{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,22]],"date-time":"2026-10-22T18:39:15Z","timestamp":1792694355774,"version":"4.5.0"},"reference-count":62,"publisher":"Oxford University Press (OUP)","issue":"4","license":[{"start":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T00:00:00Z","timestamp":1789084800000},"content-version":"vor","delay-in-days":253,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100009708","name":"Novo Nordisk Foundation","doi-asserted-by":"publisher","award":["NNF18OC0033754"],"award-info":[{"award-number":["NNF18OC0033754"]}],"id":[{"id":"10.13039\/501100009708","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,17]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:sec>\n                    <jats:title>STUDY QUESTION<\/jats:title>\n                    <jats:p>Is DNA methylation in mature spermatozoa affected by different culture media?<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>SUMMARY ANSWER<\/jats:title>\n                    <jats:p>Incubation of mature spermatozoa in hydrogen peroxide (H\u2082O\u2082) or palmitate for 24\u2009h did not alter the DNA methylome.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>WHAT IS KNOWN ALREADY<\/jats:title>\n                    <jats:p>Children conceived through ART face a higher risk of disorders linked to abnormal DNA methylation. As culture media can modulate DNA methylation in somatic cells, abnormal DNA methylation after ART may be caused by factors present in culture media during sperm incubation.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>STUDY DESIGN, SIZE, DURATION<\/jats:title>\n                    <jats:p>Sperm from four donors underwent targeted shotgun proteomics analysis, focusing on protein size ranges that include epigenetic modifying enzymes. Additionally, sperm samples from 15 men were incubated for 24\u2009h in either standard media, media with H\u2082O\u2082, or media containing palmitate, two media known to induce acute DNA methylation changes in somatic cells. DNA methylation profiles of these incubated sperm samples were then compared to those of pre-incubation sperm using Reduced Representation Bisulfite Sequencing (RRBS).<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>PARTICIPANTS\/MATERIALS, SETTING, METHODS<\/jats:title>\n                    <jats:p>Sperm samples were obtained from 15 healthy Caucasian men aged 20\u201340\u2009years. Participants using prescription medications daily or with a history of testicular disease were excluded. Motile sperm were isolated through the swim-up technique. Protein analysis was conducted via in-gel digestion followed by mass spectrometry. DNA methylation was evaluated using RRBS and validated for select genomic regions with targeted bisulfite pyrosequencing.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>MAIN RESULTS AND THE ROLE OF CHANCE<\/jats:title>\n                    <jats:p>Proteomic analysis returned 1930 proteins, including 144 newly identified proteins. Consistent with previous studies, we confirmed the presence of DNA methyltransferase 1 (DNMT1), while de novo methyltransferases DNMT3A and DNMT3B were not detected. Incubation in control, H\u2082O\u2082, and palmitate media all led to decreased sperm motility, with the decline occurring the fastest under the H\u2082O\u2082\u00a0condition. Sperm DNA methylation remained unchanged across all three culture conditions.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>LARGE SCALE DATA<\/jats:title>\n                    <jats:p>N\/A<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>LIMITATIONS, REASONS FOR CAUTION<\/jats:title>\n                    <jats:p>RRBS analysis covers only a portion of the genome, so DNA methylation changes in unexamined regions cannot be excluded, particularly at repeated regions of the genome. Moreover, since this study tested only three culture media conditions, it remains possible that other media formulations could impact DNA methylation.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>WIDER IMPLICATIONS OF THE FINDINGS<\/jats:title>\n                    <jats:p>This study indicates that the DNA methylome of mature human spermatozoa is stable despite the presence of DNA methylation machinery components. These results imply that DNA methylation changes in response to environmental stress are more likely to happen during earlier stages of spermatogenesis. While the various culture media used in ART are unlikely to alter the DNA methylation patterns of individual sperm cells, the possibility that certain sperm with specific methylation signatures are selectively favored cannot be ruled out.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>FUNDING<\/jats:title>\n                    <jats:p>This work was supported by a Challenge Programme Grant from the Novo Nordisk Foundation (NNF18OC0033754) to the Gametic Epigenetics Consortium against Obesity (GECKO). The Novo Nordisk Foundation Center for Basic Metabolic Research is an independent research center at the University of Copenhagen, partially funded by an unrestricted donation from the Novo Nordisk Foundation (NNF18CC0034900). This work was supported by the French Government (National Research Agency, ANR) through the \u2018Investments for the Future\u2019 programs LABEX SIGNALIFE ANR-11-LABX-0028-01 and IDEX UCAJedi ANR-15-IDEX-01.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>DISCLOSURES<\/jats:title>\n                    <jats:p>R.B. is a recipient of research funding granted by the Novo Nordisk Foundation. E.A. is a shareholder of ExSeed Health Ltd.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.1093\/hropen\/hoag079","type":"journal-article","created":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T11:48:31Z","timestamp":1789040911000},"source":"Crossref","is-referenced-by-count":0,"title":["<i>In vitro<\/i>\n                    incubation of human sperm does not change DNA methylation"],"prefix":"10.1093","volume":"2026","author":[{"given":"Emma Tykesson","family":"Kj\u00f8ller","sequence":"first","affiliation":[{"name":"Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen , Copenhagen,","place":["Denmark"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marie-Charlotte","family":"Dumargne","sequence":"additional","affiliation":[{"name":"Institut de Pharmacologie Mol\u00e9culaire et Cellulaire, CNRS, Inserm and Universit\u00e9 C\u00f4te d\u2018Azur , Valbonne,","place":["France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lars Roed","family":"Ingerslev","sequence":"additional","affiliation":[{"name":"Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen , Copenhagen,","place":["Denmark"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eleonora","family":"Manitta","sequence":"additional","affiliation":[{"name":"Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen , Copenhagen,","place":["Denmark"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yasmin","family":"Dehestani","sequence":"additional","affiliation":[{"name":"Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen , Copenhagen,","place":["Denmark"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marte","family":"Saupstad","sequence":"additional","affiliation":[{"name":"Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen , Copenhagen,","place":["Denmark"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alba","family":"Gonzalez-Franquesa","sequence":"additional","affiliation":[{"name":"Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen , Copenhagen,","place":["Denmark"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Atul Shahaji","family":"Deshmukh","sequence":"additional","affiliation":[{"name":"Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen , Copenhagen,","place":["Denmark"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6009-2099","authenticated-orcid":false,"given":"Emil","family":"Andersen","sequence":"additional","affiliation":[{"name":"Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen , Copenhagen,","place":["Denmark"]},{"name":"ExSeed Health ApS , Copenhagen,","place":["Denmark"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0158-519X","authenticated-orcid":false,"given":"Romain","family":"Barr\u00e8s","sequence":"additional","affiliation":[{"name":"Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen , Copenhagen,","place":["Denmark"]},{"name":"Institut de Pharmacologie Mol\u00e9culaire et Cellulaire, CNRS, Inserm and Universit\u00e9 C\u00f4te d\u2018Azur , Valbonne,","place":["France"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,9,11]]},"reference":[{"key":"2026102213582131400_hoag079-B2","doi-asserted-by":"crossref","first-page":"861","DOI":"10.2217\/epi-2019-0055","article-title":"Environmental factors influence the epigenetic signature of newborns from mothers with gestational diabetes","volume":"11","author":"Andersen","year":"2019","journal-title":"Epigenomics"},{"key":"2026102213582131400_hoag079-B3","doi-asserted-by":"crossref","first-page":"306","DOI":"10.1038\/s41366-018-0031-3","article-title":"Preadipocytes from obese humans with type 2 diabetes are epigenetically reprogrammed at genes controlling adipose tissue function","volume":"43","author":"Andersen","year":"2019","journal-title":"Int J Obes (Lond)"},{"key":"2026102213582131400_hoag079-B4","doi-asserted-by":"crossref","first-page":"1388","DOI":"10.1016\/j.fertnstert.2015.08.019","article-title":"Aberrant sperm DNA methylation predicts male fertility status and embryo quality","volume":"104","author":"Aston","year":"2015","journal-title":"Fertil Steril"},{"key":"2026102213582131400_hoag079-B5","doi-asserted-by":"crossref","first-page":"3566","DOI":"10.3390\/cells10123566","article-title":"Sperm selection for ICSI: do we have a winner?","volume":"10","author":"Baldini","year":"2021","journal-title":"Cells"},{"key":"2026102213582131400_hoag079-B6","doi-asserted-by":"crossref","first-page":"479","DOI":"10.1093\/humrep\/deaa310","article-title":"Do assisted reproductive technologies and in vitro embryo culture influence the epigenetic control of imprinted genes and transposable elements in children?","volume":"36","author":"Barberet","year":"2021","journal-title":"Hum Reprod"},{"key":"2026102213582131400_hoag079-B7","doi-asserted-by":"crossref","first-page":"1468","DOI":"10.1016\/j.fertnstert.2021.08.014","article-title":"Do frozen embryo transfers modify the epigenetic control of imprinted genes and transposable elements in newborns compared with fresh embryo transfers and natural conceptions?","volume":"116","author":"Barberet","year":"2021","journal-title":"Fertil Steril"},{"key":"2026102213582131400_hoag079-B8","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/j.cmet.2009.07.011","article-title":"Non-CpG methylation of the PGC-1alpha promoter through DNMT3B controls mitochondrial density","volume":"10","author":"Barr\u00e8s","year":"2009","journal-title":"Cell Metab"},{"key":"2026102213582131400_hoag079-B9","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1016\/j.cmet.2012.01.001","article-title":"Acute exercise remodels promoter methylation in human skeletal muscle","volume":"15","author":"Barr\u00e8s","year":"2012","journal-title":"Cell Metab"},{"key":"2026102213582131400_hoag079-B10","doi-asserted-by":"crossref","first-page":"1109","DOI":"10.1016\/j.dnarep.2004.03.002","article-title":"DNA damage responses to oxidative stress","volume":"3","author":"Barzilai","year":"2004","journal-title":"DNA Repair (Amst)"},{"key":"2026102213582131400_hoag079-B11","doi-asserted-by":"crossref","first-page":"4340","DOI":"10.1021\/acs.jproteome.7b00374","article-title":"Validating missing proteins in human sperm cells by targeted mass-spectrometry- and antibody-based methods","volume":"16","author":"Carapito","year":"2017","journal-title":"J Proteome Res"},{"key":"2026102213582131400_hoag079-B2874972","doi-asserted-by":"crossref","first-page":"hoz031","DOI":"10.1093\/hropen\/hoz031","article-title":"The impact of selected embryo culture conditions on ART treatment cycle outcomes: a UK national study","volume":"2020","author":"Castillo","year":"2020","journal-title":"Hum Reprod Open"},{"key":"2026102213582131400_hoag079-B14","doi-asserted-by":"crossref","first-page":"535","DOI":"10.1093\/humupd\/dmy017","article-title":"The contribution of human sperm proteins to the development and epigenome of the preimplantation embryo","volume":"24","author":"Castillo","year":"2018","journal-title":"Hum Reprod Update"},{"key":"2026102213582131400_hoag079-B16","doi-asserted-by":"crossref","first-page":"372","DOI":"10.1016\/j.rbmo.2019.02.007","article-title":"Differential impacts of gonadotrophins, IVF and embryo culture on mouse blastocyst development","volume":"39","author":"Chen","year":"2019","journal-title":"Reprod Biomed Online"},{"key":"2026102213582131400_hoag079-B17","doi-asserted-by":"crossref","first-page":"3189","DOI":"10.2337\/db14-0103","article-title":"Altered glucose metabolism in mouse and humans conceived by IVF","volume":"63","author":"Chen","year":"2014","journal-title":"Diabetes"},{"key":"2026102213582131400_hoag079-B18","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1016\/j.placenta.2017.11.009","article-title":"Maternal high estradiol exposure alters CDKN1C and IGF2 expression in human placenta","volume":"61","author":"Chen","year":"2018","journal-title":"Placenta"},{"key":"2026102213582131400_hoag079-B6240639","doi-asserted-by":"crossref","first-page":"e20210132","DOI":"10.1590\/1984-3143-ar2021-0132","article-title":"The embryo culture media in the era of epigenetics: is it time to go back to nature?","volume":"19","author":"Coy","year":"2022","journal-title":"Anim Reprod"},{"key":"2026102213582131400_hoag079-B21","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1093\/humrep\/deu286","article-title":"The type of culture medium and the duration of in vitro culture do not influence birthweight of ART singletons","volume":"30","author":"De Vos","year":"2015","journal-title":"Hum Reprod"},{"key":"2026102213582131400_hoag079-B23","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":"2026102213582131400_hoag079-B24","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1055\/s-0032-1307417","article-title":"Role of ART in imprinting disorders","volume":"30","author":"Eroglu","year":"2012","journal-title":"Semin Reprod Med"},{"key":"2026102213582131400_hoag079-B30","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1093\/molehr\/gap038","article-title":"Abnormal methylation at the KvDMR1 imprinting control region in clinically normal children conceived by assisted reproductive technologies","volume":"15","author":"Gomes","year":"2009","journal-title":"Mol Hum Reprod"},{"key":"2026102213582131400_hoag079-B4260323","doi-asserted-by":"crossref","first-page":"468","DOI":"10.1038\/nprot.2010.190","article-title":"Preparation of reduced representation bisulfite sequencing libraries for genome-scale DNA methylation profiling","volume":"6","author":"Gu","year":"2011","journal-title":"Nat Protoc"},{"key":"2026102213582131400_hoag079-B0522675","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1186\/1741-7015-12-103","article-title":"Effects of palmitate on genome-wide mRNA expression and DNA methylation patterns in human pancreatic islets","volume":"12","author":"Hall","year":"2014","journal-title":"BMC Med"},{"key":"2026102213582131400_hoag079-B3490349","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1186\/s13148-019-0623-3","article-title":"Association of four imprinting disorders and ART","volume":"11","author":"Hattori","year":"2019","journal-title":"Clin Epigenetics"},{"key":"2026102213582131400_hoag079-B36","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1016\/j.fertnstert.2015.09.013","article-title":"Decreased fecundity and sperm DNA methylation patterns","volume":"105","author":"Jenkins","year":"2016","journal-title":"Fertil Steril"},{"key":"2026102213582131400_hoag079-B38","doi-asserted-by":"crossref","first-page":"717729","DOI":"10.3389\/fcvm.2021.717729","article-title":"Elevated Expression of lncRNA MEG3 induces endothelial dysfunction on HUVECs of IVF born offspring via epigenetic regulation","volume":"8","author":"Jiang","year":"2021","journal-title":"Front Cardiovasc Med"},{"key":"2026102213582131400_hoag079-B39","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1080\/17501911.2024.2439782","article-title":"Impact of excess sugar on the whole genome DNA methylation pattern in human sperm","volume":"17","author":"J\u00f6nsson","year":"2025","journal-title":"Epigenomics"},{"key":"2026102213582131400_hoag079-B40","doi-asserted-by":"crossref","first-page":"g252","DOI":"10.1136\/bmj.g252","article-title":"Are we overusing IVF?","volume":"348","author":"Kamphuis","year":"2014","journal-title":"BMJ"},{"key":"2026102213582131400_hoag079-B5903018","doi-asserted-by":"crossref","first-page":"2204","DOI":"10.1093\/bioinformatics\/btq351","article-title":"BigWig and BigBed: enabling browsing of large distributed datasets","volume":"26","author":"Kent","year":"2010","journal-title":"Bioinformatics"},{"key":"2026102213582131400_hoag079-B75537319","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1093\/biostatistics\/kxy007","article-title":"Detection and accurate false discovery rate control of differentially methylated regions from whole genome bisulfite sequencing","volume":"20","author":"Korthauer","year":"2019","journal-title":"Biostatistics"},{"key":"2026102213582131400_hoag079-B42","doi-asserted-by":"crossref","first-page":"2219","DOI":"10.1093\/humrep\/dew156","article-title":"Influence of embryo culture medium (G5 and HTF) on pregnancy and perinatal outcome after IVF: a multicenter RCT","volume":"31","author":"Kleijkers","year":"2016","journal-title":"Hum Reprod"},{"key":"2026102213582131400_hoag079-B8748177","doi-asserted-by":"crossref","first-page":"hoaf041","DOI":"10.1093\/hropen\/hoaf041","article-title":"Effects of different embryo culture media on birthweight following assisted reproductive technology","volume":"2025","author":"Li","year":"2025","journal-title":"Hum Reprod Open"},{"key":"2026102213582131400_hoag079-B45","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1017\/S2040174415000161","article-title":"Association of in vitro fertilization with global and IGF2\/H19 methylation variation in newborn twins","volume":"6","author":"Loke","year":"2015","journal-title":"J Dev Orig Health Dis"},{"key":"2026102213582131400_hoag079-B46","doi-asserted-by":"crossref","first-page":"997","DOI":"10.1177\/1933719118802052","article-title":"Aberrant DNA methylation of IGF2-H19 locus in human fetus and in spermatozoa from assisted reproductive technologies","volume":"26","author":"Lou","year":"2019","journal-title":"Reprod Sci"},{"key":"2026102213582131400_hoag079-B47","doi-asserted-by":"crossref","first-page":"4064","DOI":"10.3390\/cells11244064","article-title":"Proteomic Landscape of Human Spermatozoa: Optimized Extraction Method and Application","volume":"11","author":"Luo","year":"2022","journal-title":"Cells"},{"key":"2026102213582131400_hoag079-B48","doi-asserted-by":"crossref","first-page":"1987","DOI":"10.1093\/humrep\/dew173","article-title":"No change in live birthweight of IVF singleton deliveries over an 18-year period despite significant clinical and laboratory changes","volume":"31","author":"Maas","year":"2016","journal-title":"Hum Reprod"},{"key":"2026102213582131400_hoag079-B1445561","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1152\/physiolgenomics.00065.2014","article-title":"Lipid exposure elicits differential responses in gene expression and DNA methylation in primary human skeletal muscle cells from severely obese women","volume":"47","author":"Maples","year":"2015","journal-title":"Physiol Genomics"},{"key":"2026102213582131400_hoag079-B50","doi-asserted-by":"crossref","first-page":"1354","DOI":"10.4161\/epi.6.11.17993","article-title":"DNA methylation imprinting marks and DNA methyltransferase expression in human spermatogenic cell stages","volume":"6","author":"Marques","year":"2011","journal-title":"Epigenetics"},{"key":"2026102213582131400_hoag079-B54","doi-asserted-by":"crossref","first-page":"1117","DOI":"10.1093\/humrep\/des459","article-title":"Placentas from pregnancies conceived by IVF\/ICSI have a reduced DNA methylation level at the H19 and MEST differentially methylated regions","volume":"28","author":"Nelissen","year":"2013","journal-title":"Hum Reprod"},{"key":"2026102213582131400_hoag079-B55","doi-asserted-by":"crossref","first-page":"1411","DOI":"10.1093\/humrep\/dew096","article-title":"TET enzymes are successively expressed during human spermatogenesis and their expression level is pivotal for male fertility","volume":"31","author":"Ni","year":"2016","journal-title":"Hum Reprod"},{"key":"2026102213582131400_hoag079-B5179522","doi-asserted-by":"crossref","first-page":"123484","DOI":"10.1155\/2015\/123484","article-title":"Environmental Impact on DNA Methylation in the Germline: State of the Art and Gaps of Knowledge","volume":"2015","author":"Pacchierotti","year":"2015","journal-title":"Biomed Res Int"},{"key":"2026102213582131400_hoag079-B62","doi-asserted-by":"crossref","first-page":"e250105","DOI":"10.1530\/REP-25-0105","article-title":"Towards a kingdom of reproductive life\u2014the core sperm proteome","volume":"169","author":"Pini","year":"2025","journal-title":"Reproduction"},{"key":"2026102213582131400_hoag079-B7646263","doi-asserted-by":"crossref","first-page":"328","DOI":"10.3390\/jcm13020328","article-title":"Investigating the Association of Assisted Reproduction Techniques and Adverse Perinatal Outcomes","volume":"13","author":"Potiris","year":"2024","journal-title":"J Clin Med"},{"key":"2026102213582131400_hoag079-B40779588","author":"R Core Team.","year":"2022"},{"key":"2026102213582131400_hoag079-B67","doi-asserted-by":"crossref","first-page":"2208","DOI":"10.1093\/humrep\/des151","article-title":"Methylation levels at imprinting control regions are not altered with ovulation induction or in vitro fertilization in a birth cohort","volume":"27","author":"Rancourt","year":"2012","journal-title":"Hum Reprod"},{"key":"2026102213582131400_hoag079-B31158810","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1038\/nbt.1754","article-title":"Integrative genomics viewer","volume":"29","author":"Robinson","year":"2011","journal-title":"Nat Biotechnol"},{"key":"2026102213582131400_hoag079-B69","doi-asserted-by":"crossref","first-page":"1100","DOI":"10.1016\/j.placenta.2015.08.008","article-title":"Altered gene expression of H19 and IGF2 in placentas from ART pregnancies","volume":"36","author":"Sakian","year":"2015","journal-title":"Placenta"},{"key":"2026102213582131400_hoag079-B70","doi-asserted-by":"crossref","first-page":"S109","DOI":"10.1074\/mcp.RA117.000541","article-title":"Characterization of the sperm proteome and reproductive outcomes with in vitro, fertilization after a reduction in male ejaculatory abstinence period","volume":"18","author":"Shen","year":"2019","journal-title":"Mol Cell Proteomics"},{"key":"2026102213582131400_hoag079-B95290270","doi-asserted-by":"crossref","first-page":"308","DOI":"10.1093\/nar\/29.1.308","article-title":"dbSNP: the NCBI database of genetic variation","volume":"29","author":"Sherry","year":"2001","journal-title":"Nucleic Acids Res"},{"key":"2026102213582131400_hoag079-B6475918","doi-asserted-by":"crossref","first-page":"272","DOI":"10.1038\/nrg2072","article-title":"Transposable elements and the epigenetic regulation of the genome","volume":"8","author":"Slotkin","year":"2007","journal-title":"Nat Rev Genet"},{"key":"2026102213582131400_hoag079-B72","doi-asserted-by":"crossref","first-page":"690","DOI":"10.1016\/j.rbmo.2016.08.022","article-title":"DNA methyltransferases exhibit dynamic expression during spermatogenesis","volume":"33","author":"Uysal","year":"2016","journal-title":"Reprod Biomed Online"},{"key":"2026102213582131400_hoag079-B73","doi-asserted-by":"crossref","first-page":"3998","DOI":"10.1021\/acs.jproteome.6b00400","article-title":"Looking for missing proteins in the proteome of human spermatozoa: an update","volume":"15","author":"Vandenbrouck","year":"2016","journal-title":"J Proteome Res"},{"key":"2026102213582131400_hoag079-B74","doi-asserted-by":"crossref","first-page":"1076","DOI":"10.1093\/biolre\/ioz032","article-title":"Sperm selection methods in the 21st century","volume":"101","author":"Vaughan","year":"2019","journal-title":"Biol Reprod"},{"key":"2026102213582131400_hoag079-B75","doi-asserted-by":"crossref","first-page":"114","DOI":"10.1016\/j.jprot.2012.12.008","article-title":"In-depth proteomic analysis of the human sperm reveals complex protein compositions","volume":"79","author":"Wang","year":"2013","journal-title":"J Proteomics"},{"key":"2026102213582131400_hoag079-B76","doi-asserted-by":"crossref","first-page":"5449","DOI":"10.1073\/pnas.0610317104","article-title":"Mouse embryo culture induces changes in postnatal phenotype including raised systolic blood pressure","volume":"104","author":"Watkins","year":"2007","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026102213582131400_hoag079-B77","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1038\/ng1990","article-title":"Distribution, silencing potential and evolutionary impact of promoter DNA methylation in the human genome","volume":"39","author":"Weber","year":"2007","journal-title":"Nat Genet"},{"key":"2026102213582131400_hoag079-B78","author":"WHO","year":"2010"},{"key":"2026102213582131400_hoag079-B16214924","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-319-24277-4","author":"Wickham","year":"2016","journal-title":"ggplot2: Elegant Graphics for Data Analysis"},{"key":"2026102213582131400_hoag079-B80","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1016\/S0015-0282(16)49104-1","article-title":"Sperm washing and swim-up technique using antibiotics removes microbes from human semen","volume":"45","author":"Wong","year":"1986","journal-title":"Fertil Steril"},{"key":"2026102213582131400_hoag079-B0712441","author":"World Health Organization","year":"2023","journal-title":"Infertility Prevalence Estimates, 1990\u20132021"},{"key":"2026102213582131400_hoag079-B81","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","year":"2015","journal-title":"Biotechniques"},{"key":"2026102213582131400_hoag079-B7296897","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1089\/omi.2011.0118","article-title":"clusterProfiler: an R package for comparing biological themes among gene clusters","volume":"16","author":"Yu","year":"2012","journal-title":"OMICS"},{"key":"2026102213582131400_hoag079-B82","doi-asserted-by":"crossref","first-page":"1564","DOI":"10.1002\/pmic.201400248","article-title":"Comparative analysis of macaque and human sperm proteomes: insights into sperm competition","volume":"15","author":"Zhou","year":"2015","journal-title":"Proteomics"}],"container-title":["Human Reproduction Open"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/hropen\/advance-article-pdf\/doi\/10.1093\/hropen\/hoag079\/71032706\/hoag079.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/hropen\/article-pdf\/2026\/4\/hoag079\/71032706\/hoag079.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/hropen\/article-pdf\/2026\/4\/hoag079\/71032706\/hoag079.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,22]],"date-time":"2026-10-22T17:58:34Z","timestamp":1792691914000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/hropen\/article\/doi\/10.1093\/hropen\/hoag079\/8790601"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026]]},"references-count":62,"journal-issue":{"issue":"4","published-print":{"date-parts":[[2026,9,17]]}},"URL":"https:\/\/doi.org\/10.1093\/hropen\/hoag079","relation":{},"ISSN":["2399-3529"],"issn-type":[{"value":"2399-3529","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026]]},"published":{"date-parts":[[2026]]},"article-number":"hoag079"}}