{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T16:05:40Z","timestamp":1785600340160,"version":"3.56.0"},"reference-count":91,"publisher":"MDPI AG","issue":"4","license":[{"start":{"date-parts":[[2021,11,13]],"date-time":"2021-11-13T00:00:00Z","timestamp":1636761600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["JDB"],"abstract":"<jats:p>Telomeres promote genome integrity by protecting chromosome ends from the activation of the DNA damage response and protecting chromosomes from the loss of coding sequences due to the end replication problem. Telomere length (TL) is progressively shortened as age progresses, thus resulting in cellular senescence. Therefore, TL is in strong adverse linear correlation with aging. Mounting evidence supports the notion that telomeres and male\/female infertility are in a close relationship, posing the biology of telomeres as a hot topic in the era of human-assisted reproduction. Specifically, the length of sperm telomeres is gradually increasing as men get older, while the telomere length of the oocytes seems not to follow similar patterns with that of sperm. Nonetheless, the telomere length of the embryos during the cleavage stages seems to have a paternal origin, but the telomere length can be further extended by telomerase activity during the blastocyst stage. The latter has been proposed as a new molecular biomarker with strong predictive value regarding male infertility. As far as the role of telomeres in assisted reproduction, the data is limited but the length of telomeres in both gametes seems to be affected mainly by the cause of infertility rather than the assisted reproductive therapy (ART) procedure itself. The present review aims to shed more light into the role of telomeres in human embryological parameters, including gametes and embryos and also presents opinions regarding the association between telomeres and in vitro fertilization (IVF).<\/jats:p>","DOI":"10.3390\/jdb9040049","type":"journal-article","created":{"date-parts":[[2021,11,14]],"date-time":"2021-11-14T20:52:28Z","timestamp":1636923148000},"page":"49","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":23,"title":["Insights into the Role of Telomeres in Human Embryological Parameters. Opinions Regarding IVF"],"prefix":"10.3390","volume":"9","author":[{"given":"George","family":"Anifandis","sequence":"first","affiliation":[{"name":"Department of Obstetrics and Gynecology, Faculty of Medicine, School of Health Sciences, University of Thessaly, 41222 Larisa, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria","family":"Samara","sequence":"additional","affiliation":[{"name":"Department of Anatomical Pathology, Faculty of Medicine, School of Health Sciences, University of Thessaly, 41222 Larisa, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1000-9100","authenticated-orcid":false,"given":"Mara","family":"Simopoulou","sequence":"additional","affiliation":[{"name":"Laboratory of Physiology, Medical School, National and Kapodistrian University of Athens, 11527 Athens, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0909-2365","authenticated-orcid":false,"given":"Christina I.","family":"Messini","sequence":"additional","affiliation":[{"name":"Department of Obstetrics and Gynecology, Faculty of Medicine, School of Health Sciences, University of Thessaly, 41222 Larisa, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9624-6352","authenticated-orcid":false,"given":"Katerina","family":"Chatzimeletiou","sequence":"additional","affiliation":[{"name":"Unit for Human Reproduction, 1st Department of Obstetrics and Gynecology, Faculty of Medicine, School of Health Sciences, Aristotelian University of Thessaloniki, 56403 Thessaloniki, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eleni","family":"Thodou","sequence":"additional","affiliation":[{"name":"Department of Anatomical Pathology, Faculty of Medicine, School of Health Sciences, University of Thessaly, 41222 Larisa, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alexandros","family":"Daponte","sequence":"additional","affiliation":[{"name":"Department of Obstetrics and Gynecology, Faculty of Medicine, School of Health Sciences, University of Thessaly, 41222 Larisa, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ioannis","family":"Georgiou","sequence":"additional","affiliation":[{"name":"Laboratory of Medical Genetics, Faculty of Medicine, School of Health Sciences, University of Ioannina, 45110 Ioannina, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,11,13]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1186\/s12958-015-0032-1","article-title":"A unique view on male infertility around the globe","volume":"13","author":"Agarwal","year":"2015","journal-title":"Reprod. Biol. Endocrinol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"738","DOI":"10.23736\/S0026-4784.18.04247-8","article-title":"Assisted reproductive technique in women of advanced fertility age","volume":"70","author":"Artini","year":"2018","journal-title":"Minerva Ginecol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"883","DOI":"10.1016\/j.hlc.2013.12.016","article-title":"The correlation between peripheral leukocyte telomere length and indicators of cardiovascular aging","volume":"23","author":"Zhang","year":"2014","journal-title":"Heart Lung Circ."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"161","DOI":"10.2174\/1874609808666150122153151","article-title":"Telomere length variations in aging and age-related diseases","volume":"7","author":"Rizvi","year":"2014","journal-title":"Curr. Aging Sci."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1097\/MCO.0b013e32834121b1","article-title":"Telomeres, lifestyle, cancer, and aging","volume":"14","author":"Shammas","year":"2011","journal-title":"Curr. Opin. Clin. Nutr. Metab. Care"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"492","DOI":"10.1006\/excr.1999.4602","article-title":"Normal human telomeres are not late replicating","volume":"251","author":"Wright","year":"1999","journal-title":"Exp. Cell Res."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1685","DOI":"10.1007\/s10815-015-0574-3","article-title":"A single-cell assay for telomere DNA content shows increasing telomere length heterogeneity, as well as increasing mean telomere length in human spermatozoa with advancing age","volume":"32","author":"Antunes","year":"2015","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1007\/s10815-017-1029-9","article-title":"Sub-fertile sperm cells exemplify telomere dysfunction","volume":"35","author":"Wiser","year":"2018","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"e12840","DOI":"10.1111\/and.12840","article-title":"Sperm telomere length in motile sperm selection techniques: A qFISH approach","volume":"50","author":"Lafuente","year":"2018","journal-title":"Andrologia"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"510","DOI":"10.1093\/molehr\/gat021","article-title":"Telomere lengths in human pronuclei, oocytes and spermatozoa","volume":"19","author":"Turner","year":"2013","journal-title":"Mol. Hum. Reprod."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"947","DOI":"10.1093\/molehr\/7.10.947","article-title":"Characterization of telomerase activity in the human oocyte and preimplantation embryo","volume":"7","author":"Wright","year":"2001","journal-title":"Mol. Hum. Reprod."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Treff, N.R., Su, J., Taylor, D., and Scott, R.T. (2011). Telomere DNA deficiency is associated with development of human embryonic aneuploidy. PLoS Genet., 7.","DOI":"10.1371\/journal.pgen.1002161"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"957","DOI":"10.1016\/j.fertnstert.2020.09.164","article-title":"Shorter telomere length of white blood cells is associated with higher rates of aneuploidy among infertile women undergoing in vitro fertilization","volume":"115","author":"Hanson","year":"2021","journal-title":"Fertil. Steril."},{"key":"ref_14","first-page":"1180","article-title":"Mortality and immortality at the cellular level. A review","volume":"62","author":"Hayflick","year":"1997","journal-title":"Biochemistry"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1007\/s00404-012-2632-8","article-title":"Analysis of sperm telomere length in men with idiopathic infertility","volume":"287","author":"Thilagavathi","year":"2013","journal-title":"Arch. Gynecol. Obstet."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1016\/j.aanat.2010.07.005","article-title":"Sizing the ends: Normal length of human telomeres","volume":"192","author":"Samassekou","year":"2010","journal-title":"Ann. Anat."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"R397","DOI":"10.1016\/j.cub.2016.01.056","article-title":"Shelterin","volume":"26","author":"Schmutz","year":"2016","journal-title":"Curr. Biol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1133","DOI":"10.1038\/nm1006-1133","article-title":"Telomeres and telomerase: The path from maize, Tetrahymena and yeast to human cancer and aging","volume":"12","author":"Blackburn","year":"2006","journal-title":"Nat. Med."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"966","DOI":"10.1172\/JCI64098","article-title":"Cellular senescence and the senescent secretory phenotype: Therapeutic opportunities","volume":"123","author":"Tchkonia","year":"2013","journal-title":"J. Clin. Invest."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1323","DOI":"10.1002\/jcp.24537","article-title":"Telomere length maintenance, shortening, and lengthening","volume":"229","author":"Zhao","year":"2014","journal-title":"J. Cell Physiol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"2785","DOI":"10.1016\/j.neurobiolaging.2015.06.017","article-title":"Heritability of telomere length in a study of long-lived families","volume":"36","author":"Honig","year":"2015","journal-title":"Neurobiol. Aging"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1089\/rej.2010.1085","article-title":"A natural product telomerase activator as part of a health maintenance program","volume":"14","author":"Harley","year":"2011","journal-title":"Rejuvenation Res."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"604","DOI":"10.1111\/j.1474-9726.2011.00700.x","article-title":"The telomerase activator TA-65 elongates short telomeres and increases health span of adult\/old mice without increasing cancer incidence","volume":"10","author":"Schneeberger","year":"2011","journal-title":"Aging Cell."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1101\/gad.205062.112","article-title":"Alternative lengthening of telomeres in normal mammalian somatic cells","volume":"27","author":"Neumann","year":"2013","journal-title":"Genes Dev."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1436","DOI":"10.1038\/ncb1664","article-title":"Telomere lengthening early in development","volume":"9","author":"Liu","year":"2007","journal-title":"Nat. Cell Biol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1038\/nature20099","article-title":"Break-induced telomere synthesis underlies alternative telomere maintenance","volume":"539","author":"Dilley","year":"2016","journal-title":"Nature"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"2907","DOI":"10.15252\/embj.201796889","article-title":"BLM and SLX4 play opposing roles in recombination-dependent replication at human telomeres","volume":"36","author":"Sobinoff","year":"2017","journal-title":"EMBO J."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1816","DOI":"10.1016\/j.molcel.2021.02.004","article-title":"A unified alternative telomere-lengthening pathway in yeast survivor cells","volume":"81","author":"Kockler","year":"2021","journal-title":"Mol. Cell"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1392","DOI":"10.1101\/gad.328575.119","article-title":"Break-induced replication promotes fragile telomere formation","volume":"34","author":"Yang","year":"2020","journal-title":"Genes Dev."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"777","DOI":"10.1007\/s00404-014-3281-x","article-title":"The impact of cigarette smoking and alcohol consumption on sperm parameters and sperm DNA fragmentation (SDF) measured by Halosperm(\u00ae)","volume":"290","author":"Anifandis","year":"2014","journal-title":"Arch. Gynecol. Obstet."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Anifandis, G., Amiridis, G., Dafopoulos, K., Daponte, A., Dovolou, E., Gavriil, E., Gorgogietas, V., Kachpani, E., Mamuris, Z., and Messini, C.I. (2017). The In Vitro Impact of the Herbicide Roundup on Human Sperm Motility and Sperm Mitochondria. Toxics, 6.","DOI":"10.3390\/toxics6010002"},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Anifandis, G., Katsanaki, K., Lagodonti, G., Messini, C., Simopouloum, M., Dafopoulosm, K., and Daponte, A. (2018). The Effect of Glyphosate on Human Sperm Motility and Sperm DNA Fragmentation. Int. J. Env. Res. Public Health, 15.","DOI":"10.3390\/ijerph15061117"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.etap.2017.02.024","article-title":"Human exposure to endocrine disrupting chemicals: Effects on the male and female reproductive systems","volume":"51","author":"Sifakis","year":"2017","journal-title":"Env. Toxicol. Pharmacol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1016\/j.taap.2018.03.025","article-title":"Association of pesticide exposure with human congenital abnormalities","volume":"346","author":"Kalliora","year":"2018","journal-title":"Toxicol. Appl. Pharmacol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1111\/j.2047-2927.2012.00012.x","article-title":"The BMI of men and not sperm parameters impact on embryo quality and the IVF outcome","volume":"1","author":"Anifandis","year":"2013","journal-title":"Andrology"},{"key":"ref_36","first-page":"3701","article-title":"Discovery of potent telomerase activators: Unfolding new therapeutic and anti-aging perspectives","volume":"20","author":"Tsoukalas","year":"2019","journal-title":"Mol. Med. Rep."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"32","DOI":"10.2174\/1389202916666141224205025","article-title":"Genes and Conditions Controlling Mammalian Pre- and Post-implantation Embryo Development","volume":"16","author":"Anifandis","year":"2015","journal-title":"Curr. Genom."},{"key":"ref_38","first-page":"451","article-title":"Considerations Regarding Embryo Culture Conditions: From Media to Epigenetics","volume":"32","author":"Simopoulou","year":"2018","journal-title":"In Vivo"},{"key":"ref_39","first-page":"305","article-title":"Telomere length: Lights and shadows on their role in human reproduction","volume":"100","author":"Rocca","year":"2019","journal-title":"Biol. Reprod."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"475","DOI":"10.3390\/genes12040475","article-title":"TERRA: A Novel Biomarker of Embryo Quality and Art Outcome","volume":"12","author":"Rocca","year":"2021","journal-title":"Genes"},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Minamoto, T., Nakayama, K., Ishibashi, T., Ishikawa, M., Nakamura, K., Yamashita, H., Shanta, K., Mahmud, H.M., Razia, S., and Iida, K. (2020). Pregnancy by Assisted Reproductive Technology Is Associated with Shorter Telomere Length in Neonates. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21249688"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"358","DOI":"10.3389\/fgene.2015.00358","article-title":"Insertion of Retrotransposons at Chromosome Ends: Adaptive Response to Chromosome Maintenance","volume":"6","author":"Servant","year":"2016","journal-title":"Front. Genet."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1221","DOI":"10.1093\/hmg\/ddp022","article-title":"Retrotransposon RNA expression and evidence for retrotransposition events in human oocytes","volume":"18","author":"Georgiou","year":"2009","journal-title":"Hum. Mol. Genet."},{"key":"ref_44","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_45","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1007\/s10577-005-0993-6","article-title":"Two retrotransposons maintain telomeres in Drosophila","volume":"13","author":"Pardue","year":"2005","journal-title":"Chromosome Res."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"2269","DOI":"10.21873\/invivo.12037","article-title":"Reverse Transcriptase Affects Gametogenesis and Preimplantation Development in Mouse","volume":"34","author":"Kitsou","year":"2020","journal-title":"Vivo"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1921","DOI":"10.21873\/invivo.12458","article-title":"Expression of Retroelements in Mammalian Gametes and Embryos","volume":"35","author":"Mastora","year":"2021","journal-title":"Vivo"},{"key":"ref_48","first-page":"375","article-title":"The association of female and male infertility with telomere length (Review)","volume":"44","author":"Vasilopoulos","year":"2019","journal-title":"Int. J. Mol. Med."},{"key":"ref_49","first-page":"805","article-title":"Expression of the RNA component of telomerase during human development and differentiation","volume":"9","author":"Yashima","year":"1998","journal-title":"Cell Growth Differ."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"380","DOI":"10.1016\/j.maturitas.2013.05.001","article-title":"Age-dependence of relative telomere length profiles during spermatogenesis in man","volume":"75","author":"Fedder","year":"2013","journal-title":"Maturitas"},{"key":"ref_51","first-page":"103","article-title":"Telomeric repeat-containing RNA (TERRA) and telomerase are components of telomeres during mammalian gametogenesis","volume":"90","author":"Vitelli","year":"2014","journal-title":"Biol. Reprod."},{"key":"ref_52","first-page":"20190800","article-title":"Older paternal ages and grandpaternal ages at conception predict longer telomeres in human descendants","volume":"286","author":"Eisenberg","year":"2019","journal-title":"Proc. Biol. Sci."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"10251","DOI":"10.1073\/pnas.1202092109","article-title":"Delayed paternal age of reproduction in humans is associated with longer telomeres across two generations of descendants","volume":"109","author":"Eisenberg","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1158","DOI":"10.1093\/humrep\/dew061","article-title":"Sperm telomere length as a parameter of sperm quality in normozoospermic men","volume":"31","author":"Rocca","year":"2016","journal-title":"Hum. Reprod."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"404","DOI":"10.1016\/j.rbmo.2016.06.006","article-title":"Investigation of sperm telomere length as a potential marker of paternal genome integrity and semen quality","volume":"33","author":"Cariati","year":"2016","journal-title":"Reprod. Biomed. Online"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1686","DOI":"10.1091\/mbc.e05-07-0672","article-title":"PARP1 Is a TRF2-associated poly(ADP-ribose)polymerase and protects eroded telomeres","volume":"17","author":"Gomez","year":"2006","journal-title":"Mol. Biol. Cell"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1016\/j.rbmo.2018.12.022","article-title":"Evaluation of sperm telomere length in infertile men with failed\/low fertilization after intracytoplasmic sperm injection","volume":"38","author":"Darmishonnejad","year":"2019","journal-title":"Reprod. Biomed. Online"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"2023","DOI":"10.1091\/mbc.12.7.2023","article-title":"Telomere dysfunction triggers developmentally regulated germ cell apoptosis","volume":"12","author":"Hemann","year":"2001","journal-title":"Mol. Biol. Cell"},{"key":"ref_59","first-page":"493","article-title":"Role of sperm apoptosis and oxidative stress in male infertility: A narrative review","volume":"19","author":"Asadi","year":"2021","journal-title":"Int. J. Reprod. Biomed."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"39051","DOI":"10.1038\/srep39051","article-title":"Semen preparation methods and sperm telomere length: Density gradient centrifugation versus the swim up procedure","volume":"6","author":"Zhao","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"163","DOI":"10.5662\/wjm.v6.i2.163","article-title":"Mild oxidative stress is beneficial for sperm telomere length maintenance","volume":"6","author":"Mishra","year":"2016","journal-title":"World J. Methodol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1016\/j.gene.2013.11.008","article-title":"Genetic variants in telomerase reverse transcriptase (TERT) and telomerase-associated protein 1 (TEP1) and the risk of male infertility","volume":"534","author":"Yan","year":"2014","journal-title":"Gene"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1055\/s-0035-1567823","article-title":"Telomeres and Female Reproductive Aging","volume":"33","author":"Kalmbach","year":"2015","journal-title":"Semin. Reprod. Med."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"280","DOI":"10.1097\/01.gco.0000193019.05686.49","article-title":"The telomere theory of reproductive senescence in women","volume":"18","author":"Keefe","year":"2006","journal-title":"Curr. Opin. Obstet. Gynecol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"542","DOI":"10.1016\/j.fertnstert.2014.10.035","article-title":"Inflammatory biomarkers and telomere length in women with polycystic ovary syndrome","volume":"103","author":"Pedroso","year":"2015","journal-title":"Fertil. Steril."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"861","DOI":"10.1007\/s10815-017-0945-z","article-title":"Significantly lengthened telomere in granulosa cells from women with polycystic ovarian syndrome (PCOS)","volume":"34","author":"Wei","year":"2017","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"1697","DOI":"10.1007\/s10815-015-0581-4","article-title":"Cell-free DNA and telomere length among women undergoing in vitro fertilization treatment","volume":"32","author":"Sarid","year":"2015","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"736","DOI":"10.1093\/humrep\/dey024","article-title":"Parity associated with telomere length among US reproductive age women","volume":"33","author":"Pollack","year":"2018","journal-title":"Hum. Reprod."},{"key":"ref_69","first-page":"201","article-title":"Impaired telomere length and telomerase activity in peripheral blood leukocytes and granulosa cells in patients with biochemical primary ovarian insufficiency","volume":"32","author":"Xu","year":"2017","journal-title":"Hum. Reprod."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"16","DOI":"10.3892\/mmr.2016.5231","article-title":"Telomerase activity in pregnancy complications (Review)","volume":"14","author":"Fragkiadaki","year":"2016","journal-title":"Mol. Med. Rep."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1016\/j.mad.2017.12.002","article-title":"DNA methylation dynamics in aging: How far are we from understanding the mechanisms?","volume":"174","author":"Ciccarone","year":"2018","journal-title":"Mech. Ageing Dev."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"4954","DOI":"10.1038\/srep04954","article-title":"Human leukocyte telomere length is associated with DNA methylation levels in multiple subtelomeric and imprinted loci","volume":"4","author":"Buxton","year":"2014","journal-title":"Sci. Rep."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"14630","DOI":"10.18632\/aging.203117","article-title":"Different epigenetic signatures of newborn telomere length and telomere attrition rate in early life","volume":"13","author":"Wang","year":"2021","journal-title":"Aging"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1002\/(SICI)1520-6408(1996)18:2<173::AID-DVG10>3.0.CO;2-3","article-title":"Telomerase activity in human germline and embryonic tissues and cells","volume":"18","author":"Wright","year":"1996","journal-title":"Dev Genet."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1530\/REP-15-0375","article-title":"Spermatozoa telomeres determine telomere length in early embryos and offspring","volume":"151","author":"Rizos","year":"2016","journal-title":"Reproduction"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"638","DOI":"10.1126\/science.aah6752","article-title":"TZAP: A telomere-associated protein involved in telomere length control","volume":"355","author":"Li","year":"2017","journal-title":"Science"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"685","DOI":"10.1093\/molehr\/gaq048","article-title":"Telomere lengths in human oocytes, cleavage stage embryos and blastocysts","volume":"16","author":"Turner","year":"2010","journal-title":"Mol. Hum. Reprod."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"399","DOI":"10.1002\/mrd.23336","article-title":"Listening to mother: Long-term maternal effects in mammalian development","volume":"87","author":"Ruebel","year":"2020","journal-title":"Mol. Reprod. Dev."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"866","DOI":"10.1093\/humupd\/dmab010","article-title":"Maternal germline factors associated with aneuploid pregnancy loss: A systematic review","volume":"27","author":"Blyth","year":"2021","journal-title":"Hum. Reprod. Update"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"318","DOI":"10.1093\/biolre\/ioy215","article-title":"Shedding light into the relevance of telomeres in human reproduction and male factor infertility","volume":"100","author":"Lopes","year":"2019","journal-title":"Biol. Reprod."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"8034","DOI":"10.1073\/pnas.0402400101","article-title":"Telomere length is reset during early mammalian embryogenesis","volume":"101","author":"Schaetzlein","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1016\/j.theriogenology.2020.09.019","article-title":"Evaluation of bovine sperm telomere length and association with semen quality","volume":"158","author":"Iannuzzi","year":"2020","journal-title":"Theriogenology"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1016\/j.rbmo.2015.02.016","article-title":"Processing of semen by density gradient centrifugation selects spermatozoa with longer telomeres for assisted reproduction techniques","volume":"31","author":"Yang","year":"2015","journal-title":"Reprod. Biomed. Online"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"e13546","DOI":"10.1111\/and.13546","article-title":"Relationship between sperm telomere length and sperm quality in infertile men","volume":"52","author":"Darmishonnejad","year":"2020","journal-title":"Andrologia"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"1876","DOI":"10.1093\/humrep\/dev144","article-title":"Sperm telomere length is positively associated with the quality of early embryonic development","volume":"30","author":"Yang","year":"2015","journal-title":"Hum. Reprod."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1111\/andr.12734","article-title":"Associations of sperm telomere length with semen parameters, clinical outcomes and lifestyle factors in human normozoospermic samples","volume":"8","author":"Berneau","year":"2020","journal-title":"Andrology"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"282","DOI":"10.1080\/14647273.2019.1575988","article-title":"The effect of maternal body mass index (BMI) and telomere function on in vitro fertilization (IVF) outcome: A preliminary cohort study","volume":"23","author":"Weeg","year":"2020","journal-title":"Hum. Fertil."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"649","DOI":"10.1007\/s10815-017-1104-2","article-title":"Sperm telomere length in donor samples is not related to ICSI outcome","volume":"35","author":"Bellu","year":"2018","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1007\/s00439-020-02236-1","article-title":"Disruption of human meiotic telomere complex genes TERB1, TERB2 and MAJIN in men with non-obstructive azoospermia","volume":"140","author":"Wyrwoll","year":"2021","journal-title":"Hum. Genet."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"606","DOI":"10.1016\/j.rbmo.2018.12.008","article-title":"Maternal telomere length is shorter in intrauterine growth restriction versus uncomplicated pregnancies, but not in the offspring or in IVF-conceived newborns","volume":"38","author":"Monterde","year":"2019","journal-title":"Reprod. Biomed. Online"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1080\/19485565.2019.1706150","article-title":"Biomarkers for demographic research: Sperm counts and other male infertility biomarkers","volume":"65","author":"Westerman","year":"2020","journal-title":"Biodemography Soc. Biol."}],"container-title":["Journal of Developmental Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2221-3759\/9\/4\/49\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T07:29:49Z","timestamp":1760167789000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2221-3759\/9\/4\/49"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,11,13]]},"references-count":91,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2021,12]]}},"alternative-id":["jdb9040049"],"URL":"https:\/\/doi.org\/10.3390\/jdb9040049","relation":{},"ISSN":["2221-3759"],"issn-type":[{"value":"2221-3759","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,11,13]]}}}