publisher","first-page":"751","DOI":"10.1038\/362751a0","volume":"362","author":"AC Ferguson-Smith","year":"1993","unstructured":"Ferguson-Smith AC et al (1993) Parental-origin-specific epigenetic modification of the mouse H19 gene. Nature 362(6422):751\u2013755","journal-title":"Nature"},{"issue":"6489","key":"10217_CR25","first-page":"367","volume":"368","author":"P Filippakopoulos","year":"2020","unstructured":"Filippakopoulos P, Knapp SJS (2020) Next-generation epigenetic inhibitors 368(6489):367\u2013368","journal-title":"Next-generation epigenetic inhibitors"},{"key":"10217_CR26","unstructured":"Gomes VLdS (2022) Analysis of epigenetic regulators in sperm cells from patients with male infertility"},{"issue":"6","key":"10217_CR27","doi-asserted-by":"publisher","first-page":"733","DOI":"10.1002\/wrna.1252","volume":"5","author":"LT Gou","year":"2014","unstructured":"Gou LT, Dai P, Liu MF (2014) Small noncoding RNAs and male infertility. Wiley Interdiscip Rev RNA 5(6):733\u2013745","journal-title":"Wiley Interdiscip Rev RNA"},{"issue":"6","key":"10217_CR28","doi-asserted-by":"publisher","first-page":"1090","DOI":"10.1016\/j.cell.2017.04.034","volume":"169","author":"L-T Gou","year":"2017","unstructured":"Gou L-T et al (2017) Ubiquitination-deficient mutations in human Piwi cause male infertility by impairing histone-to-protamine exchange during spermiogenesis. Cell 169(6):1090\u20131104e13","journal-title":"Cell"},{"issue":"3","key":"10217_CR29","doi-asserted-by":"publisher","first-page":"349","DOI":"10.2164\/jandrol.04146","volume":"26","author":"E Greco","year":"2005","unstructured":"Greco E et al (2005) Reduction of the incidence of sperm DNA fragmentation by oral antioxidant treatment. J Androl 26(3):349\u2013353","journal-title":"J Androl"},{"issue":"10","key":"10217_CR30","doi-asserted-by":"publisher","first-page":"590","DOI":"10.1038\/s41580-019-0159-6","volume":"20","author":"MV Greenberg","year":"2019","unstructured":"Greenberg MV, Bourc\u2019his D (2019) The diverse roles of DNA methylation in mammalian development and disease. Nat Rev Mol Cell Biol 20(10):590\u2013607","journal-title":"Nat Rev Mol Cell Biol"},{"issue":"7254","key":"10217_CR31","doi-asserted-by":"publisher","first-page":"473","DOI":"10.1038\/nature08162","volume":"460","author":"SS Hammoud","year":"2009","unstructured":"Hammoud SS et al (2009) Distinctive chromatin in human sperm packages genes for embryo development. Nature 460(7254):473\u2013478","journal-title":"Nature"},{"issue":"44","key":"10217_CR32","doi-asserted-by":"publisher","first-page":"18674","DOI":"10.1073\/pnas.0906378106","volume":"106","author":"T Han","year":"2009","unstructured":"Han T et al (2009) 26G endo-siRNAs regulate spermatogenic and zygotic gene expression in Caenorhabditis elegans. Proceedings of the National Academy of Sciences 106(44):18674\u201318679","journal-title":"Proceedings of the National Academy of Sciences"},{"issue":"4","key":"10217_CR33","doi-asserted-by":"publisher","first-page":"363","DOI":"10.4103\/aja.aja_85_20","volume":"23","author":"BM Hanson","year":"2021","unstructured":"Hanson BM et al (2021) Round spermatid injection into human oocytes: a systematic review and meta-analysis. Asian J Androl 23(4):363\u2013369","journal-title":"Asian J Androl"},{"issue":"6785","key":"10217_CR34","doi-asserted-by":"publisher","first-page":"486","DOI":"10.1038\/35013106","volume":"405","author":"AT Hark","year":"2000","unstructured":"Hark AT et al (2000) CTCF mediates methylation-sensitive enhancer-blocking activity at the H19\/Igf2 locus. Nature 405(6785):486\u2013489","journal-title":"Nature"},{"issue":"6","key":"10217_CR35","doi-asserted-by":"publisher","first-page":"627","DOI":"10.1016\/j.rbmo.2012.09.005","volume":"25","author":"XJ He","year":"2012","unstructured":"He XJ et al (2012) PRM1 variant rs35576928 (Arg\u2009>\u2009Ser) is associated with defective spermatogenesis in the Chinese Han population. Reprod Biomed Online 25(6):627\u2013634","journal-title":"Reprod Biomed Online"},{"issue":"10","key":"10217_CR36","doi-asserted-by":"publisher","first-page":"2205","DOI":"10.1002\/stem.1474","volume":"31","author":"Z He","year":"2013","unstructured":"He Z et al (2013) MiRNA-20 and mirna-106a regulate spermatogonial stem cell renewal at the post-transcriptional level via targeting STAT3 and Ccnd1. Stem Cells 31(10):2205\u20132217","journal-title":"Stem Cells"},{"key":"10217_CR37","doi-asserted-by":"crossref","unstructured":"Heerboth S et al (2014) Use of epigenetic drugs in disease: an overview. 6:pGEGS12270","DOI":"10.4137\/GEG.S12270"},{"issue":"17","key":"10217_CR38","doi-asserted-by":"publisher","first-page":"3061","DOI":"10.1242\/dev.136721","volume":"143","author":"S Hilz","year":"2016","unstructured":"Hilz S et al (2016) The roles of microRNAs and siRNAs in mammalian spermatogenesis. Development 143(17):3061\u20133073","journal-title":"Development"},{"issue":"4","key":"10217_CR39","doi-asserted-by":"publisher","first-page":"899","DOI":"10.1002\/tox.23453","volume":"37","author":"Q Hu","year":"2022","unstructured":"Hu Q et al (2022) Uranium induces kidney cells apoptosis via reactive oxygen species generation, endoplasmic reticulum stress and inhibition of PI3K\/AKT\/mTOR signaling in culture. Environ Toxicol 37(4):899\u2013909","journal-title":"Environ Toxicol"},{"issue":"9","key":"10217_CR40","doi-asserted-by":"publisher","first-page":"e14172","DOI":"10.1111\/and.14172","volume":"53","author":"TM Hussein","year":"2021","unstructured":"Hussein TM et al (2021) Assessment of serum vitamin D level and seminal vitamin D receptor gene methylation in a sample of Egyptian men with idiopathic infertility. Andrologia 53(9):e14172","journal-title":"Andrologia"},{"issue":"1","key":"10217_CR41","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1095\/biolreprod.112.103747","volume":"88","author":"JM Huszar","year":"2013","unstructured":"Huszar JM, Payne CJ (2013) MicroRNA 146 (Mir146) modulates spermatogonial differentiation by retinoic acid in mice. Biol Reprod 88(1):1\u201310","journal-title":"Biol Reprod"},{"issue":"18","key":"10217_CR42","doi-asserted-by":"publisher","first-page":"1685","DOI":"10.1056\/NEJMoa1209026","volume":"368","author":"HL Janssen","year":"2013","unstructured":"Janssen HL et al (2013) Treatment of HCV infection by targeting microRNA. N Engl J Med 368(18):1685\u20131694","journal-title":"N Engl J Med"},{"key":"10217_CR43","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s12958-020-00660-6","volume":"18","author":"M Joshi","year":"2020","unstructured":"Joshi M, Rajender S (2020) Long non-coding RNAs (lncRNAs) in spermatogenesis and male infertility. Reprod Biol Endocrinol 18:1\u201318","journal-title":"Reprod Biol Endocrinol"},{"issue":"1","key":"10217_CR44","doi-asserted-by":"publisher","first-page":"106","DOI":"10.1186\/s12958-023-01155-w","volume":"21","author":"M Joshi","year":"2023","unstructured":"Joshi M et al (2023) Small RNAs, spermatogenesis, and male infertility: a decade of retrospect. Reproductive Biology and Endocrinology 21(1):106","journal-title":"Reproductive Biology and Endocrinology"},{"issue":"2","key":"10217_CR45","first-page":"113","volume":"8","author":"Y Khazaie","year":"2014","unstructured":"Khazaie Y, Esfahani MH (2014) MicroRNA and male infertility: a potential for diagnosis. Int J fertil & sterl 8(2):113","journal-title":"Int J fertil & sterl"},{"issue":"1","key":"10217_CR46","doi-asserted-by":"publisher","first-page":"17079","DOI":"10.1038\/s41598-024-67910-0","volume":"14","author":"H Kobayashi","year":"2024","unstructured":"Kobayashi H et al (2024) A new model for determining risk of male infertility from serum hormone levels, without semen analysis. Sci Rep 14(1):17079","journal-title":"Sci Rep"},{"issue":"12","key":"10217_CR47","doi-asserted-by":"publisher","first-page":"3401","DOI":"10.1093\/humrep\/der329","volume":"26","author":"SA Krawetz","year":"2011","unstructured":"Krawetz SA et al (2011) A survey of small RNAs in human sperm. Human Reproduction 26(12):3401\u20133412","journal-title":"Human Reproduction"},{"key":"10217_CR48","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1016\/j.trsl.2022.12.006","volume":"256","author":"H-M Li","year":"2023","unstructured":"Li H-M et al (2023) Promising novel biomarkers and therapy targets: the application of cell-free seminal nucleotides in male reproduction research. Transl Res 256:73\u201386","journal-title":"Transl Res"},{"issue":"3","key":"10217_CR49","doi-asserted-by":"publisher","first-page":"535","DOI":"10.3892\/mmr.2012.967","volume":"6","author":"T Liu","year":"2012","unstructured":"Liu T et al (2012) Microarray analysis of microRNA expression patterns in the semen of infertile men with semen abnormalities. Mol Med Rep 6(3):535\u2013542","journal-title":"Mol Med Rep"},{"key":"10217_CR50","doi-asserted-by":"crossref","unstructured":"Ly L, Chan D, Trasler JM (2015) Developmental windows of susceptibility for epigenetic inheritance through the male germline. in Seminars in cell & developmental biology. Elsevier","DOI":"10.1016\/j.semcdb.2015.07.006"},{"issue":"8","key":"10217_CR51","doi-asserted-by":"publisher","DOI":"10.3390\/antiox13081020","volume":"13","author":"Z Ma","year":"2024","unstructured":"Ma Z et al (2024) Ribonuclease inhibitor 1 (RNH1) regulates sperm tsRNA generation for paternal inheritance through interacting with angiogenin in the caput epididymis. Antioxidants 13(8):1020","journal-title":"Antioxidants"},{"issue":"1","key":"10217_CR52","doi-asserted-by":"publisher","first-page":"e55387","DOI":"10.1371\/journal.pone.0055387","volume":"8","author":"M Manikkam","year":"2013","unstructured":"Manikkam M et al (2013) Plastics derived endocrine disruptors (BPA, DEHP and DBP) induce epigenetic transgenerational inheritance of obesity, reproductive disease and sperm epimutations. PLoS ONE 8(1):e55387","journal-title":"PLoS ONE"},{"issue":"1","key":"10217_CR53","doi-asserted-by":"publisher","first-page":"44","DOI":"10.1093\/humupd\/dmr041","volume":"18","author":"SC McIver","year":"2012","unstructured":"McIver SC et al (2012) miRNA and mammalian male germ cells. Hum Reprod Update 18(1):44\u201359","journal-title":"Hum Reprod Update"},{"issue":"4","key":"10217_CR54","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0035553","volume":"7","author":"SC McIver","year":"2012","unstructured":"McIver SC et al (2012) A unique combination of male germ cell miRNAs coordinates gonocyte differentiation. PLoS ONE 7(4):e35553","journal-title":"PLoS ONE"},{"issue":"3","key":"10217_CR55","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1038\/nrg2521","volume":"10","author":"TR Mercer","year":"2009","unstructured":"Mercer TR, Dinger ME, Mattick JS (2009) Long non-coding RNAs: insights into functions. Nat Rev Genet 10(3):155\u2013159","journal-title":"Nat Rev Genet"},{"key":"10217_CR56","doi-asserted-by":"crossref","unstructured":"Minh N et al (2018) Histone Deacetylase Inhibitor Corrects Histone H3K9 Modification in Round Spermatid DNA at the 2-Cell Stage and Increases the Development of ROSI Embryos. in 6th International Conference on the Development of Biomedical Engineering in Vietnam (BME6) 6. Springer","DOI":"10.1007\/978-981-10-4361-1_149"},{"key":"10217_CR57","doi-asserted-by":"publisher","first-page":"124","DOI":"10.1016\/j.placenta.2020.10.020","volume":"103","author":"HL Morgan","year":"2021","unstructured":"Morgan HL et al (2021) Paternal low protein diet and the supplementation of methyl-donors impact fetal growth and placental development in mice. Placenta 103:124\u2013133","journal-title":"Placenta"},{"issue":"10","key":"10217_CR58","doi-asserted-by":"publisher","first-page":"e12766","DOI":"10.1111\/and.12766","volume":"49","author":"F Nasri","year":"2017","unstructured":"Nasri F et al (2017) Sperm DNA methylation of H19 imprinted gene and male infertility. Andrologia 49(10):e12766","journal-title":"Andrologia"},{"issue":"3","key":"10217_CR59","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1111\/joim.13096","volume":"288","author":"D N\u00e4tt","year":"2020","unstructured":"N\u00e4tt D, \u00d6st A (2020) Male reproductive health and intergenerational metabolic responses from a small RNA perspective. Journal of Internal Medicine 288(3):305\u2013320","journal-title":"Journal of Internal Medicine"},{"issue":"7","key":"10217_CR60","doi-asserted-by":"publisher","first-page":"1411","DOI":"10.1093\/humrep\/dew096","volume":"31","author":"K Ni","year":"2016","unstructured":"Ni K et al (2016) TET enzymes are successively expressed during human spermatogenesis and their expression level is pivotal for male fertility. Hum Reprod 31(7):1411\u20131424","journal-title":"Hum Reprod"},{"issue":"31","key":"10217_CR61","doi-asserted-by":"publisher","first-page":"12740","DOI":"10.1073\/pnas.1109987108","volume":"108","author":"Z Niu","year":"2011","unstructured":"Niu Z et al (2011) MicroRNA-21 regulates the self-renewal of mouse spermatogonial stem cells. Proc Natl Acad Sci 108(31):12740\u201312745","journal-title":"Proc Natl Acad Sci"},{"issue":"4","key":"10217_CR62","doi-asserted-by":"publisher","first-page":"417","DOI":"10.1093\/humupd\/dml009","volume":"12","author":"R Oliva","year":"2006","unstructured":"Oliva R (2006) Protamines and male infertility. Hum Reprod Update 12(4):417\u2013435","journal-title":"Hum Reprod Update"},{"issue":"6","key":"10217_CR64","doi-asserted-by":"publisher","first-page":"1352","DOI":"10.1016\/j.cell.2014.11.005","volume":"159","author":"A \u00d6st","year":"2014","unstructured":"\u00d6st A et al (2014) Paternal diet defines offspring chromatin state and intergenerational obesity. Cell 159(6):1352\u20131364","journal-title":"Cell"},{"issue":"7","key":"10217_CR65","doi-asserted-by":"publisher","first-page":"e41146","DOI":"10.1371\/journal.pone.0041146","volume":"7","author":"S Panneerdoss","year":"2012","unstructured":"Panneerdoss S et al (2012) Androgen-responsive microRNAs in mouse Sertoli cells. PLoS ONE 7(7):e41146","journal-title":"PLoS ONE"},{"key":"10217_CR66","doi-asserted-by":"publisher","first-page":"101463","DOI":"10.1016\/j.molmet.2022.101463","volume":"59","author":"AS Pepin","year":"2022","unstructured":"Pepin AS et al (2022) Sperm histone H3 lysine 4 tri-methylation serves as a metabolic sensor of paternal obesity and is associated with the inheritance of metabolic dysfunction. Mol Metab 59:101463","journal-title":"Mol Metab"},{"key":"10217_CR67","doi-asserted-by":"publisher","first-page":"101463","DOI":"10.1016\/j.molmet.2022.101463","volume":"59","author":"A-S Pepin","year":"2022","unstructured":"Pepin A-S et al (2022) Sperm histone H3 lysine 4 tri-methylation serves as a metabolic sensor of paternal obesity and is associated with the inheritance of metabolic dysfunction. Mol metabolism 59:101463","journal-title":"Mol metabolism"},{"issue":"4","key":"10217_CR68","doi-asserted-by":"publisher","first-page":"642","DOI":"10.1111\/j.1365-2605.2009.01000.x","volume":"33","author":"A Poplinski","year":"2010","unstructured":"Poplinski A et al (2010) Idiopathic male infertility is strongly associated with aberrant methylation of MEST and IGF2\/H19 ICR1. Int J Androl 33(4):642\u2013649","journal-title":"Int J Androl"},{"issue":"1","key":"10217_CR69","doi-asserted-by":"publisher","first-page":"8209","DOI":"10.1038\/s41467-023-43480-z","volume":"14","author":"D Qin","year":"2023","unstructured":"Qin D et al (2023) Phase-separated CCER1 coordinates the histone-to-protamine transition and male fertility. Nat Commun 14(1):8209","journal-title":"Nat Commun"},{"issue":"2","key":"10217_CR70","doi-asserted-by":"publisher","first-page":"518","DOI":"10.1095\/biolreprod.103.020487","volume":"70","author":"J Rivlin","year":"2004","unstructured":"Rivlin J et al (2004) Role of hydrogen peroxide in sperm capacitation and acrosome reaction. Biol Reprod 70(2):518\u2013522","journal-title":"Biol Reprod"},{"issue":"44","key":"10217_CR71","doi-asserted-by":"publisher","first-page":"13699","DOI":"10.1073\/pnas.1508347112","volume":"112","author":"AB Rodgers","year":"2015","unstructured":"Rodgers AB et al (2015) Transgenerational epigenetic programming via sperm microRNA recapitulates effects of paternal stress. Proc Natl Acad Sci U S A 112(44):13699\u201313704","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"3","key":"10217_CR72","doi-asserted-by":"publisher","first-page":"dvz013","DOI":"10.1093\/eep\/dvz013","volume":"5","author":"I Sadler-Riggleman","year":"2019","unstructured":"Sadler-Riggleman I et al (2019) Epigenetic transgenerational inheritance of testis pathology and Sertoli cell epimutations: generational origins of male infertility. Environ epigenetics 5(3):dvz013","journal-title":"Environ epigenetics"},{"issue":"1","key":"10217_CR73","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1016\/j.fertnstert.2019.03.020","volume":"112","author":"S Sarkar","year":"2019","unstructured":"Sarkar S et al (2019) Array-based DNA methylation profiling reveals peripheral blood differential methylation in male infertility. Fertil sterl 112(1):61\u201372","journal-title":"Fertil sterl"},{"key":"10217_CR74","unstructured":"da Silva Gomes VL (2021) Analysis of Epigenetic Regulators in Sperm Cells From Patients with Male Infertility. Universidade do Minho (Portugal)"},{"issue":"11","key":"10217_CR75","doi-asserted-by":"publisher","first-page":"677","DOI":"10.1111\/boc.201200014","volume":"104","author":"L Smorag","year":"2012","unstructured":"Smorag L et al (2012) MicroRNA signature in various cell types of mouse spermatogenesis: evidence for stage-specifically expressed miRNA\u2010221,\u2010203 and\u201034b\u20105p mediated spermatogenesis regulation. Biol Cell 104(11):677\u2013692","journal-title":"Biol Cell"},{"issue":"32","key":"10217_CR76","doi-asserted-by":"publisher","first-page":"13159","DOI":"10.1073\/pnas.1108567108","volume":"108","author":"R Song","year":"2011","unstructured":"Song R et al (2011) Male germ cells express abundant endogenous siRNAs. Proceedings of the National Academy of Sciences 108(32):13159\u201313164","journal-title":"Proceedings of the National Academy of Sciences"},{"issue":"6","key":"10217_CR77","doi-asserted-by":"publisher","first-page":"465","DOI":"10.1038\/nrg2341","volume":"9","author":"MM Suzuki","year":"2008","unstructured":"Suzuki MM, Bird A (2008) DNA methylation landscapes: provocative insights from epigenomics. Nat Rev Genet 9(6):465\u2013476","journal-title":"Nat Rev Genet"},{"issue":"1","key":"10217_CR78","doi-asserted-by":"publisher","first-page":"11219","DOI":"10.1038\/s41598-017-11636-9","volume":"7","author":"Q Tang","year":"2017","unstructured":"Tang Q et al (2017) Idiopathic male infertility and polymorphisms in the DNA methyltransferase genes involved in epigenetic marking. Sci Rep 7(1):11219","journal-title":"Sci Rep"},{"key":"10217_CR79","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s13148-018-0568-y","volume":"10","author":"Q Tang","year":"2018","unstructured":"Tang Q et al (2018) Idiopathic male infertility is strongly associated with aberrant DNA methylation of imprinted loci in sperm: a case-control study. Clin epigenetics 10:1\u201310","journal-title":"Clin epigenetics"},{"issue":"1","key":"10217_CR80","doi-asserted-by":"publisher","first-page":"11219","DOI":"10.1038\/s41598-017-11636-9","volume":"7","author":"Q Tang","year":"2017","unstructured":"Tang Q et al (2017) Idiopathic male infertility and polymorphisms in the DNA methyltransferase genes involved in epigenetic marking. Scientific Reports (1):11219","journal-title":"Scientific Reports"},{"issue":"8","key":"10217_CR81","doi-asserted-by":"publisher","first-page":"1770","DOI":"10.3390\/ijerph15081770","volume":"15","author":"Y Tian","year":"2018","unstructured":"Tian Y et al (2018) Association of bisphenol A exposure with LINE-1 hydroxymethylation in human semen. Int J Environ Res Public Health 15(8):1770","journal-title":"Int J Environ Res Public Health"},{"key":"10217_CR82","doi-asserted-by":"publisher","first-page":"537","DOI":"10.1007\/s10815-009-9346-2","volume":"26","author":"O Tunc","year":"2009","unstructured":"Tunc O, Tremellen K (2009) Oxidative DNA damage impairs global sperm DNA methylation in infertile men. J Assist Reprod Genet 26:537\u2013544","journal-title":"J Assist Reprod Genet"},{"issue":"5","key":"10217_CR83","doi-asserted-by":"publisher","first-page":"446","DOI":"10.1242\/bio.2012844","volume":"1","author":"M De Vries","year":"2012","unstructured":"De Vries M et al (2012) Chromatin remodelling initiation during human spermiogenesis. Biology open 1(5):446\u2013457","journal-title":"Biology open"},{"key":"10217_CR84","doi-asserted-by":"publisher","first-page":"962","DOI":"10.3389\/fgene.2019.00962","volume":"10","author":"T Wang","year":"2019","unstructured":"Wang T et al (2019) Essential role of histone replacement and modifications in male fertility. Front Genet 10:962","journal-title":"Front Genet"},{"issue":"37","key":"10217_CR85","first-page":"5541","volume":"26","author":"W-S Xu","year":"2007","unstructured":"Xu W-S, Parmigiani RB, Marks PAJO (2007) Histone deacetylase inhibitors: Mol Mech action 26(37):5541\u20135552","journal-title":"Histone deacetylase inhibitors: Mol Mech action"},{"issue":"2","key":"10217_CR86","doi-asserted-by":"publisher","first-page":"212","DOI":"10.1242\/bio.201410959","volume":"4","author":"S Yuan","year":"2015","unstructured":"Yuan S et al (2015) mir-34b\/c and mir-449a\/b\/c are required for spermatogenesis, but not for the first cleavage division in mice. Biol Open 4(2):212\u2013223","journal-title":"Biol Open"},{"issue":"1","key":"10217_CR87","doi-asserted-by":"publisher","first-page":"17743","DOI":"10.1038\/srep17743","volume":"5","author":"R Zhou","year":"2015","unstructured":"Zhou R et al (2015) Mitochondria-related miR-151a-5p reduces cellular ATP production by targeting CYTB in asthenozoospermia. Sci Rep 5(1):17743","journal-title":"Sci Rep"},{"key":"10217_CR88","doi-asserted-by":"publisher","first-page":"105849","DOI":"10.1016\/j.aquatox.2021.105849","volume":"236","author":"L Zhu","year":"2021","unstructured":"Zhu L et al (2021) Chronic exposure to Bisphenol A resulted in alterations of reproductive functions via immune defense, oxidative damage and disruption DNA\/histone methylation in male rare minnow Gobiocypris rarus. Aquat Toxicol 236:105849","journal-title":"Aquat Toxicol"},{"issue":"9","key":"10217_CR89","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0107023","volume":"9","author":"C Zimmermann","year":"2014","unstructured":"Zimmermann C et al (2014) Germ cell-specific targeting of DICER or DGCR8 reveals a novel role for endo-siRNAs in the progression of mammalian spermatogenesis and male fertility. PLoS ONE 9(9):e107023","journal-title":"PLoS ONE"}],"container-title":["Mammalian Genome"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00335-026-10217-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00335-026-10217-4","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00335-026-10217-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,4,1]],"date-time":"2026-04-01T12:04:00Z","timestamp":1775045040000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00335-026-10217-4"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,1]]},"references-count":88,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2026,12]]}},"alternative-id":["10217"],"URL":"https:\/\/doi.org\/10.1007\/s00335-026-10217-4","relation":{},"ISSN":["0938-8990","1432-1777"],"issn-type":[{"value":"0938-8990","type":"print"},{"value":"1432-1777","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,4,1]]},"assertion":[{"value":"3 August 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"18 March 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"1 April 2026","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":"49"}}