{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,17]],"date-time":"2026-10-17T05:02:19Z","timestamp":1792213339856,"version":"4.4.0"},"reference-count":95,"publisher":"MDPI AG","issue":"5","license":[{"start":{"date-parts":[[2023,4,25]],"date-time":"2023-04-25T00:00:00Z","timestamp":1682380800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biomolecules"],"abstract":"<jats:p>In recent decades, male infertility has been correlated with the shortening of sperm telomeres. Telomeres regulate the reproductive lifespan by mediating the synapsis and homologous recombination of chromosomes during gametogenesis. They are composed of thousands of hexanucleotide DNA repeats (TTAGGG) that are coupled to specialized shelterin complex proteins and non-coding RNAs. Telomerase activity in male germ cells ensures that the telomere length is maintained at maximum levels during spermatogenesis, despite telomere shortening due to DNA replication or other genotoxic factors such as environmental pollutants. An emerging body of evidence has associated an exposure to pollutants with male infertility. Although telomeric DNA may be one of the important targets of environmental pollutants, only a few authors have considered it as a conventional parameter for sperm function. The aim of this review is to provide comprehensive and up-to-date data on the research carried out so far on the structure\/function of telomeres in spermatogenesis and the influence of environmental pollutants on their functionality. The link between pollutant-induced oxidative stress and telomere length in germ cells is discussed.<\/jats:p>","DOI":"10.3390\/biom13050745","type":"journal-article","created":{"date-parts":[[2023,4,26]],"date-time":"2023-04-26T02:30:52Z","timestamp":1682476252000},"page":"745","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":7,"title":["Male Germ Cell Telomeres and Chemical Pollutants"],"prefix":"10.3390","volume":"13","author":[{"given":"Gabriella","family":"Chieffi Baccari","sequence":"first","affiliation":[{"name":"Dipartimento di Scienze e Tecnologie Ambientali, Biologiche e Farmaceutiche, Universit\u00e0 della Campania \u201cLuigi Vanvitelli\u201d, 81100 Caserta, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Giuseppe","family":"Iurato","sequence":"additional","affiliation":[{"name":"Ministero dell\u2019Istruzione e del Merito, 00153 Roma, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alessandra","family":"Santillo","sequence":"additional","affiliation":[{"name":"Dipartimento di Scienze e Tecnologie Ambientali, Biologiche e Farmaceutiche, Universit\u00e0 della Campania \u201cLuigi Vanvitelli\u201d, 81100 Caserta, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Brian","family":"Dale","sequence":"additional","affiliation":[{"name":"Centro Fecondazione Assistita (CFA-Italia), 80127 Napoli, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2023,4,25]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"597","DOI":"10.1038\/s41585-022-00626-w","article-title":"Spatiotemporal trends in human semen quality","volume":"19","author":"Ager","year":"2022","journal-title":"Nat. Rev. Urol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1012119","DOI":"10.3389\/fendo.2022.1012119","article-title":"Testicular aging, male fertility and beyond","volume":"13","author":"Dong","year":"2022","journal-title":"Front. Endocrinol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1095\/biolreprod.115.131318","article-title":"Male Rat germ cells display age-dependent and cell-specific susceptibility in response to oxidative stress challenges","volume":"93","author":"Selvaratnam","year":"2015","journal-title":"Biol. Reprod."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"e13242","DOI":"10.1111\/acel.13242","article-title":"A germ cell-specific ageing pattern in otherwise healthy men","volume":"19","author":"Laurentino","year":"2020","journal-title":"Aging Cell"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"744","DOI":"10.1038\/ng1382","article-title":"BubR1 insufficiency causes early onset of aging-associated phenotypes and infertility in mice","volume":"36","author":"Baker","year":"2004","journal-title":"Nat. Genet."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"360","DOI":"10.1080\/19396368.2017.1391888","article-title":"Aging deteriorates quality of sperm produced by male mice with partial yq deletion","volume":"63","author":"Kotarska","year":"2017","journal-title":"Syst. Biol. Reprod. Med."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Cheng, Y., Vechtova, P., Fussy, Z., Sterba, J., Linhartova, Z., Rodina, M., Tu\u010dkov\u00e1, V., Gela, D., Samarin, A.M., and Lebeda, I. (2021). Changes in phenotypes and DNA methylation of In Vitro aging sperm in common carp Cyprinus carpio. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22115925"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1269","DOI":"10.1095\/biolreprod.111.094219","article-title":"Aging results in differential regulation of DNA repair pathways in pachytene spermatocytes in the brown Norway rat","volume":"85","author":"Paul","year":"2011","journal-title":"Biol. Reprod."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1053\/j.seminhematol.2013.03.030","article-title":"Telomere dynamics in mice and humans","volume":"50","author":"Calado","year":"2013","journal-title":"Semin. Hematol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"2100","DOI":"10.1101\/gad.1346005","article-title":"Shelterin: The protein complex that shapes and safeguards human telomeres","volume":"19","year":"2005","journal-title":"Genes Dev."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"188","DOI":"10.1046\/j.1432-0436.2002.690412.x","article-title":"Telomerase and differentiation in multicellular organisms: Turn it off, turn it on, and turn it off again","volume":"69","author":"Forsyth","year":"2002","journal-title":"Differentiation"},{"key":"ref_12","first-page":"1496","article-title":"The principle of marginotomy in the matrix synthesis of polynucleotides","volume":"201","author":"Olovnikov","year":"1971","journal-title":"Dokl. Akad. Nauk SSSR"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1016\/0022-5193(73)90198-7","article-title":"A Theory of marginotomy. The incomplete copying of template margin in enzymic synthesis of poly nucleotides and biological significance of the phenomenon","volume":"41","author":"Olovnikov","year":"1973","journal-title":"J. Theor. Biol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"7405","DOI":"10.1002\/anie.201002387","article-title":"Telomeres and telomerase: The means to the end (Nobel lecture)","volume":"49","author":"Blackburn","year":"2010","journal-title":"Angew. Chem. Int. Ed. Engl."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"7422","DOI":"10.1002\/anie.201002408","article-title":"Telomerase discovery: The excitement of putting together pieces of the puzzle (Nobel lecture)","volume":"49","author":"Greider","year":"2010","journal-title":"Angew. Chem. Int. Ed. Engl."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"7386","DOI":"10.1002\/anie.201000635","article-title":"DNA ends: Just the beginning (Nobel lecture)","volume":"49","author":"Szostak","year":"2010","journal-title":"Angew. Chem. Int. Ed. Engl."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1095\/biolreprod.114.124008","article-title":"Telomerase activity and telomere length in male germ cells","volume":"92","author":"Ozturk","year":"2015","journal-title":"Biol. Reprod."},{"key":"ref_18","first-page":"181","article-title":"The role of epigenetics in spermatogenesis","volume":"39","year":"2013","journal-title":"Turk. J. Urol."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Cescon, M., Chianese, R., and Tavares, R.S. (2020). Environmental Impact on Male (In)Fertility via Epigenetic Route. J. Clin. Med., 9.","DOI":"10.3390\/jcm9082520"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1016\/j.ygeno.2004.06.001","article-title":"Novel human testis-specific histone H2B encoded by the interrupted gene on the X chromosome","volume":"84","author":"Churikov","year":"2004","journal-title":"Genomics"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1111\/and.12008","article-title":"Telomere length in reproduction","volume":"45","author":"Thilagavathi","year":"2013","journal-title":"Andrologia"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"728","DOI":"10.1016\/j.fertnstert.2014.06.005","article-title":"Telomere homeostasis is compromised in spermatocytes from patients with idiopathic infertility","volume":"102","author":"Capilla","year":"2014","journal-title":"Fertil. Steril."},{"key":"ref_23","first-page":"375","article-title":"The association of female and male infertility with telomere length","volume":"44","author":"Vasilopoulos","year":"2019","journal-title":"Int. J. Mol. Med."},{"key":"ref_24","first-page":"561","article-title":"Impact of ovarian aging in reproduction: From telomeres and mice models to ovarian rejuvenation","volume":"93","author":"Polonio","year":"2020","journal-title":"Biol. Med."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1097\/GCO.0000000000000779","article-title":"Telomeres, aging and reproduction","volume":"34","author":"Varela","year":"2022","journal-title":"Curr. Opin. Obstet. Gynecol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"2420","DOI":"10.1101\/gad.271783.115","article-title":"High telomerase is a hallmark of undifferentiated spermatogonia and is required for maintenance of male germline stem cells","volume":"29","author":"Pech","year":"2015","journal-title":"Genes Dev."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1071\/RD08229","article-title":"Telomeres and reproductive aging","volume":"21","author":"Keefe","year":"2009","journal-title":"Reprod. Fertil. Dev."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Ribas-Maynou, J., Mateo-Otero, Y., Sanchez-Quijada, M., Recuero, S., Delgado-Berm\u00fadez, A., Llavanera, M., and Yeste, M. (2022). Telomere length in pig sperm is related to in vitro embryo development outcomes. Animals, 12.","DOI":"10.3390\/ani12020204"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1080\/14647273.2018.1456682","article-title":"Telomeres as a molecular marker of male infertility","volume":"22","author":"Cybulska","year":"2019","journal-title":"Hum. Fertil."},{"key":"ref_30","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_31","doi-asserted-by":"crossref","first-page":"3370","DOI":"10.1093\/humrep\/det392","article-title":"In young men sperm telomere length is related to sperm number and parental age","volume":"28","author":"Ferlin","year":"2013","journal-title":"Hum. Reprod."},{"key":"ref_32","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":"2013","journal-title":"Hum. Reprod."},{"key":"ref_33","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_34","doi-asserted-by":"crossref","first-page":"706","DOI":"10.1016\/S0015-0282(99)00603-2","article-title":"Telomerase activity and expression of telomerase subunits in the testicular tissue of infertile patients","volume":"73","author":"Schrader","year":"2000","journal-title":"Fertil. Steril."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1007\/s00412-015-0555-4","article-title":"Telomere homeostasis in mammalian germ cells: A review","volume":"125","year":"2016","journal-title":"Chromosoma"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"F125","DOI":"10.1530\/REP-22-0189","article-title":"Oxidative stress and reproductive function: Sperm telomeres, oxidative stress, and infertility","volume":"164","author":"Moazamian","year":"2022","journal-title":"Reproduction"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Fattet, A.J., Chaillot, M., and Koscinski, I. (2023). Telomere Length, a New Biomarker of Male (in)Fertility? A Systematic Review of the Literature. Genes, 14.","DOI":"10.3390\/genes14020425"},{"key":"ref_38","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. Repr."},{"key":"ref_39","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":"Barragan","year":"2018","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"2581","DOI":"10.1007\/s10815-020-01897-1","article-title":"Discordance between human sperm quality and telomere length following differential gradient separation\/swim-up","volume":"37","author":"Lopes","year":"2020","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Gentiluomo, M., Luddi, A., Cingolani, A., Fornili, M., Governini, L., Lucenteforte, E., Baglietto, L., Piomboni, P., and Campa, D. (2021). Telomere Length and Male Fertility. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22083959"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1079966","DOI":"10.3389\/fendo.2022.1079966","article-title":"Sperm telomere length as a novel biomarker of male infertility and embryonic development: A systematic review and meta-analysis","volume":"13","author":"Yuan","year":"2023","journal-title":"Front. Endocrinol."},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Garanina, A.S., Alieva, I.B., Bragina, E.E., Blanchard, E., Arbeille, B., Guerif, F., Uzbekova, S., and Uzbekov, R.E. (2019). The Centriolar Adjunct\u2014Appearance and Disassembly in Spermiogenesis and the Potential Impact on Fertility. Cells, 8.","DOI":"10.3390\/cells8020180"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"636","DOI":"10.1016\/j.envpol.2018.09.130","article-title":"Air pollution and telomere length in adults: A systematic review and meta-analysis of observational studies","volume":"244","author":"Miri","year":"2019","journal-title":"Environ. Pollut."},{"key":"ref_45","first-page":"377","article-title":"Environmental exposure to non-persistent endocrine disrupting chemicals and semen quality: An overview of the current epidemiological evidence","volume":"31","author":"Zamkowska","year":"2018","journal-title":"Int. J. Occup. Med. Environ. Health"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"5957","DOI":"10.1021\/acs.est.8b06942","article-title":"Association of exposure to ambient fine particulate matter constituents with semen quality among men attending a fertility center in China","volume":"53","author":"Huang","year":"2019","journal-title":"Environ. Sci. Technol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"4095","DOI":"10.1007\/s11356-020-11589-z","article-title":"Pollutants and sperm quality: A systematic review and meta-analysis","volume":"28","author":"Pizzol","year":"2021","journal-title":"Environ. Sci. Pollut. Res. Int."},{"key":"ref_48","first-page":"291","article-title":"Does GSTP1 polymorphism contribute to genetic damage caused by ageing and occupational exposure?","volume":"62","author":"Eshkoor","year":"2011","journal-title":"Arch. Ind. Hyg. Toxicol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1016\/j.envint.2012.06.020","article-title":"Air pollution exposure and telomere length in highly exposed subjects in Beijing, China: A repeated-measure study","volume":"48","author":"Hou","year":"2012","journal-title":"Environ. Int."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1016\/j.envint.2016.08.021","article-title":"Air pollution, ethnicity and telomere length in east London schoolchildren: An observational study","volume":"96","author":"Walton","year":"2016","journal-title":"Environ. Int."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"12806","DOI":"10.1021\/acs.est.7b03289","article-title":"Inverse association between ambient sulfur dioxide exposure and semen quality in Wuhan, China","volume":"51","author":"Liu","year":"2017","journal-title":"Environ. Sci. Technol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1136\/oemed-2017-104529","article-title":"Exposure to ambient fine particulate matter and semen quality in Taiwan","volume":"75","author":"Lao","year":"2018","journal-title":"Occup. Environ. Med."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"501","DOI":"10.4103\/aja.aja_116_18","article-title":"Ambient ozone pollution is associated with decreased semen quality: Longitudinal analysis of 8945 semen samples from 2015 to 2018 and during pollution-control period in Beijing, China","volume":"21","author":"Zhang","year":"2019","journal-title":"Asian J. Androl."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"105483","DOI":"10.1016\/j.envint.2020.105483","article-title":"Associations of ambient air pollutant exposure with seminal plasma MDA, sperm mtDNA copy number, and mtDNA integrity","volume":"136","author":"Zhang","year":"2020","journal-title":"Environ. Int."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"e2148684","DOI":"10.1001\/jamanetworkopen.2021.48684","article-title":"Association of exposure to particulate matter air pollution with semen quality among men in China","volume":"5","author":"Zhao","year":"2022","journal-title":"JAMA Netw. Open"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1146","DOI":"10.18632\/aging.101454","article-title":"Roles of three Es-Caspases during spermatogenesis and Cadmium-induced apoptosis in Eriocheir sinensis","volume":"10","author":"Xu","year":"2018","journal-title":"Aging"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"114485","DOI":"10.1016\/j.ecoenv.2022.114485","article-title":"Establishing the relationship between Polycyclic Aromatic Hydrocarbons (PAHs) exposure and male infertility: A systematic review","volume":"250","author":"Kakavandi","year":"2023","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"915","DOI":"10.13075\/ijomeh.1896.00598","article-title":"Sperm quality and DNA integrity of coke oven workers exposed to polycyclic aromatic hydrocarbons","volume":"29","author":"Jeng","year":"2016","journal-title":"Int. J. Occup. Med. Environ. Health"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"153017","DOI":"10.1016\/j.tox.2021.153017","article-title":"Environmental and occupational pesticide exposure and human sperm parameters: A Navigation Guide review","volume":"465","author":"Knapke","year":"2022","journal-title":"Toxicology"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1515\/reveh-2021-0074","article-title":"Electronic waste exposure and DNA damage: A systematic review and meta-analysis","volume":"38","author":"Issah","year":"2021","journal-title":"Rev. Environ. Health"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"641","DOI":"10.1007\/s40618-019-01152-0","article-title":"Perfluoro-octanoic acid impairs spermmotility through the alteration of plasmamembrane","volume":"43","author":"Cosci","year":"2020","journal-title":"J. Endocrinol. Investig."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"633","DOI":"10.4103\/1008-682X.153850","article-title":"Are sperm capacitation and apoptosis the opposite ends of a continuum driven by oxidative stress?","volume":"17","author":"Aitken","year":"2015","journal-title":"Asian J. Androl."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1093\/mutage\/geq024","article-title":"In vitro evaluation of baseline and induced DNA damage in human sperm exposed to benzo[a]pyrene or its metabolite benzo[a]pyrene-7,8-diol-9,10-epoxide, using the comet assay","volume":"25","author":"Sipinen","year":"2010","journal-title":"Mutagenesis"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1124\/mol.108.053645","article-title":"The spermastatic and microbicidal actions of quinones and maleimides: Toward a dual-purpose contraceptive agent","volume":"76","author":"Hughes","year":"2009","journal-title":"Mol. Pharmacol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"124308","DOI":"10.1016\/j.jhazmat.2020.124308","article-title":"Sperm mtDNA copy number, telomere length, and seminal spermatogenic cells in relation to ambient air pollution: Results of a cross-sectional study in Jing-Jin-Ji region of China","volume":"406","author":"Zhou","year":"2021","journal-title":"J. Hazard. Mater."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Vecoli, C., Montano, L., Borghini, A., Notari, T., Guglielmino, A., Mercuri, A., Turchi, S., and Andreassi, M.G. (2017). Effects of Highly Polluted Environment on Sperm Telomere Length: A Pilot Study. Int. J. Mol. Sci., 18.","DOI":"10.3390\/ijms18081703"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"114737","DOI":"10.1016\/j.envres.2022.114737","article-title":"Blood trihalomethane concentrations in relation to sperm mitochondrial DNA copy number and telomere length among 958 healthy men","volume":"216","author":"Liu","year":"2023","journal-title":"Environ. Res."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1016\/j.envint.2016.08.001","article-title":"Shorter sperm telomere length in association with exposure to polycyclic aromatic hydrocarbons: Results from the MARHCS cohort study in Chongqing, China and in vivo animal experiments","volume":"95","author":"Ling","year":"2016","journal-title":"Environ. Int."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"836","DOI":"10.1016\/j.envpol.2017.12.099","article-title":"TERT regulates telomere-related senescence and apoptosis through DNA damage response in male germ cells exposed to BPDE in vitro and to B[a]P in vivo","volume":"235","author":"Ling","year":"2018","journal-title":"Environ. Pollut."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"106307","DOI":"10.1016\/j.envint.2020.106307","article-title":"BDE-209 and DBDPE induce male reproductive toxicity through telomere-related cell senescence and apoptosis in SD rat","volume":"146","author":"Li","year":"2021","journal-title":"Environ. Int."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"e201900374","DOI":"10.26508\/lsa.201900374","article-title":"ROS amplification drives mouse spermatogonial stem cell selfrenewal","volume":"2","author":"Morimoto","year":"2019","journal-title":"Life Sci. Alliance"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"250","DOI":"10.1101\/gad.339903.120","article-title":"An interplay of NOX1-derived ROS and oxygen determines the spermatogonial stem cell self-renewal efficiency under hypoxia","volume":"35","author":"Morimoto","year":"2021","journal-title":"Genes Dev."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1038\/s41421-020-0183-x","article-title":"A bioenergetic shift is required for spermatogonial differentiation","volume":"6","author":"Chen","year":"2020","journal-title":"Cell Discov."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1321","DOI":"10.1089\/scd.2009.0313","article-title":"Effects of oxidative stress on mouse embryonic stem cell proliferation, apoptosis, senescence, and self-renewal","volume":"19","author":"Guo","year":"2010","journal-title":"Stem Cells Dev."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"517","DOI":"10.1095\/biolreprod.109.076836","article-title":"DNA damage in human spermatozoa is highly correlated with the efficiency of chromatin remodeling and the formation of 8-hydroxy-2\u2032-deoxyguanosine, a marker of oxidative stress","volume":"81","author":"Thomson","year":"2009","journal-title":"Biol. Reprod."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1071\/RD15325","article-title":"Causes and consequences of oxidative stress in spermatozoa","volume":"28","author":"Aitken","year":"2016","journal-title":"Reprod. Fertil. Dev."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"e13012","DOI":"10.1111\/and.13012","article-title":"Reactive oxygen species impact on sperm DNA and its role in male infertility","volume":"50","author":"Bui","year":"2018","journal-title":"Andrologia"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1016\/j.mad.2018.03.013","article-title":"The impact of oxidative DNA damage and stress on telomere homeostasis","volume":"177","author":"Barnes","year":"2019","journal-title":"Mech. Ageing Dev."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"246","DOI":"10.1016\/0168-9525(93)90089-Z","article-title":"Mutagenesis by 8-oxoguanine: An enemy within","volume":"9","author":"Grollman","year":"1993","journal-title":"Trends Genet."},{"key":"ref_80","doi-asserted-by":"crossref","unstructured":"Coluzzi, E., Leone, S., and Sgura, A. (2019). Oxidative stress induces telomere dysfunction and senescence by replication fork arrest. Cells, 8.","DOI":"10.3390\/cells8010019"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1016\/S0968-0004(02)02110-2","article-title":"Oxidative stress shortens telomeres","volume":"27","year":"2002","journal-title":"Trends Biochem. Sci."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1146\/annurev.genet.41.110306.130350","article-title":"How shelterin protects mammalian telomeres","volume":"42","author":"Palm","year":"2008","journal-title":"Annu. Rev. Genet."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"1659","DOI":"10.1128\/MCB.20.5.1659-1668.2000","article-title":"Control of human telomere length by TRF1 and TRF2","volume":"20","author":"Smogorzewska","year":"2000","journal-title":"Mol. Cell Biol."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"1230","DOI":"10.1093\/nar\/gki273","article-title":"Oxidative damage in telomeric DNA disrupts recognition by TRF1 and TRF2","volume":"33","author":"Opresko","year":"2005","journal-title":"Nucleic Acids Res."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1016\/j.diff.2017.12.002","article-title":"Impact of oxidative stress on telomere biology","volume":"99","author":"Ahmed","year":"2018","journal-title":"Differentiation"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"3107","DOI":"10.1016\/j.celrep.2016.11.071","article-title":"Peroxiredoxin 1 protects telomeres from oxidative damage and preserves telomeric DNA for extension by telomerase","volume":"17","author":"Abey","year":"2016","journal-title":"Cell Rep."},{"key":"ref_87","first-page":"1488","article-title":"The presence of a truncated base excision repair pathway in human spermatozoa that is mediated by OGG1","volume":"126","author":"Smith","year":"2013","journal-title":"J. Cell Sci."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"1518","DOI":"10.1016\/j.fertnstert.2014.10.020","article-title":"Male infertility testing: Reactive oxygen species and antioxidant capacity","volume":"102","author":"Ko","year":"2014","journal-title":"Fertil. Steril."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"88","DOI":"10.5653\/cerm.2018.45.2.88","article-title":"Seminal reactive oxygen species and total antioxidant capacity: Correlations with sperm parameters and impact on male infertility","volume":"45","author":"Subramanian","year":"2018","journal-title":"Clin. Exp. Reprod. Med."},{"key":"ref_90","doi-asserted-by":"crossref","unstructured":"Berby, B., Bichara, C., Rives-Feraille, A., Jumeau, F., Di Pizio, P.D., Setif, V., Sibert, L., Dumont, L., Rondanino, C., and Rives, N. (2021). Oxidative stress is associated with telomere interaction impairment and chromatin defect in spermatozoa of infertile males. Antioxidants, 10.","DOI":"10.3390\/antiox10040593"},{"key":"ref_91","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_92","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1016\/j.aju.2017.11.001","article-title":"Role of reactive oxygen species in male infertility: An updated review of literature","volume":"16","author":"Wagner","year":"2018","journal-title":"Arab. J. Urol."},{"key":"ref_93","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_94","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1111\/j.1439-0272.2009.01032.x","article-title":"Impact of body mass index on seminal oxidative stress","volume":"43","author":"Tunc","year":"2011","journal-title":"Andrologia"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"45457","DOI":"10.1038\/srep45457","article-title":"Study on the short-term effects of increased alcohol and cigarette consumption in healthy young men\u2019s seminal quality","volume":"7","author":"Silva","year":"2017","journal-title":"Sci. Rep."}],"container-title":["Biomolecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2218-273X\/13\/5\/745\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T19:23:27Z","timestamp":1760124207000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2218-273X\/13\/5\/745"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,4,25]]},"references-count":95,"journal-issue":{"issue":"5","published-online":{"date-parts":[[2023,5]]}},"alternative-id":["biom13050745"],"URL":"https:\/\/doi.org\/10.3390\/biom13050745","relation":{},"ISSN":["2218-273X"],"issn-type":[{"value":"2218-273X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,4,25]]}}}