{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T11:31:39Z","timestamp":1791977499059,"version":"4.4.0"},"reference-count":56,"publisher":"Wiley","license":[{"start":{"date-parts":[[2026,7,20]],"date-time":"2026-07-20T00:00:00Z","timestamp":1784505600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"},{"start":{"date-parts":[[2026,7,20]],"date-time":"2026-07-20T00:00:00Z","timestamp":1784505600000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/100000066","name":"National Institute of Environmental Health Sciences","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000066","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01 ES028298"],"award-info":[{"award-number":["R01 ES028298"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01 HD110462"],"award-info":[{"award-number":["R01 HD110462"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Andrology"],"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:sec>\n                    <jats:title>Background<\/jats:title>\n                    <jats:p>Sperm mitochondrial DNA copy number is an emerging marker of male fertility, with elevated mtDNAcn associated with poor semen quality across various populations. Sperm DNA methylation has also been previously demonstrated to be negatively correlated with semen parameters in IVF cohorts.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Objective<\/jats:title>\n                    <jats:p>To determine whether mtDNAcn is associated with nuclear DNA methylation across three population\u2010based and clinical fertility cohorts.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Design<\/jats:title>\n                    <jats:p>\n                      The association between sperm mtDNAcn and genome\u2010wide DNA methylation was investigated in 748 men from a general population cohort (LIFE;\n                      <jats:italic>n<\/jats:italic>\n                      = 379) and two infertility cohorts (SEEDS;\n                      <jats:italic>n<\/jats:italic>\n                      = 170 and EARTH;\n                      <jats:italic>n<\/jats:italic>\n                      = 199). Sperm DNA methylation and mtDNAcn were quantified by the Illumina EPIC (v1) array and triplex probe\u2010based digital PCR, respectively. Covariate\u2010adjusted linear regressions were conducted within each cohort and then meta\u2010analyzed using METAL. Gene ontology analyses explored the biological relevance of differentially methylated CpGs (DMCs) and regions (DMRs).\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>Using a Holm\u2013Bonferroni significance threshold, our analyses identified 1351, 224, and 258 DMCs and 2582, 1536, and 1284 DMRs in LIFE, SEEDS, and EARTH, respectively, indicating that higher sperm mtDNAcn was associated with DNA hypermethylation. In meta\u2010analysis, 576 DMCs were associated with mtDNAcn. Notably, the genes corresponding with DMCs and DMRs are known critical regulators of male fertility and were enriched in the regulation of male meiosis, germline protection, spermatogenesis pathways, and early embryonic development.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Discussion and Conclusion<\/jats:title>\n                    <jats:p>Our results, observed across three independent cohorts and supported by meta\u2010analysis, indicate a consistent association between the sperm mitochondrial genome and nuclear DNA methylation, suggesting that mtDNAcn and methylation patterns may be linked during spermatogenesis. Further research is warranted to elucidate the mechanisms underlying these associations and to evaluate their relevance for reproductive outcomes.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.1111\/andr.70320","type":"journal-article","created":{"date-parts":[[2026,7,20]],"date-time":"2026-07-20T07:25:26Z","timestamp":1784532326000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Sperm Mitochondrial DNA Copy Number and Nuclear DNA Methylation Across Clinical and Non\u2010Clinical Preconception Cohorts"],"prefix":"10.1111","author":[{"ORCID":"https:\/\/orcid.org\/0009-0000-9157-5771","authenticated-orcid":false,"given":"Savni","family":"Sawant","sequence":"first","affiliation":[{"name":"Department of Biochemistry Microbiology Immunology School of Medicine Wayne State University  Detroit Michigan USA"},{"name":"Department of Obstetrics and Gynecology C.S. Mott Centre for Human Growth and Development School of Medicine Wayne State University  Detroit Michigan USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9971-0023","authenticated-orcid":false,"given":"Karolina","family":"Nowak","sequence":"additional","affiliation":[{"name":"Department of Obstetrics and Gynecology C.S. Mott Centre for Human Growth and Development School of Medicine Wayne State University  Detroit Michigan USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amanda L.","family":"Paskavitz","sequence":"additional","affiliation":[{"name":"Department of Obstetrics and Gynecology C.S. Mott Centre for Human Growth and Development School of Medicine Wayne State University  Detroit Michigan USA"},{"name":"Center For Molecular Medicine and Genetics School of Medicine Wayne State University  Detroit Michigan USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Emily","family":"Houle","sequence":"additional","affiliation":[{"name":"Department of Obstetrics and Gynecology C.S. Mott Centre for Human Growth and Development School of Medicine Wayne State University  Detroit Michigan USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6892-9244","authenticated-orcid":false,"given":"Ricardo P.","family":"Bertolla","sequence":"additional","affiliation":[{"name":"Department of Obstetrics and Gynecology C.S. Mott Centre for Human Growth and Development School of Medicine Wayne State University  Detroit Michigan USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2370-3900","authenticated-orcid":false,"given":"Lidia","family":"Minguez\u2010Alarcon","sequence":"additional","affiliation":[{"name":"Department of Medicine Brigham and Women's Hospital and Harvard Medical School  Boston Massachusetts USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Russ","family":"Hauser","sequence":"additional","affiliation":[{"name":"Department of Environmental Health Harvard T. H. Chan School of Public Health  Boston Massachusetts USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4541-6267","authenticated-orcid":false,"given":"J. Richard","family":"Pilsner","sequence":"additional","affiliation":[{"name":"Department of Obstetrics and Gynecology C.S. Mott Centre for Human Growth and Development School of Medicine Wayne State University  Detroit Michigan USA"},{"name":"Institute of Environmental Health Sciences Wayne State University  Detroit Michigan USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,7,20]]},"reference":[{"key":"e_1_2_9_2_1","volume-title":"Infertility Prevalence Estimates, 1990\u20132021","author":"WHO","year":"2023"},{"issue":"10271","key":"e_1_2_9_3_1","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1016\/S0140-6736(20)32667-2","article-title":"Male Infertility","volume":"397","author":"Agarwal A.","year":"2021","journal-title":"Lancet"},{"issue":"2","key":"e_1_2_9_4_1","doi-asserted-by":"crossref","first-page":"1567","DOI":"10.3390\/cimb46020101","article-title":"Associations Between Sperm Epigenetic Age and Semen Parameters: An Evaluation of Clinical and Non\u2010Clinical Cohorts","volume":"46","author":"Sawant S.","year":"2024","journal-title":"Current Issues in Molecular Biology"},{"issue":"2","key":"e_1_2_9_5_1","doi-asserted-by":"crossref","first-page":"324","DOI":"10.1016\/j.eururo.2012.04.048","article-title":"European Association of Urology Guidelines on Male Infertility: The 2012 Update","volume":"62","author":"Jungwirth A.","year":"2012","journal-title":"European Urology"},{"issue":"19","key":"e_1_2_9_6_1","doi-asserted-by":"crossref","first-page":"1388","DOI":"10.1056\/NEJMoa003005","article-title":"Sperm Morphology, Motility, and Concentration in Fertile and Infertile Men","volume":"345","author":"Guzick D. S.","year":"2001","journal-title":"New England Journal of Medicine"},{"issue":"11","key":"e_1_2_9_7_1","doi-asserted-by":"crossref","first-page":"2619","DOI":"10.1093\/humrep\/deaa191","article-title":"Sperm Mitochondrial DNA Biomarkers and Couple Fecundity","volume":"35","author":"Rosati A. J.","year":"2020","journal-title":"Human Reproduction"},{"issue":"1","key":"e_1_2_9_8_1","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1093\/humrep\/dey330","article-title":"Associations of Sperm Mitochondrial DNA Copy Number and Deletion Rate With Fertilization and Embryo Development in a Clinical Setting","volume":"34","author":"Wu H.","year":"2019","journal-title":"Human Reproduction"},{"key":"e_1_2_9_9_1","doi-asserted-by":"crossref","first-page":"270","DOI":"10.1016\/j.xfre.2025.05.002","article-title":"A Machine Learning Approach Using Semen Parameters and Sperm Mitochondrial DNA Copy Number to Predict Couples\u2032 Fecundity","volume":"6","author":"Sawant S.","year":"2025","journal-title":"F&S Reports"},{"issue":"2","key":"e_1_2_9_10_1","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1093\/humrep\/15.suppl_2.86","article-title":"Regulation of Mitochondrial DNA Copy Number During Spermatogenesis","volume":"15","author":"Rantanen A.","year":"2000","journal-title":"Human Reproduction"},{"issue":"1","key":"e_1_2_9_11_1","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1080\/17501911.2025.2603883","article-title":"Mitochondrial DNA Copy Number Reduction via in Vitro TFAM Knockout Remodels the Nuclear Epigenome and Transcriptome","volume":"18","author":"Nguyen J.","year":"2026","journal-title":"Epigenomics"},{"issue":"6760","key":"e_1_2_9_12_1","doi-asserted-by":"crossref","first-page":"371","DOI":"10.1038\/46466","article-title":"Ubiquitin Tag for Sperm Mitochondria","volume":"402","author":"Sutovsky P.","year":"1999","journal-title":"Nature"},{"issue":"10","key":"e_1_2_9_13_1","doi-asserted-by":"crossref","first-page":"1632","DOI":"10.1038\/s41588-023-01505-9","article-title":"Molecular Basis for Maternal Inheritance of Human Mitochondrial DNA","volume":"55","author":"Lee W.","year":"2023","journal-title":"Nature Genetics"},{"issue":"1","key":"e_1_2_9_14_1","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1016\/j.rbmo.2018.10.004","article-title":"Sperm Mitochondrial DNA Measures and Semen Parameters Among Men Undergoing Fertility Treatment","volume":"38","author":"Wu H.","year":"2019","journal-title":"Reproductive Biomedicine Online"},{"issue":"3","key":"e_1_2_9_15_1","doi-asserted-by":"crossref","first-page":"550","DOI":"10.1093\/humrep\/deg096","article-title":"Increased Sperm Mitochondrial DNA Content in Male Infertility","volume":"18","author":"May\u2010Panloup P.","year":"2003","journal-title":"Human Reproduction"},{"issue":"13","key":"e_1_2_9_16_1","doi-asserted-by":"crossref","first-page":"2098","DOI":"10.1111\/1471-0528.17139","article-title":"Sperm Mitochondrial DNA Copy Number in Relation to Semen Quality: A Cross\u2010Sectional Study of 1164 Potential Sperm Donors","volume":"129","author":"Sun B.","year":"2022","journal-title":"BJOG"},{"issue":"9","key":"e_1_2_9_17_1","doi-asserted-by":"crossref","first-page":"1434","DOI":"10.1111\/1471-0528.17078","article-title":"Sperm Mitochondrial DNA Copy Numbers in Normal and Abnormal Semen Analysis: A Systematic Review and Meta\u2010Analysis","volume":"129","author":"Popova D.","year":"2022","journal-title":"BJOG"},{"key":"e_1_2_9_18_1","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1016\/j.mito.2020.07.006","article-title":"Associations of Mitochondrial DNA Copy Number and Deletion Rate With Early Pregnancy Loss","volume":"55","author":"Ye M.","year":"2020","journal-title":"Mitochondrion"},{"key":"e_1_2_9_19_1","doi-asserted-by":"crossref","DOI":"10.1016\/j.mce.2024.112393","article-title":"Status of Sperm Mitochondrial Functions and DNA Methylation in Infertile Men With Clinical Varicocele Before and After Treatment","volume":"595","author":"Arya D.","year":"2025","journal-title":"Molecular and Cellular Endocrinology"},{"issue":"8","key":"e_1_2_9_20_1","doi-asserted-by":"crossref","first-page":"759","DOI":"10.1007\/s10815-012-9785-z","article-title":"Influence of Microsurgical Varicocelectomy on Human Sperm Mitochondrial DNA Copy Number: A Pilot Study","volume":"29","author":"Gabriel M. S.","year":"2012","journal-title":"Journal of Assisted Reproduction and Genetics"},{"issue":"24","key":"e_1_2_9_21_1","doi-asserted-by":"crossref","first-page":"2141","DOI":"10.2217\/epi-2020-0253","article-title":"Association Between Sperm Mitochondarial DNA Copy Number and Nuclear DNA Methylation","volume":"12","author":"Oluwayiose O. A.","year":"2020","journal-title":"Epigenomics"},{"issue":"1575","key":"e_1_2_9_22_1","doi-asserted-by":"crossref","first-page":"2266","DOI":"10.1098\/rstb.2011.0042","article-title":"Epigenetic Reprogramming in the Germline: Towards the Ground State of the Epigenome","volume":"366","author":"Hajkova P.","year":"2011","journal-title":"Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences"},{"issue":"5","key":"e_1_2_9_23_1","doi-asserted-by":"crossref","first-page":"413","DOI":"10.1111\/j.1365-3016.2011.01205.x","article-title":"Designing Prospective Cohort Studies for Assessing Reproductive and Developmental Toxicity During Sensitive Windows of Human Reproduction and Development\u2013The LIFE Study","volume":"25","author":"Buck Louis G. M.","year":"2011","journal-title":"Paediatric and Perinatal Epidemiology"},{"issue":"2","key":"e_1_2_9_24_1","doi-asserted-by":"crossref","DOI":"10.1093\/hropen\/hoy001","article-title":"The Environment and Reproductive Health (EARTH) Study: A Prospective Preconception Cohort","volume":"2018","author":"Messerlian C.","year":"2018","journal-title":"Human Reproduction Open"},{"issue":"2","key":"e_1_2_9_25_1","doi-asserted-by":"crossref","first-page":"453","DOI":"10.1016\/j.fertnstert.2013.10.022","article-title":"Semen Quality and Time to Pregnancy: The Longitudinal Investigation of Fertility and the Environment Study","volume":"101","author":"Buck Louis G. M.","year":"2014","journal-title":"Fertility and Sterility"},{"issue":"1","key":"e_1_2_9_26_1","doi-asserted-by":"crossref","first-page":"3216","DOI":"10.1038\/s41598-020-80857-2","article-title":"Sperm DNA Methylation Mediates the Association of Male Age on Reproductive Outcomes Among Couples Undergoing Infertility Treatment","volume":"11","author":"Oluwayiose O. A.","year":"2021","journal-title":"Scientific Reports"},{"key":"e_1_2_9_27_1","first-page":"271","volume-title":"WHO Laboratory Manual for the Examination and Processing of Human Semen","author":"World Health Organization","year":"2010"},{"issue":"6","key":"e_1_2_9_28_1","doi-asserted-by":"crossref","first-page":"293","DOI":"10.2144\/000114280","article-title":"Rapid Method for the Isolation of Mammalian Sperm DNA","volume":"58","author":"Wu H.","year":"2015","journal-title":"Biotechniques"},{"issue":"7","key":"e_1_2_9_29_1","first-page":"1581","article-title":"Sperm Epigenetic Clock Associates With Pregnancy Outcomes in the General Population","volume":"37","author":"Pilsner J. R.","year":"2022","journal-title":"Human Reproduction"},{"issue":"9","key":"e_1_2_9_30_1","doi-asserted-by":"crossref","first-page":"1280","DOI":"10.1093\/bioinformatics\/bts124","article-title":"CpGassoc: An R Function for Analysis of DNA Methylation Microarray Data","volume":"28","author":"Barfield R. T.","year":"2012","journal-title":"Bioinformatics"},{"issue":"1","key":"e_1_2_9_31_1","doi-asserted-by":"crossref","first-page":"763","DOI":"10.1186\/s12864-018-5153-4","article-title":"Paternal Germ Line Aging: DNA Methylation Age Prediction From Human Sperm","volume":"19","author":"Jenkins T. G.","year":"2018","journal-title":"BMC Genomics"},{"issue":"2","key":"e_1_2_9_32_1","doi-asserted-by":"crossref","first-page":"146","DOI":"10.1080\/19396368.2018.1434838","article-title":"Pre\u2010Screening Method for Somatic Cell Contamination in Human Sperm Epigenetic Studies","volume":"64","author":"Jenkins T. G.","year":"2018","journal-title":"Systems Biology in Reproductive Medicine"},{"issue":"1","key":"e_1_2_9_33_1","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1186\/s13059-016-1131-9","article-title":"Controlling Bias and Inflation in Epigenome\u2010 and Transcriptome\u2010Wide Association Studies Using the Empirical Null Distribution","volume":"18","author":"Iterson M.","year":"2017","journal-title":"Genome biology"},{"issue":"5","key":"e_1_2_9_34_1","doi-asserted-by":"crossref","first-page":"711","DOI":"10.1093\/bioinformatics\/btaa732","article-title":"ipDMR: Identification of Differentially Methylated Regions With Interval P\u2010Values","volume":"37","author":"Xu Z.","year":"2021","journal-title":"Bioinformatics"},{"issue":"17","key":"e_1_2_9_35_1","doi-asserted-by":"crossref","first-page":"2190","DOI":"10.1093\/bioinformatics\/btq340","article-title":"METAL: Fast and Efficient Meta\u2010Analysis of Genomewide Association Scans","volume":"26","author":"Willer C. J.","year":"2010","journal-title":"Bioinformatics"},{"key":"e_1_2_9_36_1","doi-asserted-by":"crossref","DOI":"10.1016\/j.envint.2024.109049","article-title":"Urinary Concentrations of Phthalate and Phthalate Alternative Metabolites and Sperm DNA Methylation: A Multi\u2010Cohort and Meta\u2010Analysis of Men in Preconception Studies","volume":"192","author":"Nowak K.","year":"2024","journal-title":"Environment International"},{"issue":"1","key":"e_1_2_9_37_1","doi-asserted-by":"crossref","first-page":"1523","DOI":"10.1038\/s41467-019-09234-6","article-title":"Metascape Provides a Biologist\u2010Oriented Resource for the Analysis of Systems\u2010Level Datasets","volume":"10","author":"Zhou Y.","year":"2019","journal-title":"Nature Communications"},{"issue":"3","key":"e_1_2_9_38_1","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1093\/humupd\/dmp048","article-title":"World Health Organization Reference Values for Human Semen Characteristics","volume":"16","author":"Cooper T. G.","year":"2010","journal-title":"Human Reproduction Update"},{"issue":"1","key":"e_1_2_9_39_1","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1038\/s41467-018-08182-x","article-title":"Loss\u2010of\u2010Function Mutations in QRICH2 Cause Male Infertility With Multiple Morphological Abnormalities of the Sperm Flagella","volume":"10","author":"Shen Y.","year":"2019","journal-title":"Nature Communications"},{"key":"e_1_2_9_40_1","doi-asserted-by":"crossref","DOI":"10.3389\/fgene.2022.1017302","article-title":"A Homozygous Missense Variant in DND1 Causes Non\u2010Obstructive Azoospermia in Humans","volume":"13","author":"Xie X.","year":"2022","journal-title":"Frontiers in Genetics"},{"key":"e_1_2_9_41_1","doi-asserted-by":"crossref","DOI":"10.7717\/peerj.18240","article-title":"Interaction of CTCF and CTCFL in Genome Regulation Through Chromatin Architecture During the Spermatogenesis and Carcinogenesis","volume":"12","author":"Tong X.","year":"2024","journal-title":"PeerJ"},{"issue":"1","key":"e_1_2_9_42_1","doi-asserted-by":"crossref","first-page":"3846","DOI":"10.1038\/s41467-021-24140-6","article-title":"The Combined Action of CTCF and Its Testis\u2010Specific Paralog BORIS Is Essential for Spermatogenesis","volume":"12","author":"Rivero\u2010Hinojosa S.","year":"2021","journal-title":"Nature Communications"},{"issue":"14","key":"e_1_2_9_43_1","doi-asserted-by":"crossref","first-page":"1688","DOI":"10.1016\/j.febslet.2015.03.023","article-title":"Functional and Structural Insights Into the piRNA Factor Maelstrom","volume":"589","author":"Sato K.","year":"2015","journal-title":"Febs Letters"},{"key":"e_1_2_9_44_1","doi-asserted-by":"crossref","first-page":"E292","DOI":"10.1016\/j.fertnstert.2016.07.830","article-title":"Epigenetic Dysregulation of Mael Gene Contributes to One of the Causes of Non\u2010Obstructive Azoospermia With Hypospermatogenesis","volume":"106","author":"Lu Y.","year":"2016","journal-title":"Fertility and Sterility"},{"issue":"7","key":"e_1_2_9_45_1","doi-asserted-by":"crossref","first-page":"1286","DOI":"10.1111\/andr.13408","article-title":"The MAEL Expression in Mitochondria of Human Spermatozoa and the Association With Asthenozoospermia","volume":"11","author":"Cheng Y. S.","year":"2023","journal-title":"Andrology"},{"issue":"1","key":"e_1_2_9_46_1","doi-asserted-by":"crossref","first-page":"6637","DOI":"10.1038\/s41467-024-50930-9","article-title":"Inherited Defects of piRNA Biogenesis Cause Transposon De\u2010Repression, Impaired Spermatogenesis, and Human Male Infertility","volume":"15","author":"Stallmeyer B.","year":"2024","journal-title":"Nature Communications"},{"issue":"11","key":"e_1_2_9_47_1","doi-asserted-by":"crossref","first-page":"1322","DOI":"10.1016\/j.jgg.2024.06.014","article-title":"Pathogenic TDRD12 Variants Cause Defective piRNA Pathway and Male Infertility in Humans and Mice","volume":"51","author":"Bao Z.","year":"2024","journal-title":"Journal of Genetics and Genomics"},{"issue":"8","key":"e_1_2_9_48_1","doi-asserted-by":"crossref","first-page":"1014","DOI":"10.1016\/j.modgep.2006.04.007","article-title":"Sohlh2 is a Germ Cell\u2010Specific bHLH Transcription Factor","volume":"6","author":"Ballow D. J.","year":"2006","journal-title":"Gene Expression Patterns"},{"key":"e_1_2_9_49_1","doi-asserted-by":"crossref","DOI":"10.1038\/srep20980","article-title":"SOHLH2 is Essential for Synaptonemal Complex Formation During Spermatogenesis in Early Postnatal Mouse Testes","volume":"6","author":"Park M.","year":"2016","journal-title":"Scientific Reports"},{"issue":"1","key":"e_1_2_9_50_1","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1016\/j.ajhg.2021.11.022","article-title":"Bi\u2010Allelic Variants in DNHD1 Cause Flagellar Axoneme Defects and Asthenoteratozoospermia in Humans and Mice","volume":"109","author":"Tan C.","year":"2022","journal-title":"American Journal of Human Genetics"},{"issue":"3","key":"e_1_2_9_51_1","doi-asserted-by":"crossref","first-page":"2559","DOI":"10.3390\/ijms24032559","article-title":"New Mutations in DNHD1 Cause Multiple Morphological Abnormalities of the Sperm Flagella","volume":"24","author":"Martinez G.","year":"2023","journal-title":"International Journal of Molecular Sciences"},{"issue":"6","key":"e_1_2_9_52_1","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1002\/mrd.22650","article-title":"A Genomic Region Transcribed Into a Long Noncoding RNA Interacts With the Prss42\/Tessp\u20102 Promoter in Spermatocytes During Mouse Spermatogenesis, and Its Flanking Sequences Can Function as Enhancers","volume":"83","author":"Yoneda R.","year":"2016","journal-title":"Molecular Reproduction and Development"},{"issue":"4","key":"e_1_2_9_53_1","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0124293","article-title":"A Novel Transcriptional Factor Nkapl Is a Germ Cell\u2010Specific Suppressor of Notch Signaling and Is Indispensable for Spermatogenesis","volume":"10","author":"Okuda H.","year":"2015","journal-title":"PLoS ONE"},{"issue":"1","key":"e_1_2_9_54_1","doi-asserted-by":"crossref","first-page":"791","DOI":"10.1038\/s41467-024-55579-y","article-title":"NKAPL Facilitates Transcription Pause\u2010Release and Bridges Elongation to Initiation During Meiosis Exit","volume":"16","author":"Kang Z.","year":"2025","journal-title":"Nature Communications"},{"issue":"1","key":"e_1_2_9_55_1","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1186\/s13072-018-0223-z","article-title":"Modulation of Mitochondrial DNA Copy Number in a Model of Glioblastoma Induces Changes to DNA Methylation and Gene Expression of the Nuclear Genome in Tumours","volume":"11","author":"Sun X. S.","year":"2018","journal-title":"Epigenetics & Chromatin"},{"issue":"1","key":"e_1_2_9_56_1","doi-asserted-by":"crossref","first-page":"2083","DOI":"10.1038\/s41598-024-52373-0","article-title":"Nuclear and Mitochondrial Genetic Variants Associated With Mitochondrial DNA Copy Number","volume":"14","author":"Koller A.","year":"2024","journal-title":"Scientific Reports"},{"issue":"1","key":"e_1_2_9_57_1","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1186\/s13073-020-00778-7","article-title":"Mitochondrial DNA Copy Number Can Influence Mortality and Cardiovascular Disease via Methylation of Nuclear DNA CpGs","volume":"12","author":"Castellani C. A.","year":"2020","journal-title":"Genome Medicine"}],"container-title":["Andrology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/andr.70320","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1111\/andr.70320","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/andr.70320","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,20]],"date-time":"2026-07-20T07:25:34Z","timestamp":1784532334000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1111\/andr.70320"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,20]]},"references-count":56,"alternative-id":["10.1111\/andr.70320"],"URL":"https:\/\/doi.org\/10.1111\/andr.70320","archive":["Portico"],"relation":{},"ISSN":["2047-2919","2047-2927"],"issn-type":[{"value":"2047-2919","type":"print"},{"value":"2047-2927","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,20]]},"assertion":[{"value":"2026-01-16","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-07-04","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-07-20","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"andr.70320"}}