{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T17:11:05Z","timestamp":1790097065590,"version":"4.0.1"},"reference-count":95,"publisher":"MDPI AG","issue":"18","license":[{"start":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T00:00:00Z","timestamp":1789948800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100007628","name":"Uludag University Scientific Research Projects Unit, Bursa, Turkey","doi-asserted-by":"publisher","award":["\u00d6NAP (TOA-609) 2021\/11"],"award-info":[{"award-number":["\u00d6NAP (TOA-609) 2021\/11"]}],"id":[{"id":"10.13039\/501100007628","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100007628","name":"Uludag University Scientific Research Projects Unit, Bursa, Turkey","doi-asserted-by":"publisher","award":["GAP III (TGA3) 2025\/2590"],"award-info":[{"award-number":["GAP III (TGA3) 2025\/2590"]}],"id":[{"id":"10.13039\/501100007628","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Int. J. Mol. Sci.","IJMS"],"abstract":"<jats:p>This study aimed to evaluate the effects of nicotinic acid (NA) and folic acid (FA) as antioxidants on post-thaw spermatological characteristics and OGG1\/ROMO1 gene expression and methylation. Collected ram semen was pooled and divided into four equal aliquots, diluted with different dose groups: Control, 10 mM NA with no FA (NA10), 10 mM NA + 50 nM FA (NA10FA50) and 20 mM NA + 50 nM FA (NA20FA50). Although not statistically significant (p &gt; 0.05), the NA10FA50 (53.34%) group showed the highest numerical mean motility, whereas the control group (43.44%) showed the lowest numerical mean motility. There were no statistical differences in terms of plasma membrane and DNA integrity, malondialdehyde and total antioxidant capacity rates. The lowest acrosomal damage was obtained in the NA10FA50 group (36.50%), which was only statistically different from the control (48.00%) (p &lt; 0.05). The lowest mitochondrial membrane potential was observed in the control group (13.80%) (p &lt; 0.05). NA10FA50 and NA20FA50 altered OGG1 gene expression, which was accompanied by corresponding changes in DNA methylation, suggesting a potential epigenetic regulatory relationship. ROMO1 expression was significantly higher in the NA10 compared to the NA10FA50 (p &lt; 0.01), accompanied by differences in DNA methylation patterns (p &lt; 0.05). This study suggests that the addition of 10 mM NA or a combination with 50 nM FA showed beneficial effects on the post-thaw quality of ram semen, particularly with respect to acrosomal integrity and mitochondrial membrane potential and selected molecular markers.<\/jats:p>","DOI":"10.3390\/ijms27188422","type":"journal-article","created":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T13:32:12Z","timestamp":1790083932000},"page":"8422","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Nicotinic and Folic Acid Supplementation Improves Cryopreserved Ram Sperm Quality, Potentially Through Modulation of OGG1 and ROMO1 Expression and DNA Methylation"],"prefix":"10.3390","volume":"27","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-6050-791X","authenticated-orcid":false,"given":"Mehmet Melih","family":"Y\u0131lmaz","sequence":"first","affiliation":[{"name":"Department of Reproduction and Artificial Insemination, Faculty of Veterinary Medicine, Bursa Uludag University, 16059 Bursa, T\u00fcrkiye"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5280-9360","authenticated-orcid":false,"given":"Tugaycan","family":"Cimen","sequence":"additional","affiliation":[{"name":"Department of Reproduction and Artificial Insemination, Faculty of Veterinary Medicine, Bursa Uludag University, 16059 Bursa, T\u00fcrkiye"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2758-5945","authenticated-orcid":false,"given":"Sena","family":"Ardicli","sequence":"additional","affiliation":[{"name":"Swiss Institute of Allergy and Asthma Research (SIAF), University of Zurich, CH-7265 Davos, Switzerland"},{"name":"Department of Genetics, Faculty of Veterinary Medicine, Bursa Uludag University, 16059 Bursa, T\u00fcrkiye"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"software","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4125-6841","authenticated-orcid":false,"given":"Oguzhan","family":"Huraydin","sequence":"additional","affiliation":[{"name":"Department of Reproduction and Artificial Insemination, Faculty of Veterinary Medicine, Bursa Uludag University, 16059 Bursa, T\u00fcrkiye"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7678-3289","authenticated-orcid":false,"given":"Elif","family":"Gokce","sequence":"additional","affiliation":[{"name":"Department of Reproduction and Artificial Insemination, Faculty of Veterinary Medicine, Nam\u0131k Kemal University, 59030 Tekirdag, T\u00fcrkiye"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1884-1874","authenticated-orcid":false,"given":"Deniz","family":"Karakci","sequence":"additional","affiliation":[{"name":"Department of Biochemistry, Faculty of Veterinary Medicine, Nam\u0131k Kemal University, 59030 Tekirdag, T\u00fcrkiye"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2975-2594","authenticated-orcid":false,"given":"Ahmet","family":"Aktar","sequence":"additional","affiliation":[{"name":"Department of Reproduction and Artificial Insemination, Faculty of Veterinary Medicine, Bursa Uludag University, 16059 Bursa, T\u00fcrkiye"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"resources","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1438-221X","authenticated-orcid":false,"given":"Zekariya","family":"Nur","sequence":"additional","affiliation":[{"name":"Department of Reproduction and Artificial Insemination, Faculty of Veterinary Medicine, Bursa Uludag University, 16059 Bursa, T\u00fcrkiye"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"validation","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7707-2705","authenticated-orcid":false,"given":"Muhammed","family":"Duman","sequence":"additional","affiliation":[{"name":"Department of Aquatic Animal Diseases, Faculty of Veterinary Medicine, Bursa Uludag University, 16059 Bursa, T\u00fcrkiye"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"project-administration","vocabulary":"credit"}]},{"given":"Burcu","family":"Ustuner","sequence":"additional","affiliation":[{"name":"Department of Reproduction and Artificial Insemination, Faculty of Veterinary Medicine, Bursa Uludag University, 16059 Bursa, T\u00fcrkiye"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,21]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/S0093-691X(99)00239-3","article-title":"Fundamental Aspects of Sperm Cryobiology: The Importance of Species and Individual Differences","volume":"53","author":"Holt","year":"2000","journal-title":"Theriogenology"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1267","DOI":"10.1016\/j.theriogenology.2009.12.007","article-title":"Effects of Different Cryoprotective Agents on Ram Sperm Morphology and DNA Integrity","volume":"73","author":"Nur","year":"2010","journal-title":"Theriogenology"},{"key":"ref_3","first-page":"393","article-title":"Effect of Egg Yolk and Soybean Lecithin on Tris-Based Extender in Post-Thaw Ram Semen Quality and in Vitro Fertility","volume":"20","author":"Ustuner","year":"2014","journal-title":"Kafkas Univ. Vet. Fak. Derg."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1016\/j.cryobiol.2017.07.005","article-title":"Effect of Lecithin Nanoliposome or Soybean Lecithin Supplemented by Pomegranate Extract on Post-Thaw Flow Cytometric, Microscopic and Oxidative Parameters in Ram Semen","volume":"78","author":"Mehdipour","year":"2017","journal-title":"Cryobiology"},{"key":"ref_5","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_6","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1016\/j.fertnstert.2015.11.037","article-title":"Sperm DNA Fragmentation and Mitochondrial Membrane Potential Combined Are Better for Predicting Natural Conception than Standard Sperm Parameters","volume":"105","author":"Golob","year":"2016","journal-title":"Fertil. Steril."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1007\/s10561-016-9566-5","article-title":"The Role of Antioxidants in Sperm Freezing: A Review","volume":"17","author":"Amidi","year":"2016","journal-title":"Cell Tissue Bank."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1016\/j.anireprosci.2018.09.019","article-title":"Supplementation of Extender with Coenzyme Q10 Improves the Function and Fertility Potential of Rooster Spermatozoa after Cryopreservation","volume":"198","author":"Masoudi","year":"2018","journal-title":"Anim. Reprod. Sci."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"62","DOI":"10.1016\/j.cryobiol.2019.10.198","article-title":"Effects of Supplementation of Tris-Egg Yolk Extender with Different Sugars and Antioxidants on Freezability of Ram Semen","volume":"92","author":"Rostami","year":"2020","journal-title":"Cryobiology"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1016\/j.cryobiol.2020.06.010","article-title":"Caffeic Acid Improves Microscopic Sperm Parameters and Antioxidant Status of Buffalo (Bubalus Bubalis) Bull Semen Following Freeze-Thawing Process","volume":"95","author":"Soleimanzadeh","year":"2020","journal-title":"Cryobiology"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1253","DOI":"10.1185\/030079904125004402","article-title":"Raising High-Density Lipoprotein Cholesterol with Reduction of Cardiovascular Risk: The Role of Nicotinic Acid\u2014A Position Paper Developed by the European Consensus Panel on HDL-C*","volume":"20","author":"Chapman","year":"2004","journal-title":"Curr. Med. Res. Opin."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"1791","DOI":"10.1002\/vms3.1158","article-title":"The Influence of L-proline and Fulvic Acid on Oxidative Stress and Semen Quality of Buffalo Bull Semen Following Cryopreservation","volume":"9","author":"Ramazani","year":"2023","journal-title":"Vet. Med. Sci."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1016\/j.atherosclerosis.2008.04.044","article-title":"Niacin Inhibits Vascular Oxidative Stress, Redox-Sensitive Genes, and Monocyte Adhesion to Human Aortic Endothelial Cells","volume":"202","author":"Ganji","year":"2009","journal-title":"Atherosclerosis"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1741","DOI":"10.1017\/S0007114510005520","article-title":"Dietary Nicotinic Acid Supplementation Improves Hepatic Zinc Uptake and Offers Hepatoprotection against Oxidative Damage","volume":"105","author":"Tupe","year":"2011","journal-title":"Br. J. Nutr."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1251","DOI":"10.1111\/rda.13508","article-title":"Effect of Nicotinic Acid on the Plasma Membrane Function and Polyunsaturated Fatty Acids Composition during Cryopreservation in Boar Sperm","volume":"54","author":"Lee","year":"2019","journal-title":"Reprod. Domest. Anim."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"103236","DOI":"10.1016\/j.jevs.2020.103236","article-title":"Nicotinic Acid (Niacin) Supplementation in Cooling and Freezing Extenders Enhances Stallion Semen Characteristics","volume":"94","author":"Bahrami","year":"2020","journal-title":"J. Equine Vet. Sci."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Picciano, M.F., Stokstad, E.L.R., and Gregory, J.F. (1991). Folic Acid Metabolism in Health and Disease (Contemporary Issues in Clinical Nutrition Vol. 13), Wiley-Liss. FEBS Lett 291.","DOI":"10.1016\/0014-5793(91)81133-S"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1390","DOI":"10.1016\/S0891-5849(01)00543-3","article-title":"Free Radical Scavenging Behavior of Folic Acid: Evidence for Possible Antioxidant Activity","volume":"30","author":"Joshi","year":"2001","journal-title":"Free Radic. Biol. Med."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1007\/s10561-019-09775-6","article-title":"Correlation between Sperm Motility and Sperm Chromatin\/DNA Damage before and after Cryopreservation and the Effect of Folic Acid and Nicotinic Acid on Post-Thaw Sperm Quality in Normozoospermic Men","volume":"20","author":"Rarani","year":"2019","journal-title":"Cell Tissue Bank."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"15","DOI":"10.52547\/sjku.27.2.15","article-title":"Evaluation of the Effects of Nicotinic Acid and Folic Acid on Morphology, Membrane Integrity and Quality of Sperm Chromatin Concentration in Normozoospermic Men during Cryopreservation","volume":"27","author":"Hashemi","year":"2022","journal-title":"SJKU"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"62","DOI":"10.1111\/j.1365-2605.2005.00620.x","article-title":"Semen Quality in Fertile US Men in Relation to Geographical Area and Pesticide Exposure","volume":"29","author":"Swan","year":"2006","journal-title":"Int. J. Androl."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"2631","DOI":"10.1016\/j.fertnstert.2010.03.021","article-title":"Reproducibility and Reliability of Repeated Semen Analyses in Male Partners of Subfertile Couples","volume":"94","author":"Leushuis","year":"2010","journal-title":"Fertil. Steril."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1109","DOI":"10.1021\/cr960421s","article-title":"Oxidative Nucleobase Modifications Leading to Strand Scission","volume":"98","author":"Burrows","year":"1998","journal-title":"Chem. Rev."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1021\/tx0600043","article-title":"Mechanisms of Formation, Genotoxicity, and Mutation of Guanine Oxidation Products","volume":"19","author":"Neeley","year":"2006","journal-title":"Chem. Res. Toxicol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"941","DOI":"10.1038\/nature05978","article-title":"Base-Excision Repair of Oxidative DNA Damage","volume":"447","author":"David","year":"2007","journal-title":"Nature"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"481","DOI":"10.1016\/j.dnarep.2006.10.012","article-title":"Oxidative Damage to Purines in DNA: Role of Mammalian Ogg1","volume":"6","author":"Klungland","year":"2007","journal-title":"DNA Repair"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1016\/j.bbrc.2013.07.012","article-title":"Romo1 Expression Contributes to Oxidative Stress-Induced Death of Lung Epithelial Cells","volume":"439","author":"Shin","year":"2013","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"ra10","DOI":"10.1126\/scisignal.2004374","article-title":"ROMO1 Is an Essential Redox-Dependent Regulator of Mitochondrial Dynamics","volume":"7","author":"Norton","year":"2014","journal-title":"Sci. Signal."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1864","DOI":"10.1093\/humrep\/13.7.1864","article-title":"Oxidative Damage to DNA in Human Spermatozoa Does Not Preclude Pronucleus Formation at Intracytoplasmic Sperm Injection","volume":"13","author":"Twigg","year":"1998","journal-title":"Hum. Reprod."},{"key":"ref_30","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_31","doi-asserted-by":"crossref","first-page":"12053","DOI":"10.1038\/s41598-025-96413-9","article-title":"The Synergistic Activity of Fisetin on Quercetin Improves Testicular Recover in Ischemia-Reperfusion Injury in Rats","volume":"15","author":"Zolfaghari","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"14224","DOI":"10.1038\/s41598-025-97730-9","article-title":"Allicin and Hesperidin Protect Sperm Production from Environmental Toxins in Mice","volume":"15","author":"Behnejad","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1007\/s11250-025-04311-z","article-title":"Resveratrol and Antifreeze Protein Type I Show No Synergistic Benefits in Cryopreserved Ram Sperm","volume":"57","author":"Alfradique","year":"2025","journal-title":"Trop. Anim. Health Prod."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"e70172","DOI":"10.1002\/vms3.70172","article-title":"The Effect of Cholesterol-Loaded Cyclodextrin and Resveratrol Compounds on Post-Thawing Quality of Ram Semen","volume":"11","author":"Eser","year":"2025","journal-title":"Vet. Med. Sci."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1089\/bio.2022.0081","article-title":"Does Rainbow Trout Seminal Plasma-Cysteine Combination Affect the Cryo-Survivability and Post-Thaw Incubation Resilience of Ram Semen?","volume":"21","author":"Gokce","year":"2023","journal-title":"Biopreserv. Biobank."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"104593","DOI":"10.1016\/j.cryobiol.2023.104593","article-title":"Synergistic Effect of Extracellular Adenosine Triphosphate and Quercetin on Post-Thaw Quality and Fertilization Potential of Lohi Ram Sperm","volume":"113","author":"Khan","year":"2023","journal-title":"Cryobiology"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"878","DOI":"10.1002\/mrd.20686","article-title":"Impact of Cryopreservation and Reactive Oxygen Species on DNA Integrity, Lipid Peroxidation, and Functional Parameters in Ram Sperm","volume":"74","author":"Peris","year":"2007","journal-title":"Mol. Reprod. Dev."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"10","DOI":"10.3329\/bjas.v44i1.23113","article-title":"Effect of Preservation Time on the Quality of Frozen Semen in Indigenous Rams","volume":"44","author":"Mahmuda","year":"2015","journal-title":"Bang. J. Anim. Sci."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1016\/j.anireprosci.2015.11.017","article-title":"Effect of Rainbow Trout (Oncorhynchus mykiss) Seminal Plasma on the Post-Thaw Quality of Ram Semen Cryopreserved in a Soybean Lecithin-Based or Egg Yolk-Based Extender","volume":"164","author":"Ustuner","year":"2016","journal-title":"Anim. Reprod. Sci."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/j.cryobiol.2016.12.008","article-title":"Use of Antioxidants Reduce Lipid Peroxidation and Improve Quality of Crossbred Ram Sperm during Its Cryopreservation","volume":"74","author":"Banday","year":"2017","journal-title":"Cryobiology"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"36","DOI":"10.1016\/j.cryobiol.2022.12.019","article-title":"Protective Effects of Different Doses of MitoQ Separately and Combined with Trehalose on Oxidative Stress and Sperm Function of Cryopreserved Markhoz Goat Semen","volume":"110","author":"Rezaei","year":"2023","journal-title":"Cryobiology"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"10526","DOI":"10.1038\/s41598-025-95820-2","article-title":"Mitochondrial Specific Antioxidant MitoPBN Mitigates Oxidative Stress and Improves Mitochondrial Function in Cryopreserved Ram Sperm","volume":"15","author":"Mehdipour","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1016\/j.cryobiol.2016.05.001","article-title":"Cryopreservation of Ram Semen with Antioxidant Supplemented Soybean Lecithin-Based Extenders and Impacts on Incubation Resilience","volume":"72","author":"Toker","year":"2016","journal-title":"Cryobiology"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"e14547","DOI":"10.1111\/and.14249","article-title":"Effects of Alpha-lipoic Acid on Ram Semen Cryopreservation and Post-thaw Life Span","volume":"54","author":"Onder","year":"2022","journal-title":"Andrologia"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"891","DOI":"10.5187\/jast.2023.e54","article-title":"Comparison Various Level Ascorbic Acid and Lycopene Additions in Semen Diluent Enhanced Sperm Quality of Sapudi Ram","volume":"66","author":"Sigit","year":"2024","journal-title":"J. Anim. Sci. Technol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"135","DOI":"10.14710\/jitaa.44.2.135-145","article-title":"Characteristics of the Post-Thawed Balinese Bull Semen Extended in Three Different Extenders and Equilibration Times","volume":"44","author":"Amal","year":"2019","journal-title":"J. Indones. Trop. Anim. Agric."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1016\/j.anireprosci.2017.04.002","article-title":"Seminal Plasma Removal by Density-Gradient Centrifugation Is Superior for Goat Sperm Preservation Compared with Classical Sperm Washing","volume":"181","author":"Esteso","year":"2017","journal-title":"Anim. Reprod. Sci."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1111\/j.1439-0531.2010.01603.x","article-title":"Objective Sperm Motion Analysis to Assess Dairy Bull Fertility Using Computer-Aided System\u2014A Review","volume":"46","author":"Kathiravan","year":"2011","journal-title":"Reprod. Domest. Anim."},{"key":"ref_49","first-page":"153","article-title":"The factors affecting motility and kinematic values of spermatozoa in examinations by Computer Aided Semen Analysis (CASA) system","volume":"36","author":"Sonmez","year":"2022","journal-title":"F\u0131rat Univ. Sagl\u0131k Bilim. Vet. Derg."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1016\/0005-2728(90)90026-Z","article-title":"Quantification of Aerobic Energy Turnover in Epididymal Bull Spermatozoa","volume":"1015","author":"Halangk","year":"1990","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1111\/j.1365-2605.1990.tb01051.x","article-title":"Analysis of the Relationship between Reactive Oxygen Species Production and Leucocyte Infiltration in Fractions of Human Semen Separated on Percoll Gradients","volume":"13","author":"Aitken","year":"1990","journal-title":"Int. J. Androl."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1027","DOI":"10.1038\/s42003-022-03954-0","article-title":"Predicting Fertility from Sperm Motility Landscapes","volume":"5","author":"Garriga","year":"2022","journal-title":"Commun. Biol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"106697","DOI":"10.1016\/j.anireprosci.2021.106697","article-title":"Effects of Resveratrol in Bull Semen Extender on Post-Thaw Sperm Quality and Capacity for Fertilization and Embryo Development","volume":"226","author":"Mendes","year":"2021","journal-title":"Anim. Reprod. Sci."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"799","DOI":"10.1071\/RD17432","article-title":"CASA-Mot in Mammals: An Update","volume":"30","author":"Silvestre","year":"2018","journal-title":"Reprod. Fertil. Dev."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"525","DOI":"10.1093\/jn\/135.3.525","article-title":"Low Concentrations of Flavonoids Are Protective in Rat H4IIE Cells Whereas High Concentrations Cause DNA Damage and Apoptosis","volume":"135","author":"Michels","year":"2005","journal-title":"J. Nutr."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"228","DOI":"10.4161\/oxim.3.4.12858","article-title":"Exogenous Antioxidants\u2014Double-Edged Swords in Cellular Redox State: Health Beneficial Effects at Physiologic Doses versus Deleterious Effects at High Doses","volume":"3","author":"Bouayed","year":"2010","journal-title":"Oxid. Med. Cell. Longev."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1530\/jrf.0.0700219","article-title":"Development of an Assay to Assess the Functional Integrity of the Human Sperm Membrane and Its Relationship to Other Semen Characteristics","volume":"70","author":"Jeyendran","year":"1984","journal-title":"Reproduction"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1016\/j.theriogenology.2022.03.015","article-title":"Identification and Validation of Ram Sperm Proteins Associated with Cryoinjuries Caused by the Cryopreservation Process","volume":"184","author":"Jia","year":"2022","journal-title":"Theriogenology"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1111\/j.1439-0531.2011.01818.x","article-title":"Effect of Cryopreservation on Apoptotic-like Events and Its Relationship with Cryocapacitation of Buffalo (Bubalus bubalis) Sperm","volume":"47","author":"Kadirvel","year":"2012","journal-title":"Reprod. Domest. Anim."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.cryobiol.2018.10.266","article-title":"Dimethylacetamide and Trehalose for Ram Semen Cryopreservation","volume":"85","author":"Bittencourt","year":"2018","journal-title":"Cryobiology"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"107517","DOI":"10.1016\/j.anireprosci.2024.107517","article-title":"The Effects of MitoQ as a Mitochondrial-Targeted Antioxidant in a Plant-Based Extender on Buck Sperm Quality Parameters during Cryopreservation","volume":"266","author":"Hatami","year":"2024","journal-title":"Anim. Reprod. Sci."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"380","DOI":"10.1093\/humrep\/del399","article-title":"Kinetics of Occurrence of Some Features of Apoptosis during the Cryopreservation Process of Bovine Spermatozoa","volume":"22","author":"Martin","year":"2007","journal-title":"Hum. Reprod."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1089\/ars.2021.0238","article-title":"Mitochondria-Targeted Compounds to Assess and Improve Human Sperm Function","volume":"37","author":"Cristo","year":"2022","journal-title":"Antioxid. Redox Signal."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"272","DOI":"10.1111\/j.1439-0272.2011.01177.x","article-title":"Antioxidant Supplementations in Vitro Improve Rat Sperm Parameters and Enhance Antioxidant Enzyme Activities against Dimethoate-Induced Sperm Damages: Vitamin Effects on Reproductive Toxicology","volume":"44","author":"Fetoui","year":"2012","journal-title":"Andrologia"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"535","DOI":"10.2165\/11532990-000000000-00000","article-title":"Folic Acid and L-5-Methyltetrahydrofolate: Comparison of Clinical Pharmacokinetics and Pharmacodynamics","volume":"49","author":"Pietrzik","year":"2010","journal-title":"Clin. Pharmacokinet."},{"key":"ref_66","first-page":"52","article-title":"Folic Acid Supplementation and Pregnancy: More than Just Neural Tube Defect Prevention","volume":"4","author":"Greenberg","year":"2011","journal-title":"Rev. Obstet. Gynecol."},{"key":"ref_67","first-page":"1488","article-title":"The Presence of a Truncated Base Excision Repair Pathway in Human Spermatozoa, Mediated by OGG1","volume":"126","author":"Smith","year":"2013","journal-title":"J. Cell Sci."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"e12968","DOI":"10.1111\/and.12968","article-title":"Effect of Sperm Pooling with Seminal Plasma Collected in Breeding or Nonbreeding Season on Saanen Goat Sperm Cryosurvival","volume":"50","author":"Ustuner","year":"2018","journal-title":"Andrologia"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1016\/j.cryobiol.2020.03.008","article-title":"The Synergistic Effect of Trehalose and Low Concentrations of Cryoprotectants Can Improve Post-Thaw Ram Sperm Parameters","volume":"95","author":"Ozturk","year":"2020","journal-title":"Cryobiology"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"e14547","DOI":"10.1111\/and.14547","article-title":"Thymoquinone Improves Motility, Plasma Membrane Integrity and DNA Integrity of Frozen\u2013Thawed Ram Semen","volume":"54","author":"Inanc","year":"2022","journal-title":"Andrologia"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"468","DOI":"10.1016\/j.rvsc.2009.04.014","article-title":"Effects of Hypotaurine, Cysteamine and Aminoacids Solution on Post-Thaw Microscopic and Oxidative Stress Parameters of Angora Goat Semen","volume":"87","author":"Bucak","year":"2009","journal-title":"Res. Vet. Sci."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1016\/j.cryobiol.2020.05.015","article-title":"Effects of Kinetin Supplementation on the Post-Thaw Motility, Viability, and Structural Integrity of Dog Sperm","volume":"95","author":"Qamar","year":"2020","journal-title":"Cryobiology"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"973","DOI":"10.1292\/jvms.21-0125","article-title":"Resveratrol Supplementation into Extender Protects against Cryodamage in Dog Post-Thaw Sperm","volume":"83","author":"Bang","year":"2021","journal-title":"J. Vet. Med. Sci."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"e13115","DOI":"10.1111\/and.13115","article-title":"Impact of Polymorphism in DNA Repair Genes OGG1 and XRCC1 on Seminal Parameters and Human Male Infertility","volume":"50","author":"Serrano","year":"2018","journal-title":"Andrologia"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1038\/npp.2012.112","article-title":"DNA Methylation and Its Basic Function","volume":"38","author":"Moore","year":"2013","journal-title":"Neuropsychopharmacology"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"e24561","DOI":"10.1002\/jcla.24561","article-title":"OGG1 Contributes to Hepatocellular Carcinoma by Promoting Cell Cycle-Related Protein Expression and Enhancing DNA Oxidative Damage Repair in Tumor Cells","volume":"36","author":"Zhang","year":"2022","journal-title":"J. Clin. Lab. Anal."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"102667","DOI":"10.1016\/j.dnarep.2019.102667","article-title":"Roles of OGG1 in Transcriptional Regulation and Maintenance of Metabolic Homeostasis","volume":"81","author":"Sampath","year":"2019","journal-title":"DNA Repair"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"31","DOI":"10.4103\/1008-682X.122203","article-title":"Oxidative Stress and Male Reproductive Health","volume":"16","author":"Aitken","year":"2014","journal-title":"Asian J. Androl."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"e12876","DOI":"10.1111\/and.12876","article-title":"Chronic Hypobaric Hypoxia Diminishes the Expression of Base Excision Repair OGG1 Enzymes in Spermatozoa","volume":"50","author":"Farias","year":"2018","journal-title":"Andrologia"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1016\/j.toxlet.2013.01.017","article-title":"OGG1 Ser326Cys Polymorphism Interacts with Cigarette Smoking to Increase Oxidative DNA Damage in Human Sperm and the Risk of Male Infertility","volume":"218","author":"Ji","year":"2013","journal-title":"Toxicol. Lett."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.ydbio.2015.07.024","article-title":"Fertilization Stimulates 8-Hydroxy-2\u2032-Deoxyguanosine Repair and Antioxidant Activity to Prevent Mutagenesis in the Embryo","volume":"406","author":"Lord","year":"2015","journal-title":"Dev. Biol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"e14519","DOI":"10.1111\/rda.14519","article-title":"Ameliorative Effect of Crocin on Post-thaw Quality, Fertility-associated Gene Expression and Fertilization Potential of Buffalo (Bubalus bubalis) Bull Sperm","volume":"59","author":"Khan","year":"2024","journal-title":"Reprod. Domest. Anim."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"192","DOI":"10.1038\/s41598-021-03946-w","article-title":"Network Models of Prostate Cancer Immune Microenvironments Identify ROMO1 as Heterogeneity and Prognostic Marker","volume":"12","author":"Wang","year":"2022","journal-title":"Sci. Rep."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"927","DOI":"10.1016\/j.bbrc.2016.08.091","article-title":"Spermine Reduces Reactive Oxygen Species Levels and Decreases Cryocapacitation in Canine Sperm Cryopreservation","volume":"479","author":"Setyawan","year":"2016","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"728","DOI":"10.1038\/s41576-022-00512-6","article-title":"Generating Specificity in Genome Regulation through Transcription Factor Sensitivity to Chromatin","volume":"23","author":"Isbel","year":"2022","journal-title":"Nat. Rev. Genet."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"310","DOI":"10.1038\/s41392-023-01528-y","article-title":"Methylation across the Central Dogma in Health and Diseases: New Therapeutic Strategies","volume":"8","author":"Liu","year":"2023","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Qamar, A.Y., Fang, X., Kim, M.J., and Cho, J. (2019). Myoinositol Supplementation of Freezing Medium Improves the Quality-Related Parameters of Dog Sperm. Animals, 9.","DOI":"10.3390\/ani9121038"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"1011","DOI":"10.1111\/rda.13744","article-title":"Effects of Crocin on Frozen-thawed Sperm Apoptosis, Protamine Expression and Membrane Lipid Oxidation in Yanbian Yellow Cattle","volume":"55","author":"Lv","year":"2020","journal-title":"Reprod. Domest. Anim."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"1163","DOI":"10.1111\/rda.13758","article-title":"The Effect of Astaxanthin Supplementation on the Post-thaw Quality of Dog Semen","volume":"55","author":"Qamar","year":"2020","journal-title":"Reprod. Domest. Anim."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"5194","DOI":"10.1038\/s41467-026-71693-5","article-title":"The Genetic Basis for DNA Methylation Variation across Tissues and Development","volume":"17","author":"Rosenski","year":"2026","journal-title":"Nat. Commun."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"20130512","DOI":"10.1098\/rstb.2013.0512","article-title":"DNA Modifications in the Mammalian Brain","volume":"369","author":"Shin","year":"2014","journal-title":"Philos. Trans. R. Soc. B Biol. Sci."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"1053","DOI":"10.1016\/S0093-691X(00)00251-X","article-title":"Effect of Trehalose and Edta on Cryoprotective Action of Ram Semen Diluents","volume":"53","author":"Aisen","year":"2000","journal-title":"Theriogenology"},{"key":"ref_93","first-page":"681","article-title":"Ko\u00e7 Spermas\u0131n\u0131n K\u0131sa S\u00fcreli Saklanmas\u0131na Konsantrasyonun ve Muhafaza S\u00fcresinin Etkisi","volume":"28","author":"Dayanikli","year":"2022","journal-title":"Kafkas Univ. Vet. Fak. Derg."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"118","DOI":"10.17221\/146\/2018-VETMED","article-title":"Cholesterol-Loaded Cyclodextrin plus Trehalose Improves Quality of Frozen-Thawed Ram Sperm","volume":"64","author":"Inanc","year":"2019","journal-title":"Vet. Med."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1006\/meth.2001.1262","article-title":"Analysis of Relative Gene Expression Data Using Real-Time Quantitative PCR and the 2\u2212\u0394\u0394CT Method","volume":"25","author":"Livak","year":"2001","journal-title":"Methods"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/18\/8422\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T16:41:15Z","timestamp":1790095275000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/18\/8422"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,21]]},"references-count":95,"journal-issue":{"issue":"18","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["ijms27188422"],"URL":"https:\/\/doi.org\/10.3390\/ijms27188422","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,21]]}}}