{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T02:59:01Z","timestamp":1790218741179,"version":"4.1.0"},"reference-count":156,"publisher":"MDPI AG","issue":"7","license":[{"start":{"date-parts":[[2025,7,16]],"date-time":"2025-07-16T00:00:00Z","timestamp":1752624000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"USDA-NIFA Evans Allen project","award":["ARX01-6012"],"award-info":[{"award-number":["ARX01-6012"]}]}],"content-domain":{"domain":["www.mdpi.com"],"crossmark-restriction":true},"short-container-title":["Life"],"abstract":"<jats:p>DNA methylation is a well-studied epigenetic modification that regulates gene expression, maintains genome integrity, and influences cell fate. It is strictly regulated by a group of enzymes known as DNA methyltransferases (DNMTs). Most DNA methylation occurs at cytosines within symmetrical CpG dinucleotide base pairs, often located at gene promoters or other regulatory elements. Thus, methylation of a promoter CpG island leads to stable transcriptional repression of the associated gene. Nonetheless, abnormal gene expression caused by alterations in DNA methylation has been linked to infertility in both males and females, as well as to reproductive potential and improper post-fertilization embryo development. Recent epigenetic advancements have highlighted the significant association between epigenetic modification and reproductive health outcomes, garnering considerable attention. In this review, we explore significant advancements in understanding DNA methylation, emphasizing its establishment, maintenance, and functions in male and female reproductive sex cells. We also shed light on the recent discoveries on the influence of environmental exposures, nutrition, infection, stress, and lifestyle choices on DNA methylation. Finally, we discuss the latest insights and future directions concerning the diverse functions of DNA methylation in reproductive outcomes.<\/jats:p>","DOI":"10.3390\/life15071109","type":"journal-article","created":{"date-parts":[[2025,7,16]],"date-time":"2025-07-16T10:46:05Z","timestamp":1752662765000},"page":"1109","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":11,"title":["DNA Methylation: A Key Regulator in Male and Female Reproductive Outcomes"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6611-126X","authenticated-orcid":false,"given":"Adedeji","family":"Adetunji","sequence":"first","affiliation":[{"name":"Department of Agriculture, University of Arkansas at Pine Bluff, Pine Bluff, AR 71601, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Henrietta","family":"Owusu","sequence":"additional","affiliation":[{"name":"Department of Agriculture, University of Arkansas at Pine Bluff, Pine Bluff, AR 71601, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0513-6009","authenticated-orcid":false,"given":"Esiosa","family":"Adewale","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Louisville, Louisville, KY 40292, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6253-1558","authenticated-orcid":false,"given":"Precious","family":"Adesina","sequence":"additional","affiliation":[{"name":"Graduate School of Integrated Sciences for Life, Hiroshima University, 1-4-4 Kagamiyama, Higashihiroshima 739-8528, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christian","family":"Xedzro","sequence":"additional","affiliation":[{"name":"Laboratory of Food Microbiology and Hygiene, Hiroshima University, Higashihiroshima 739-8528, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tolulope","family":"Saliu","sequence":"additional","affiliation":[{"name":"Department of Physiology, College of Medicine, University of Kentucky, Lexington, KY 40536, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3131-6692","authenticated-orcid":false,"given":"Shahidul","family":"Islam","sequence":"additional","affiliation":[{"name":"Department of Agriculture, University of Arkansas at Pine Bluff, Pine Bluff, AR 71601, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhendong","family":"Zhu","sequence":"additional","affiliation":[{"name":"College of Animal Science and Technology, Qingdao Agricultural University, Qingdao 266109, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Olanrewaju","family":"Morenikeji","sequence":"additional","affiliation":[{"name":"Department of Biological and Health Sciences, University of Pittsburgh, Bradford, PA 16701, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,7,16]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"590","DOI":"10.1038\/s41580-019-0159-6","article-title":"The diverse roles of DNA methylation in mammalian development and disease","volume":"20","author":"Greenberg","year":"2019","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"e322","DOI":"10.1038\/emm.2017.10","article-title":"DNA methylation: An epigenetic mark of cellular memory","volume":"49","author":"Kim","year":"2017","journal-title":"Exp. Mol. Med."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"570","DOI":"10.1093\/biolre\/ioab085","article-title":"Epigenetic transgenerational inheritance, gametogenesis and germline development","volume":"105","author":"Nilsson","year":"2021","journal-title":"Biol. Reprod."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"2578","DOI":"10.1038\/s12276-024-01359-z","article-title":"Epigenetic reprogramming in mouse and human primordial germ cells","volume":"56","author":"Lee","year":"2024","journal-title":"Exp. Mol. Med."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"e13586","DOI":"10.1111\/and.13586","article-title":"Role of genetics and epigenetics in male infertility","volume":"53","author":"Gunes","year":"2021","journal-title":"Andrologia"},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Shacfe, G., Turko, R., Syed, H.H., Masoud, I., Tahmaz, Y., Samhan, L.M., Alkattan, K., Shafqat, A., and Yaqinuddin, A. (2023). A DNA Methylation Perspective on Infertility. Genes, 14.","DOI":"10.3390\/genes14122132"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1038\/s41588-022-01258-x","article-title":"Dynamics of DNA hydroxymethylation and methylation during mouse embryonic and germline development","volume":"55","author":"Yan","year":"2023","journal-title":"Nat. Genet."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"738","DOI":"10.1093\/nar\/gks1117","article-title":"Accumulation and loss of asymmetric non-CpG methylation during male germ-cell development","volume":"41","author":"Ichiyanagi","year":"2013","journal-title":"Nucleic Acids Res."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1569","DOI":"10.1016\/j.stem.2023.09.010","article-title":"Epigenetic regulation of early human embryo development","volume":"30","author":"Wilkinson","year":"2023","journal-title":"Cell Stem Cell"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1080\/15592294.2019.1695341","article-title":"The dynamics of DNA methylation during epigenetic reprogramming of primordial germ cells in medaka (Oryzias latipes)","volume":"15","author":"Wang","year":"2020","journal-title":"Epigenetics"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1111\/j.1365-2605.2007.00796.x","article-title":"Epigenetic transgenerational toxicology and germ cell disease","volume":"30","author":"Skinner","year":"2007","journal-title":"Int. J. Androl."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Tibben, B.M., and Rothbart, S.B. (2024). Mechanisms of DNA Methylation Regulatory Function and Crosstalk with Histone Lysine Methylation. J. Mol. Biol., 436.","DOI":"10.1016\/j.jmb.2023.168394"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"908","DOI":"10.1038\/nature06397","article-title":"The SRA protein Np95 mediates epigenetic inheritance by recruiting Dnmt1 to methylated DNA","volume":"450","author":"Sharif","year":"2007","journal-title":"Nature"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"eadu8116","DOI":"10.1126\/sciadv.adu8116","article-title":"Histone modification-driven structural remodeling unleashes DNMT3B in DNA methylation","volume":"11","author":"Cho","year":"2025","journal-title":"Sci. Adv."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"980","DOI":"10.1038\/s41422-020-0359-9","article-title":"Kinetics and mechanisms of mitotic inheritance of DNA methylation and their roles in aging-associated methylome deterioration","volume":"30","author":"Ming","year":"2020","journal-title":"Cell Res."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1038\/s41422-018-0125-4","article-title":"Integrative single-cell analysis of transcriptome, DNA methylome and chromatin accessibility in mouse oocytes","volume":"29","author":"Gu","year":"2019","journal-title":"Cell Res."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"542","DOI":"10.3892\/ol.2023.14129","article-title":"Mechanistic insights into UHRF1-mediated DNA methylation by structure-based functional clarification of UHRF1 domains","volume":"26","author":"Song","year":"2023","journal-title":"Oncol. Lett."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1155","DOI":"10.1038\/nsmb.2391","article-title":"Association of UHRF1 with methylated H3K9 directs the maintenance of DNA methylation","volume":"19","author":"Rothbart","year":"2012","journal-title":"Nat. Struct. Mol. Biol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"144166","DOI":"10.1016\/j.gene.2019.144166","article-title":"Epigenetic inheritance and intergenerational effects in mollusks","volume":"729","author":"Fallet","year":"2020","journal-title":"Gene"},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Nava-Rivera, L.E., Betancourt-Mart\u00ednez, N.D., Lozoya-Mart\u00ednez, R., Carranza-Rosales, P., Guzm\u00e1n-Delgado, N.E., Carranza-Torres, I.E., Delgado-Aguirre, H., Zambrano-Ort\u00edz, J.O., and Mor\u00e1n-Mart\u00ednez, J. (2021). Transgenerational effects in DNA methylation, genotoxicity and reproductive phenotype by chronic arsenic exposure. Sci. Rep., 11.","DOI":"10.1038\/s41598-021-87677-y"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1038\/npp.2012.112","article-title":"DNA methylation and its basic function","volume":"38","author":"Moore","year":"2013","journal-title":"Neuropsychopharmacology"},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Wang, Y., Riaz, F., Wang, W., Pu, J., Liang, Y., Wu, Z., Pan, S., Song, J., Yang, L., and Zhang, Y. (2024). Functional significance of DNA methylation: Epigenetic insights into Sj\u00f6gren\u2019s syndrome. Front. Immunol., 15.","DOI":"10.3389\/fimmu.2024.1289492"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"490","DOI":"10.1038\/nature10716","article-title":"DNA-binding factors shape the mouse methylome at distal regulatory regions","volume":"480","author":"Stadler","year":"2011","journal-title":"Nature"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"eaaj2239","DOI":"10.1126\/science.aaj2239","article-title":"Impact of cytosine methylation on DNA binding specificities of human transcription factors","volume":"356","author":"Yin","year":"2017","journal-title":"Science"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"145998","DOI":"10.1016\/j.scitotenv.2021.145998","article-title":"Epigenetic regulation of gene expression in response to environmental exposures: From bench to model","volume":"776","author":"Dong","year":"2021","journal-title":"Sci. Total Environ."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Marasca, F., Bodega, B., and Orlando, V. (2018). How Polycomb-Mediated Cell Memory Deals with a Changing Environment: Variations in PcG complexes and proteins assortment convey plasticity to epigenetic regulation as a response to environment. Bioessays, 40.","DOI":"10.1002\/bies.201700137"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1101\/gr.108498.110","article-title":"LSH and G9a\/GLP complex are required for developmentally programmed DNA methylation","volume":"21","author":"Myant","year":"2011","journal-title":"Genome Res."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"e70085","DOI":"10.1111\/tpj.70085","article-title":"INCREASED DNA METHYLATION 3 forms a potential chromatin remodelling complex with HAIRPLUS to regulate DNA methylation and trichome development in tomato","volume":"121","author":"Gyula","year":"2025","journal-title":"Plant J."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"5626","DOI":"10.1073\/pnas.1017000108","article-title":"Lsh, chromatin remodeling family member, modulates genome-wide cytosine methylation patterns at nonrepeat sequences","volume":"108","author":"Tao","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"2145","DOI":"10.1161\/CIRCULATIONAHA.110.956839","article-title":"Epigenetic modifications: Basic mechanisms and role in cardiovascular disease","volume":"123","author":"Handy","year":"2011","journal-title":"Circulation"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"e21099","DOI":"10.1016\/j.heliyon.2023.e21099","article-title":"Impact of DNA methylation of the human mesoderm-specific transcript (MEST) on male infertility","volume":"9","author":"Amjadian","year":"2023","journal-title":"Heliyon"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"2211","DOI":"10.1007\/s10815-020-01839-x","article-title":"Profiling the DNA methylation patterns of imprinted genes in abnormal semen samples by next-generation bisulfite sequencing","volume":"37","author":"He","year":"2020","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1038\/cr.2016.128","article-title":"DNA methylation and chromatin accessibility profiling of mouse and human fetal germ cells","volume":"27","author":"Guo","year":"2017","journal-title":"Cell Res."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"606","DOI":"10.1038\/nature13544","article-title":"The DNA methylation landscape of human early embryos","volume":"511","author":"Guo","year":"2014","journal-title":"Nature"},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Zeng, Y., and Chen, T. (2019). DNA Methylation Reprogramming during Mammalian Development. Genes, 10.","DOI":"10.3390\/genes10040257"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"944","DOI":"10.1016\/j.celrep.2015.01.021","article-title":"The Dnmt3L ADD Domain Controls Cytosine Methylation Establishment during Spermatogenesis","volume":"10","author":"Georgios","year":"2015","journal-title":"Cell Rep."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Ross, S.E., Hesselson, D., and Bogdanovic, O. (2021). Developmental Accumulation of Gene Body and Transposon Non-CpG Methylation in the Zebrafish Brain. Front. Cell Dev. Biol., 9.","DOI":"10.3389\/fcell.2021.643603"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1038\/nature13551","article-title":"Abnormalities in human pluripotent cells due to reprogramming mechanisms","volume":"511","author":"Ma","year":"2014","journal-title":"Nature"},{"key":"ref_39","first-page":"433","article-title":"Mouse oocyte methylomes at base resolution reveal genome-wide accumulation of non-CpG methylation and role of DNA methyltransferases","volume":"3","author":"Shirane","year":"2013","journal-title":"Cell Rep."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"811","DOI":"10.1242\/dev.061416","article-title":"Dynamic stage-specific changes in imprinted differentially methylated regions during early mammalian development and prevalence of non-CpG methylation in oocytes","volume":"138","author":"Tomizawa","year":"2011","journal-title":"Development"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"691","DOI":"10.1042\/EBC20190043","article-title":"The role and mechanisms of DNA methylation in the oocyte","volume":"63","author":"Kelsey","year":"2019","journal-title":"Essays Biochem."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1146\/annurev-genom-090413-025437","article-title":"Non-CG Methylation in the Human Genome","volume":"16","author":"He","year":"2015","journal-title":"Annu. Rev. Genomics Hum. Genet."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1895","DOI":"10.1242\/dev.070771","article-title":"Tet family proteins and 5-hydroxymethylcytosine in development and disease","volume":"139","author":"Tan","year":"2012","journal-title":"Development"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"e323","DOI":"10.1038\/emm.2017.5","article-title":"TET family dioxygenases and DNA demethylation in stem cells and cancers","volume":"49","author":"An","year":"2017","journal-title":"Exp. Mol. Med."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1458","DOI":"10.1038\/s41556-024-01475-y","article-title":"Distinct dynamics of parental 5-hydroxymethylcytosine during human preimplantation development regulate early lineage gene expression","volume":"26","author":"Liang","year":"2024","journal-title":"Nat. Cell Biol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"979","DOI":"10.1016\/j.cell.2014.04.017","article-title":"Programming and inheritance of parental DNA methylomes in mammals","volume":"157","author":"Wang","year":"2014","journal-title":"Cell"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"335","DOI":"10.2217\/epi-2023-0104","article-title":"The link between 5-hydroxymethylcytosine and DNA demethylation in early embryos","volume":"15","author":"Pfeifer","year":"2023","journal-title":"Epigenomics"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1016\/j.fertnstert.2011.12.036","article-title":"Epigenetics of the male gamete","volume":"97","author":"Carrell","year":"2012","journal-title":"Fertil. Steril."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"392","DOI":"10.1038\/nature25964","article-title":"Epigenetic reprogramming enables the transition from primordial germ cell to gonocyte","volume":"555","author":"Hill","year":"2018","journal-title":"Nature"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"2558","DOI":"10.1093\/humrep\/der192","article-title":"Genome-wide analysis identifies changes in histone retention and epigenetic modifications at developmental and imprinted gene loci in the sperm of infertile men","volume":"26","author":"Hammoud","year":"2011","journal-title":"Hum. Reprod."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"811","DOI":"10.1038\/ng.864","article-title":"Dynamic CpG island methylation landscape in oocytes and preimplantation embryos","volume":"43","author":"Smallwood","year":"2011","journal-title":"Nat. Genet."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1399","DOI":"10.2217\/epi-2016-0056","article-title":"Establishment and functions of DNA methylation in the germline","volume":"8","author":"Stewart","year":"2016","journal-title":"Epigenomics"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"732","DOI":"10.1016\/j.fertnstert.2020.09.009","article-title":"Identification of a unique epigenetic profile in women with diminished ovarian reserve","volume":"115","author":"Olsen","year":"2021","journal-title":"Fertil. Steril."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1777","DOI":"10.1016\/j.celrep.2013.04.032","article-title":"Bivalent chromatin marks developmental regulatory genes in the mouse embryonic germline in vivo","volume":"3","author":"Sachs","year":"2013","journal-title":"Cell Rep."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Seisenberger, S., Peat, J.R., Hore, T.A., Santos, F., Dean, W., and Reik, W. (2013). Reprogramming DNA methylation in the mammalian life cycle: Building and breaking epigenetic barriers. Philos. Trans. R. Soc. B Biol. Sci., 368.","DOI":"10.1098\/rstb.2011.0330"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1038\/nature10960","article-title":"A unique regulatory phase of DNA methylation in the early mammalian embryo","volume":"484","author":"Smith","year":"2012","journal-title":"Nature"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"909","DOI":"10.1016\/j.molcel.2012.07.010","article-title":"Protection against de novo methylation is instrumental in maintaining parent-of-origin methylation inherited from the gametes","volume":"47","author":"Proudhon","year":"2012","journal-title":"Mol. Cell"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1038\/s41467-024-55441-1","article-title":"DNA methylation modulates nucleosome retention in sperm and H3K4 methylation deposition in early mouse embryos","volume":"16","author":"Fanourgakis","year":"2025","journal-title":"Nat. Commun."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"812","DOI":"10.1101\/gad.234294.113","article-title":"DNA methylation dynamics during epigenetic reprogramming in the germline and preimplantation embryos","volume":"28","author":"Messerschmidt","year":"2014","journal-title":"Genes Dev."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"676","DOI":"10.1016\/j.tig.2022.03.010","article-title":"DNA methylation: A historical perspective","volume":"38","author":"Mattei","year":"2022","journal-title":"Trends Genet."},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Rotondo, J.C., Lanzillotti, C., Mazziotta, C., Tognon, M., and Martini, F. (2021). Epigenetics of Male Infertility: The Role of DNA Methylation. Front. Cell Dev. Biol., 9.","DOI":"10.3389\/fcell.2021.689624"},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Yamazaki, W., Badescu, D., Tan, S.L., Ragoussis, J., and Taketo, T. (2021). Effects of the Sex Chromosome Complement, XX, XO, or XY, on the Transcriptome and Development of Mouse Oocytes During Follicular Growth. Front. Genet., 12.","DOI":"10.3389\/fgene.2021.792604"},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Mani, S., Srivastava, V., Shandilya, C., Kaushik, A., and Singh, K.K. (2024). Mitochondria: The epigenetic regulators of ovarian aging and longevity. Front. Endocrinol., 15.","DOI":"10.3389\/fendo.2024.1424826"},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Safti\u0107 Martinovi\u0107, L., Mladeni\u0107, T., Lovri\u0107, D., Ostoji\u0107, S., and Devi\u0107 Pavli\u0107, S. (2024). Decoding the Epigenetics of Infertility: Mechanisms, Environmental Influences, and Therapeutic Strategies. Epigenomes, 8.","DOI":"10.3390\/epigenomes8030034"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"e13942","DOI":"10.1111\/and.13942","article-title":"Methylation patterns of methylenetetrahydrofolate reductase gene promoter in infertile males","volume":"53","author":"Kulac","year":"2021","journal-title":"Andrologia"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1093\/biolre\/ioac087","article-title":"Epigenetic regulation of cell fate transition: Learning from early embryo development and somatic cell reprogramming","volume":"107","author":"Chen","year":"2022","journal-title":"Biol. Reprod."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"175","DOI":"10.2217\/epi-2023-0298","article-title":"DNA methylation as a window into female reproductive aging","volume":"16","author":"Knight","year":"2024","journal-title":"Epigenomics"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1007\/s13237-021-00367-y","article-title":"DNA methylation and regulation of gene expression: Guardian of our health","volume":"64","author":"Dhar","year":"2021","journal-title":"Nucleus"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"R9","DOI":"10.1530\/REP-18-0009","article-title":"Epigenetic reprogramming during spermatogenesis and male factor infertility","volume":"156","author":"McSwiggin","year":"2018","journal-title":"Reproduction"},{"key":"ref_70","first-page":"4447","article-title":"Developmental features of DNA methylation in CpG islands of human gametes and preimplantation embryos","volume":"17","author":"Huang","year":"2019","journal-title":"Exp. Ther. Med."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Tang, Y., Gan, H., Wang, B., Wang, X., Li, M., Yang, Q., Geng, M., Zhu, P., Shao, S., and Tao, F. (2023). Mediating effects of DNA methylation in the association between sleep quality and infertility among women of childbearing age. BMC Public Health, 23.","DOI":"10.1186\/s12889-023-16681-w"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"R27","DOI":"10.1530\/REP-18-0449","article-title":"DNA methylation in the pathogenesis of polycystic ovary syndrome","volume":"158","year":"2019","journal-title":"Reproduction"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"e13402","DOI":"10.1111\/and.13402","article-title":"Chromosomal and Y-chromosome microdeletion analysis in 1300 infertile males and the fertility outcome of patients with AZFc microdeletions","volume":"51","author":"Abur","year":"2019","journal-title":"Andrologia"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"602","DOI":"10.1111\/andr.12742","article-title":"Increased DNA methylation in the spermatogenesis-associated (SPATA) genes correlates with infertility","volume":"8","author":"Sujit","year":"2020","journal-title":"Andrology"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"1726","DOI":"10.2174\/13816128113199990517","article-title":"DNA methylation, an epigenetic mode of gene expression regulation in reproductive science","volume":"20","author":"Calicchio","year":"2014","journal-title":"Curr. Pharm. Des."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"912","DOI":"10.1016\/j.bbrc.2019.04.100","article-title":"Absence of mitochondrial DNA methylation in mouse oocyte maturation, aging and early embryo development","volume":"513","author":"Fan","year":"2019","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Jenkins, T.G., Aston, K.I., Cairns, B., Smith, A., and Carrell, D.T. (2018). Paternal germ line aging: DNA methylation age prediction from human sperm. BMC Genom., 19.","DOI":"10.1186\/s12864-018-5153-4"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"e13349","DOI":"10.1111\/acel.13349","article-title":"Epigenetic clock and methylation study of oocytes from a bovine model of reproductive aging","volume":"20","author":"Kordowitzki","year":"2021","journal-title":"Aging Cell"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"1965","DOI":"10.1093\/humrep\/dez149","article-title":"Reproduction, DNA methylation and biological age","volume":"34","author":"Kresovich","year":"2019","journal-title":"Hum. Reprod."},{"key":"ref_80","doi-asserted-by":"crossref","unstructured":"Li Piani, L., Vigano, P., and Somigliana, E. (2023). Epigenetic clocks and female fertility timeline: A new approach to an old issue?. Front. Cell Dev. Biol., 11.","DOI":"10.3389\/fcell.2023.1121231"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1093\/humupd\/dmac033","article-title":"Age-associated epigenetic changes in mammalian sperm: Implications for offspring health and development","volume":"29","author":"Ashapkin","year":"2023","journal-title":"Hum. Reprod. Update"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"841","DOI":"10.1093\/humupd\/dmaa025","article-title":"DNA methylation in human sperm: A systematic review","volume":"26","author":"Danson","year":"2020","journal-title":"Hum. Reprod. Update"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"192","DOI":"10.1186\/s13148-020-00988-1","article-title":"High-resolution analyses of human sperm dynamic methylome reveal thousands of novel age-related epigenetic alterations","volume":"12","author":"Cao","year":"2020","journal-title":"Clin. Epigenet."},{"key":"ref_84","first-page":"1581","article-title":"Sperm epigenetic clock associates with pregnancy outcomes in the general population","volume":"37","author":"Pilsner","year":"2022","journal-title":"Hum. Reprod."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"1408","DOI":"10.1016\/j.fertnstert.2006.04.024","article-title":"Sperm protamine 1\/protamine 2 ratios are related to in vitro fertilization pregnancy rates and predictive of fertilization ability","volume":"86","author":"Aoki","year":"2006","journal-title":"Fertil. Steril."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"282","DOI":"10.1007\/BF02765830","article-title":"Human protamines and male infertility","volume":"14","author":"Khara","year":"1997","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Sharma, R., Agarwal, A., Rohra, V.K., Assidi, M., Abu-Elmagd, M., and Turki, R.F. (2015). Effects of increased paternal age on sperm quality, reproductive outcome and associated epigenetic risks to offspring. Reprod. Biol. Endocrinol., 13.","DOI":"10.1186\/s12958-015-0028-x"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"662","DOI":"10.1016\/j.tig.2022.03.006","article-title":"Transgenerational epigenetic impacts of parental infection on offspring health and disease susceptibility","volume":"38","author":"Elizabeth","year":"2022","journal-title":"Trends Genet."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"1830","DOI":"10.37506\/ijfmt.v15i3.15581","article-title":"Molecular Association of Sexually Transmitted Infections and Methylation Pattern of SNRPN Gene Promoter among Males with Abnormal Semen","volume":"15","author":"Smith","year":"2021","journal-title":"Indian J. Forensic Med. Toxicol."},{"key":"ref_90","doi-asserted-by":"crossref","unstructured":"Jasmin, P., Swarupa, M., Neha, M., Aman, T., and Negi, V.D. (2023). Pregnancy, infection, and epigenetic regulation: A complex scenario. Biochim. Biophys. Acta (BBA)-Mol. Basis Dis., 1869.","DOI":"10.1016\/j.bbadis.2023.166768"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"750","DOI":"10.1080\/15592294.2016.1221569","article-title":"Epigenome-wide differential DNA methylation between HIV-infected and uninfected individuals","volume":"11","author":"Pradhan","year":"2016","journal-title":"Epigenetics"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"9753","DOI":"10.1128\/JVI.75.20.9753-9761.2001","article-title":"Infection of Lymphoid Cells by Integration-Defective Human Immunodeficiency Virus Type 1 Increases De Novo Methylation","volume":"75","author":"Fang","year":"2001","journal-title":"J. Virol."},{"key":"ref_93","doi-asserted-by":"crossref","unstructured":"Arumugam, T., Ramphal, U., Adimulam, T., Chinniah, R., and Ramsuran, V. (2021). Deciphering DNA Methylation in HIV Infection. Front. Immunol., 12.","DOI":"10.3389\/fimmu.2021.795121"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"S128","DOI":"10.1093\/infdis\/jiab088","article-title":"Is There a Hidden Burden of Disease as a Result of Epigenetic Epithelial-to-Mesenchymal Transition Following Chlamydia trachomatis Genital Tract Infection?","volume":"224","author":"Horner","year":"2021","journal-title":"J. Infect. Dis."},{"key":"ref_95","doi-asserted-by":"crossref","unstructured":"Igietseme, J.U., Omosun, Y., Nagy, T., Stuchlik, O., Reed, M.S., He, Q., Partin, J., Joseph, K., Ellerson, D., and George, Z. (2018). Molecular Pathogenesis of Chlamydia Disease Complications: Epithelial-Mesenchymal Transition and Fibrosis. Infect. Immun., 86.","DOI":"10.1128\/IAI.00585-17"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"1194","DOI":"10.1038\/s41467-019-09144-7","article-title":"Chronic Chlamydia infection in human organoids increases stemness and promotes age-dependent CpG methylation","volume":"10","author":"Kessler","year":"2019","journal-title":"Nat. Commun."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"610","DOI":"10.1038\/s41564-019-0659-3","article-title":"The microbiota programs DNA methylation to control intestinal homeostasis and inflammation","volume":"5","author":"Ansari","year":"2020","journal-title":"Nat. Microbiol."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1097\/COH.0b013e3283504941","article-title":"Quality and quantity: Mucosal CD4+ T cells and HIV susceptibility","volume":"7","author":"McKinnon","year":"2012","journal-title":"Curr. Opin. HIV AIDS"},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"e25346","DOI":"10.1002\/jia2.25346","article-title":"Mechanisms of sexually transmitted infection-induced inflammation in women: Implications for HIV risk","volume":"22","author":"Mwatelah","year":"2019","journal-title":"J. Int. AIDS Soc."},{"key":"ref_100","doi-asserted-by":"crossref","unstructured":"Qin, W., Scicluna, B.P., and van der Poll, T. (2021). The Role of Host Cell DNA Methylation in the Immune Response to Bacterial Infection. Front. Immunol., 12.","DOI":"10.3389\/fimmu.2021.696280"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"641","DOI":"10.1165\/rcmb.2014-0327OC","article-title":"Regulatory T cell DNA methyltransferase inhibition accelerates resolution of lung inflammation","volume":"52","author":"Singer","year":"2015","journal-title":"Am. J. Respir. Cell Mol. Biol."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"148329","DOI":"10.1016\/j.gene.2024.148329","article-title":"Epigenetic disruptions in the offspring hypothalamus in response to maternal infection","volume":"910","author":"Samah","year":"2024","journal-title":"Gene"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"e13400","DOI":"10.1111\/aji.13400","article-title":"DNA damage contributes to transcriptional and immunological dysregulation of testicular cells during Chlamydia infection","volume":"86","author":"Bryan","year":"2021","journal-title":"Am. J. Reprod. Immunol."},{"key":"ref_104","first-page":"1196","article-title":"Curcumin as an indirect methylation inhibitor modulates the effects of Toxoplasma gondii on genes involved in male fertility","volume":"19","author":"Saki","year":"2020","journal-title":"EXCLI J."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1530\/REP-15-0212","article-title":"Arginine methyltransferases mediate an epigenetic ovarian response to endometriosis","volume":"150","author":"Baumann","year":"2015","journal-title":"Reproduction"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"103193","DOI":"10.1016\/j.jri.2020.103193","article-title":"Impact of lipopolysaccharide administration on luteinizing hormone\/choriogonadotropin receptor (Lhcgr) expression in mouse ovaries","volume":"142","author":"Adedeji","year":"2020","journal-title":"J. Reprod. Immunol."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1016\/j.fertnstert.2016.05.006","article-title":"Assisted reproductive technology alters deoxyribonucleic acid methylation profiles in bloodspots of newborn infants","volume":"106","author":"Estill","year":"2016","journal-title":"Fertil. Steril."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"2897","DOI":"10.1007\/s11033-012-2305-5","article-title":"Oct4 and the small molecule inhibitor, SC1, regulates Tet2 expression in mouse embryonic stem cells","volume":"40","author":"Wu","year":"2013","journal-title":"Mol. Biol. Rep."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1007\/s00281-020-00793-1","article-title":"Infection-induced epigenetic changes and their impact on the pathogenesis of diseases","volume":"42","author":"Fischer","year":"2020","journal-title":"Semin. Immunopathol."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"e13595","DOI":"10.1111\/and.13595","article-title":"Do lifestyle practices impede male fertility?","volume":"53","author":"Leisegang","year":"2021","journal-title":"Andrologia"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"19415","DOI":"10.18632\/oncotarget.24767","article-title":"Anabolic androgenic steroids and carcinogenicity focusing on Leydig cell: A literature review","volume":"9","author":"Salerno","year":"2018","journal-title":"Oncotarget"},{"key":"ref_112","first-page":"93","article-title":"The effect of nandrolone decanoate on the body, testis and epididymis weight and semen parameters in adult male rats","volume":"32","author":"Mesbah","year":"2007","journal-title":"Iran. J. Med. Sci."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"122","DOI":"10.1111\/j.1600-0463.2005.apm1130205.x","article-title":"High doses of nandrolone decanoate reduce volume of testis and length of seminiferous tubules in rats","volume":"113","author":"Noorafshan","year":"2005","journal-title":"Apmis"},{"key":"ref_114","first-page":"165","article-title":"Nandrolone decanoate administration to male rats induces oxidative stress, seminiferous tubules abnormalities, and sperm DNA fragmentation","volume":"3","author":"Tahtamouni","year":"2010","journal-title":"Jordan J. Biol. Sci."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"961","DOI":"10.1093\/humupd\/dmaa022","article-title":"The effect of paternal exposure to immunosuppressive drugs on sexual function, reproductive hormones, fertility, pregnancy and offspring outcomes: A systematic review","volume":"26","author":"Dolhain","year":"2020","journal-title":"Hum. Reprod. Update"},{"key":"ref_116","doi-asserted-by":"crossref","unstructured":"Gorczyca, G., Wartalski, K., Wiater, J., Samiec, M., Tabarowski, Z., and Duda, M. (2021). Anabolic Steroids-Driven Regulation of Porcine Ovarian Putative Stem Cells Favors the Onset of Their Neoplastic Transformation. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms222111800"},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1016\/j.cyto.2016.07.006","article-title":"Sex steroid-induced DNA methylation changes and inflammation response in prostate cancer","volume":"86","author":"Songping","year":"2016","journal-title":"Cytokine"},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"E952","DOI":"10.1152\/ajpendo.00451.2005","article-title":"DHT and testosterone, but not DHEA or E2, differentially modulate IGF-I, IGFBP-2, and IGFBP-3 in human prostatic stromal cells","volume":"290","author":"Le","year":"2006","journal-title":"Am. J. Physiol.-Endocrinol. Metab."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1530\/JOE-15-0506","article-title":"Supra-physiological dose of testosterone induces pathological cardiac hypertrophy","volume":"229","author":"Pirompol","year":"2016","journal-title":"J. Endocrinol."},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1016\/j.fertnstert.2018.10.013","article-title":"Pathogenesis of endometriosis: The genetic\/epigenetic theory","volume":"111","author":"Koninckx","year":"2019","journal-title":"Fertil. Steril."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"59","DOI":"10.5653\/cerm.2016.43.2.59","article-title":"Epigenetics: A key paradigm in reproductive health","volume":"43","author":"Bunkar","year":"2016","journal-title":"Clin. Exp. Reprod. Med."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"74","DOI":"10.1016\/j.tox.2011.07.011","article-title":"Hypermethylation of estrogen receptor promoter region in adult testis of rats exposed neonatally to bisphenol A","volume":"289","author":"Doshi","year":"2011","journal-title":"Toxicology"},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"105384","DOI":"10.1016\/j.envint.2019.105384","article-title":"Life cycle exposure to propiconazole reduces fecundity by disrupting the steroidogenic pathway and altering DNA methylation in zebrafish (Danio rerio)","volume":"135","author":"Miaomiao","year":"2020","journal-title":"Environ. Int."},{"key":"ref_124","doi-asserted-by":"crossref","unstructured":"Zhou, Y., Gu, B., Brichant, G., Singh, J.P., Yang, H., Chang, H., Zhao, Y., Cheng, C., Liu, Z.-W., and Alderman, M.H. (2022). The steroid hormone estriol (E3) regulates epigenetic programming of fetal mouse brain and reproductive tract. BMC Biol., 20.","DOI":"10.1186\/s12915-022-01293-4"},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"1013","DOI":"10.1016\/j.jmb.2004.05.052","article-title":"A HOXA10 estrogen response element (ERE) is differentially regulated by 17 beta-estradiol and diethylstilbestrol (DES)","volume":"340","author":"Akbas","year":"2004","journal-title":"J. Mol. Biol."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"3376","DOI":"10.1210\/en.2009-0071","article-title":"Hypermethylation of homeobox A10 by in utero diethylstilbestrol exposure: An epigenetic mechanism for altered developmental programming","volume":"150","author":"Bromer","year":"2009","journal-title":"Endocrinology"},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"2273","DOI":"10.1096\/fj.09-140533","article-title":"Bisphenol-A exposure in utero leads to epigenetic alterations in the developmental programming of uterine estrogen response","volume":"24","author":"Bromer","year":"2010","journal-title":"FASEB J."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"1575","DOI":"10.1007\/s10815-015-0553-8","article-title":"Marijuana, phytocannabinoids, the endocannabinoid system, and male fertility","volume":"32","author":"Agarwal","year":"2015","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1038\/nrurol.2010.6","article-title":"The effect of obesity on sperm disorders and male infertility","volume":"7","author":"Cabler","year":"2010","journal-title":"Nat. Rev. Urol."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"105","DOI":"10.2174\/1874255601103010105","article-title":"Is there a link between exercise and male factor infertility","volume":"3","author":"Kashou","year":"2011","journal-title":"Open Reprod. Sci. J."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1186\/s13072-018-0178-0","article-title":"Alterations in sperm DNA methylation, non-coding RNA and histone retention associate with DDT-induced epigenetic transgenerational inheritance of disease","volume":"11","author":"Skinner","year":"2018","journal-title":"Epigenet. Chromatin"},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/j.trsl.2014.02.003","article-title":"Environmentally induced epigenetic transgenerational inheritance of disease susceptibility","volume":"165","author":"Nilsson","year":"2015","journal-title":"Transl. Res."},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1016\/j.aju.2017.12.004","article-title":"Lifestyle causes of male infertility","volume":"16","author":"Durairajanayagam","year":"2018","journal-title":"Arab J. Urol."},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1038\/nrurol.2015.112","article-title":"Effects of psychological stress on male fertility","volume":"12","author":"Nargund","year":"2015","journal-title":"Nat. Rev. Urol."},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1038\/s41421-021-00343-5","article-title":"Sperm epigenetic alterations contribute to inter- and transgenerational effects of paternal exposure to long-term psychological stress via evading offspring embryonic reprogramming","volume":"7","author":"Zheng","year":"2021","journal-title":"Cell Discov."},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"1299","DOI":"10.1007\/s10815-019-01483-0","article-title":"Epigenetically regulated imprinted gene expression associated with IVF and infertility: Possible influence of prenatal stress and depression","volume":"36","author":"Litzky","year":"2019","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"194","DOI":"10.1038\/s41398-018-0252-1","article-title":"Exposure to childhood abuse is associated with human sperm DNA methylation","volume":"8","author":"Roberts","year":"2018","journal-title":"Transl. Psychiatry"},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1016\/j.psyneuen.2018.08.014","article-title":"Discrimination exposure and DNA methylation of stress-related genes in Latina mothers","volume":"98","author":"Hudson","year":"2018","journal-title":"Psychoneuroendocrinology"},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1289\/ehp.1307047","article-title":"DNA Methylation in Oocytes and Liver of Female Mice and Their Offspring: Effects of High-Fat-Diet\u2013Induced Obesity","volume":"122","author":"Ge","year":"2014","journal-title":"Environ. Health Perspect."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"2889","DOI":"10.1038\/ncomms3889","article-title":"Low paternal dietary folate alters the mouse sperm epigenome and is associated with negative pregnancy outcomes","volume":"4","author":"Lambrot","year":"2013","journal-title":"Nat. Commun."},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"513","DOI":"10.1530\/REP-19-0549","article-title":"Maternal peri-conceptional undernourishment perturbs offspring sperm methylome","volume":"159","author":"Toschi","year":"2020","journal-title":"Reproduction"},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1016\/j.cmet.2015.11.004","article-title":"Obesity and Bariatric Surgery Drive Epigenetic Variation of Spermatozoa in Humans","volume":"23","author":"Donkin","year":"2016","journal-title":"Cell Metab."},{"key":"ref_143","doi-asserted-by":"crossref","unstructured":"Hou, Y.-J., Zhu, C.-C., Duan, X., Liu, H.-L., Wang, Q., and Sun, S.-C. (2016). Both diet and gene mutation induced obesity affect oocyte quality in mice. Sci. Rep., 6.","DOI":"10.1038\/srep18858"},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"410","DOI":"10.1016\/j.rbmo.2010.11.009","article-title":"Oocyte developmental competence and embryo development: Impact of lifestyle and environmental risk factors","volume":"22","author":"Alex","year":"2011","journal-title":"Reprod. Biomed. Online"},{"key":"ref_145","doi-asserted-by":"crossref","unstructured":"Sciorio, R., Tramontano, L., Adel, M., and Fleming, S. (2024). Decrease in Sperm Parameters in the 21st Century: Obesity, Lifestyle, or Environmental Factors? An Updated Narrative Review. J. Pers. Med., 14.","DOI":"10.3390\/jpm14020198"},{"key":"ref_146","doi-asserted-by":"crossref","unstructured":"Brie\u00f1o-Enr\u00edquez, M.A., Garc\u00eda-L\u00f3pez, J., C\u00e1rdenas, D.B., Guibert, S., Cleroux, E., D\u011bd, L., Hourcade Jde, D., P\u011bknicov\u00e1, J., Weber, M., and Del Mazo, J. (2015). Exposure to endocrine disruptor induces transgenerational epigenetic deregulation of microRNAs in primordial germ cells. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0124296"},{"key":"ref_147","doi-asserted-by":"crossref","unstructured":"Iqbal, K., Tran, D.A., Li, A.X., Warden, C., Bai, A.Y., Singh, P., Wu, X., Pfeifer, G.P., and Szab\u00f3, P.E. (2015). Deleterious effects of endocrine disruptors are corrected in the mammalian germline by epigenome reprogramming. Genome Biol., 16.","DOI":"10.1186\/s13059-015-0619-z"},{"key":"ref_148","doi-asserted-by":"crossref","unstructured":"Manikkam, M., Tracey, R., Guerrero-Bosagna, C., and Skinner, M.K. (2013). Plastics derived endocrine disruptors (BPA, DEHP and DBP) induce epigenetic transgenerational inheritance of obesity, reproductive disease and sperm epimutations. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0055387"},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"108295","DOI":"10.1016\/j.mrrev.2019.108295","article-title":"Role of oxidative stress, genome damage and DNA methylation as determinants of pathological conditions in the newborn: An overview from conception to early neonatal stage","volume":"783","author":"Roberto","year":"2020","journal-title":"Mutat. Res.\/Rev. Mutat. Res."},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1016\/j.coph.2014.10.010","article-title":"Transgenerational effects of obesogens and the obesity epidemic","volume":"19","author":"Blumberg","year":"2014","journal-title":"Curr. Opin. Pharmacol."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"296","DOI":"10.1289\/ehp.1103927","article-title":"In utero exposures, infant growth, and DNA methylation of repetitive elements and developmentally related genes in human placenta","volume":"120","author":"Houseman","year":"2012","journal-title":"Environ. Health Perspect."},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"539","DOI":"10.4161\/epi.5.6.12378","article-title":"Global DNA hypomethylation is associated with in utero exposure to cotinine and perfluorinated alkyl compounds","volume":"5","author":"Goldman","year":"2010","journal-title":"Epigenetics"},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"1473","DOI":"10.4161\/15592294.2014.971593","article-title":"Fetal lung and placental methylation is associated with in utero nicotine exposure","volume":"9","author":"Chhabra","year":"2014","journal-title":"Epigenetics"},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1186\/s13148-025-01815-1","article-title":"How do lifestyle and environmental factors influence the sperm epigenome? Effects on sperm fertilising ability, embryo development, and offspring health","volume":"17","author":"Akhatova","year":"2025","journal-title":"Clin. Epigenet."},{"key":"ref_155","doi-asserted-by":"crossref","unstructured":"Song, B., Chen, Y., Wang, C., Li, G., Wei, Z., He, X., and Cao, Y. (2022). Poor semen parameters are associated with abnormal methylation of imprinted genes in sperm DNA. Reprod. Biol. Endocrinol., 20.","DOI":"10.1186\/s12958-022-01028-8"},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"R13","DOI":"10.1530\/RAF-20-0046","article-title":"Effects of lifestyle factors on fertility: Practical recommendations for modification","volume":"2","author":"Emokpae","year":"2021","journal-title":"Reprod. Fertil."}],"updated-by":[{"DOI":"10.3390\/life15121898","type":"correction","label":"Correction","source":"publisher","updated":{"date-parts":[[2025,7,16]],"date-time":"2025-07-16T00:00:00Z","timestamp":1752624000000}}],"container-title":["Life"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2075-1729\/15\/7\/1109\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,12,12]],"date-time":"2025-12-12T08:47:31Z","timestamp":1765529251000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2075-1729\/15\/7\/1109"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,7,16]]},"references-count":156,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2025,7]]}},"alternative-id":["life15071109"],"URL":"https:\/\/doi.org\/10.3390\/life15071109","relation":{},"ISSN":["2075-1729"],"issn-type":[{"value":"2075-1729","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,7,16]]}}}