{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,24]],"date-time":"2026-10-24T01:00:37Z","timestamp":1792803637622,"version":"4.5.0"},"reference-count":76,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2022,4,28]],"date-time":"2022-04-28T00:00:00Z","timestamp":1651104000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Red Meat Research and Development Trust of South Africa","award":["2021-67015-33404"],"award-info":[{"award-number":["2021-67015-33404"]}],"id":[{"id":"https:\/\/ror.org\/033633g49","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Red Meat Research and Development Trust of South Africa","award":["1R01HD084353"],"award-info":[{"award-number":["1R01HD084353"]}],"id":[{"id":"https:\/\/ror.org\/033633g49","id-type":"ROR","asserted-by":"crossref"}]},{"name":"USDA National Institute of Food and Agriculture","award":["2021-67015-33404"],"award-info":[{"award-number":["2021-67015-33404"]}],"id":[{"id":"https:\/\/ror.org\/05qx3fv49","id-type":"ROR","asserted-by":"crossref"}]},{"name":"USDA National Institute of Food and Agriculture","award":["1R01HD084353"],"award-info":[{"award-number":["1R01HD084353"]}],"id":[{"id":"https:\/\/ror.org\/05qx3fv49","id-type":"ROR","asserted-by":"crossref"}]},{"name":"NIH\/NICHD","award":["2021-67015-33404"],"award-info":[{"award-number":["2021-67015-33404"]}]},{"name":"NIH\/NICHD","award":["1R01HD084353"],"award-info":[{"award-number":["1R01HD084353"]}]},{"name":"University of Missouri South African Education Program","award":["2021-67015-33404"],"award-info":[{"award-number":["2021-67015-33404"]}],"id":[{"id":"https:\/\/ror.org\/02ymw8z06","id-type":"ROR","asserted-by":"crossref"}]},{"name":"University of Missouri South African Education Program","award":["1R01HD084353"],"award-info":[{"award-number":["1R01HD084353"]}],"id":[{"id":"https:\/\/ror.org\/02ymw8z06","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Faculty of Veterinary Sciences, University of Pretoria","award":["2021-67015-33404"],"award-info":[{"award-number":["2021-67015-33404"]}],"id":[{"id":"https:\/\/ror.org\/00g0p6g84","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Faculty of Veterinary Sciences, University of Pretoria","award":["1R01HD084353"],"award-info":[{"award-number":["1R01HD084353"]}],"id":[{"id":"https:\/\/ror.org\/00g0p6g84","id-type":"ROR","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Animals"],"abstract":"<jats:p>Cryopreservation and storage of semen for artificial insemination (AI) result in excessive accumulation of reactive oxygen species (ROS). This leads to a shortened life span and reduced motility of spermatozoa post-thawing, with consequent impairment of their function. However, certain levels of ROS are essential to facilitate the capacitation of spermatozoa required for successful fertilisation. Tannins, as well-known antioxidant compounds, may act as ROS binders\/acceptors\/scavengers to inhibit the damaging effects of ROS. This review comprises an analysis of the semen cryopreservation protocol and health functions of tannins, as well as the effects of ROS on fresh and cryopreserved semen\u2019s longevity and fertilisation. Additionally, we surveyed available evidence of the effects of tannin extract feed supplementation on male fertility. We furthermore interrogated existing theories on tannin use as a potential additive to semen extenders, its relationship with semen quality, and to what degree existing theories have been investigated to develop testable new hypotheses. Emphasis was placed on the effects of tannins on ROS, their involvement in regulating sperm structure and function during cryopreservation, and on post-thaw sperm motility, capacitation, and fertilising ability. The diverse effects of tannins on the reproductive system as a result of their potential metal ion chelation, protein precipitation, and biological antioxidant abilities have been identified. The current data are the first to support the further investigation of the incorporation of tannin-rich plant extracts into semen extenders to enhance the post-thaw survival, motility, and fertilising ability of cryopreserved spermatozoa.<\/jats:p>","DOI":"10.3390\/ani12091130","type":"journal-article","created":{"date-parts":[[2022,4,28]],"date-time":"2022-04-28T12:06:01Z","timestamp":1651147561000},"page":"1130","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":15,"title":["Potential Use of Tannin Extracts as Additives in Semen Destined for Cryopreservation: A Review"],"prefix":"10.3390","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1671-1224","authenticated-orcid":false,"given":"Mohammed S.","family":"Liman","sequence":"first","affiliation":[{"name":"Department of Production Animal Studies, Faculty of Veterinary Science, University of Pretoria, Pretoria 0110, South Africa"},{"name":"Niger State Livestock and Fisheries Institute, Ministry of Livestock and Fisheries Development, Minna 920001, Niger State, Nigeria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8240-3414","authenticated-orcid":false,"given":"Abubeker","family":"Hassen","sequence":"additional","affiliation":[{"name":"Department of Animal and Wildlife Sciences, Faculty of Natural and Agricultural Science, University of Pretoria, Pretoria 0028, South Africa"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8447-0613","authenticated-orcid":false,"given":"Lyndy J.","family":"McGaw","sequence":"additional","affiliation":[{"name":"Phytomedicine Programme, Department of Paraclinical Sciences, Faculty of Veterinary Science, University of Pretoria, Pretoria 0110, South Africa"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9231-2823","authenticated-orcid":false,"given":"Peter","family":"Sutovsky","sequence":"additional","affiliation":[{"name":"Division of Animal Sciences, Department of Obstetrics, Gynecology and Women\u2019s Health, University of Missouri, Columbia, MO 65211-5300, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dietmar E.","family":"Holm","sequence":"additional","affiliation":[{"name":"Department of Production Animal Studies, Faculty of Veterinary Science, University of Pretoria, Pretoria 0110, South Africa"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2022,4,28]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1007\/s13273-017-0015-8","article-title":"Actions and mechanisms of reactive oxygen species and antioxidative system in semen","volume":"13","author":"Gao","year":"2017","journal-title":"Mol. Cell. Toxicol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"659","DOI":"10.1071\/RD9950659","article-title":"Free radicals, lipid peroxidation and sperm function","volume":"7","author":"Aitken","year":"1995","journal-title":"Reprod. Fertil. Dev."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"759","DOI":"10.1530\/REP-11-0310","article-title":"Sperm surface changes and physiological consequences induced by sperm handling and storage","volume":"142","author":"Leahy","year":"2011","journal-title":"Reproduction"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1006\/cryo.1995.1014","article-title":"Comparison of the efficacy of conventional slow freezing and rapid cryopreservation methods for bovine embryos","volume":"32","author":"Kruip","year":"1995","journal-title":"Cryobiology"},{"key":"ref_5","first-page":"791","article-title":"Quality and fertility of the frozen-thawed bull semen as affected by the different cryoprotectants and glutathione levels","volume":"7","author":"Eid","year":"2011","journal-title":"J. Am. Sci."},{"key":"ref_6","first-page":"39","article-title":"In vitro evaluation of sperm quality related to in vivo function and fertility","volume":"64","author":"Barth","year":"2007","journal-title":"Soc. Reprod. Fertil. Suppl."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1016\/j.cryobiol.2012.10.006","article-title":"Effects of various cryoprotectants on bull sperm quality, DNA integrity and oxidative stress parameters","volume":"66","author":"Tuncer","year":"2013","journal-title":"Cryobiology"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"871","DOI":"10.1071\/RD9950871","article-title":"Recent developments and concepts in the cryopreservation of spermatozoa and the assessment of their post-thawing function","volume":"7","author":"Watson","year":"1995","journal-title":"Reprod. Fertil. Dev."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1103090461","DOI":"10.1292\/jvms.10-0460","article-title":"Effect of butylated hydroxytoluene on dog sperm longevity in chilling storage and cryopreservation","volume":"73","author":"Sahashi","year":"2011","journal-title":"J. Vet. Med. Sci."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1016\/j.cryobiol.2012.12.010","article-title":"Effects of different concentrations of BHT on microscopic and oxidative parameters of Mahabadi goat semen following the freeze\u2013thaw process","volume":"66","author":"Naijian","year":"2013","journal-title":"Cryobiology"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"57","DOI":"10.3109\/19396368.2014.968267","article-title":"Influence of BHT inclusion on post-thaw attributes of human semen","volume":"61","author":"Ghorbani","year":"2015","journal-title":"Syst. Biol. Reprod. Med."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"746","DOI":"10.1111\/rda.12359","article-title":"Comparison of Cryoprotective Effects of Lycopene and Cysteamine in Different Cryoprotectants on Bull Semen and Fertility Results","volume":"49","author":"Tuncer","year":"2014","journal-title":"Reprod. Domest. Anim."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1262\/jrd1955.18.15","article-title":"Cryoprotective action of some amide solutions on spermatozoa of domestic animals. I. Effects of formamide, acetamide and lactamide on the motility of bovine spermatozoa after pellet freezing","volume":"18","author":"Nagase","year":"1972","journal-title":"Jpn. J. Anim. Reprod."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Ros-Santaella, J.L., and Pintus, E. (2021). Plant Extracts as Alternative Additives for Sperm Preservation. Antioxidants, 10.","DOI":"10.3390\/antiox10050772"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.anireprosci.2012.04.002","article-title":"The potential influence of plant-based feed supplements on sperm quantity and quality in livestock: A review","volume":"132","author":"Witschi","year":"2012","journal-title":"Anim. Reprod. Sci."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"895","DOI":"10.1002\/j.1939-4640.2000.tb03420.x","article-title":"The effect of reactive oxygen species on equine sperm motility, viability, acrosomal integrity, mitochondrial membrane potential, and membrane lipid peroxidation","volume":"21","author":"Baumber","year":"2000","journal-title":"J. Androl."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1111\/rda.12554","article-title":"New approaches to boar semen evaluation, processing and improvement","volume":"50","author":"Sutovsky","year":"2015","journal-title":"Reprod. Domest. Anim."},{"key":"ref_18","unstructured":"Senger, P.L. (2012). Endocrinology of the male and spermatogenesis. Pathways to Pregnancy and Parturation, Current Conceptions. [3rd ed.]."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"2061","DOI":"10.1038\/s41467-018-04523-y","article-title":"Zinc ion flux during mammalian sperm capacitation","volume":"9","author":"Kerns","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Kerns, K., Sharif, M., Zigo, M., Xu, W., Hamilton, L.E., Sutovsky, M., Ellersieck, M., Drobnis, E.Z., Bovin, N., and Oko, R. (2020). Sperm cohort-specific zinc signature acquisition and capacitation-induced zinc flux regulate sperm-oviduct and sperm-zona pellucida interactions. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21062121"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1095\/biolreprod.102.012716","article-title":"Endogenous redox activity in mouse spermatozoa and its role in regulating the tyrosine phosphorylation events associated with sperm capacitation","volume":"69","author":"Ecroyd","year":"2003","journal-title":"Biol. Reprod."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"528","DOI":"10.1016\/j.freeradbiomed.2006.04.027","article-title":"Positive role of reactive oxygen species in mammalian sperm capacitation: Triggering and modulation of phosphorylation events","volume":"41","author":"Gagnon","year":"2006","journal-title":"Free Radic. Biol. Med."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"785","DOI":"10.1093\/molehr\/gat067","article-title":"Sperm capacitation: A distant landscape glimpsed but unexplored","volume":"19","author":"Aitken","year":"2013","journal-title":"Mol. Hum. Reprod."},{"key":"ref_24","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_25","first-page":"1","article-title":"A review on medicinal plants used for improvement of spermatogenesis","volume":"8","author":"Khojasteh","year":"2016","journal-title":"Biol. Med."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1037","DOI":"10.1095\/biolreprod59.5.1037","article-title":"Relative impact of oxidative stress on the functional competence and genomic integrity of human spermatozoa","volume":"59","author":"Aitken","year":"1998","journal-title":"Biol. Reprod."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"785","DOI":"10.1530\/jrf.0.0830785","article-title":"Selenium and glutathione peroxidase in seminal plasma of men and bulls","volume":"83","author":"Kantola","year":"1988","journal-title":"Reproduction"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1016\/S0093-691X(97)84066-6","article-title":"Effects of antioxidants on motility and membrane integrity of chilled-stored stallion semen","volume":"48","author":"Aurich","year":"1997","journal-title":"Theriogenology"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1093\/molehr\/3.3.175","article-title":"Capacitation as a regulatory event that primes spermatozoa for the acrosome reaction and fertilization","volume":"3","author":"Leclerc","year":"1997","journal-title":"Mol. Hum. Reprod."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"426","DOI":"10.1016\/j.rvsc.2010.03.025","article-title":"Influence of methionine and dithioerythritol on sperm motility, lipid peroxidation and antioxidant capacities during liquid storage of ram semen","volume":"89","author":"Bucak","year":"2010","journal-title":"Res. Vet. Sci."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"759","DOI":"10.1016\/S0093-691X(03)00048-7","article-title":"Effect of milk-and TRIS-based extenders on the fertility of sheep inseminated vaginally once or twice with liquid semen","volume":"60","author":"Paulenz","year":"2003","journal-title":"Theriogenology"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"557","DOI":"10.1530\/rep.1.00049","article-title":"Birth of lambs of a pre-determined sex after in vitro production of embryos using frozen\u2013thawed sex-sorted and re-frozen\u2013thawed ram spermatozoa","volume":"127","author":"Hollinshead","year":"2004","journal-title":"Reproduction"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"974","DOI":"10.1016\/j.theriogenology.2006.02.042","article-title":"Effect of solid storage on caprine semen conservation at 5 C","volume":"66","author":"Salvador","year":"2006","journal-title":"Theriogenology"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/j.ydbio.2008.03.015","article-title":"Impaired sperm fertilizing ability in mice lacking Cysteine-RIch Secretory Protein 1 (CRISP1)","volume":"320","author":"Maldera","year":"2008","journal-title":"Dev. Biol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"481","DOI":"10.1016\/S0378-4320(00)00099-3","article-title":"The causes of reduced fertility with cryopreserved semen","volume":"60\u201361","author":"Watson","year":"2000","journal-title":"Anim. Reprod. Sci."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1016\/j.theriogenology.2018.06.006","article-title":"Sublethal sperm freezing damage: Manifestations and solutions","volume":"118","author":"Pini","year":"2018","journal-title":"Theriogenology"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/j.theriogenology.2015.09.047","article-title":"Sperm cryopreservation update: Cryodamage, markers, and factors affecting the sperm freezability in pigs","volume":"85","author":"Yeste","year":"2016","journal-title":"Theriogenology"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1111\/rda.12530","article-title":"Boar differences in artificial insemination outcomes: Can they be minimized?","volume":"50","author":"Roca","year":"2015","journal-title":"Reprod. Domest. Anim."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1111\/rda.12569","article-title":"Recent advances in boar sperm cryopreservation: State of the art and current perspectives","volume":"50","author":"Yeste","year":"2015","journal-title":"Reprod. Domest. Anim."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"535","DOI":"10.14737\/journal.aavs\/2014\/2.9.535.542","article-title":"Fertility of commercial sexed semen and the economic analyses of its application in Holstein heifers","volume":"2","author":"Abdalla","year":"2014","journal-title":"Adv. Anim. Vet. Sci."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Y\u00e1nez-Ortiz, I., Catal\u00e1n, J., Rodr\u00edguez-Gil, J.E., Mir\u00f3, J., and Yeste, M. (2021). Advances in sperm cryopreservation in farm animals: Cattle, horse, pig and sheep. Anim. Reprod. Sci., 106904.","DOI":"10.1016\/j.anireprosci.2021.106904"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1016\/j.theriogenology.2021.02.018","article-title":"Gene promoter polymorphisms in boar spermatozoa differing in freezability","volume":"166","author":"Brym","year":"2021","journal-title":"Theriogenology"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"436","DOI":"10.2174\/138955708784223486","article-title":"Condensed and hydrolysable tannins as antioxidants influencing the health","volume":"8","author":"Koleckar","year":"2008","journal-title":"Mini Rev. Med. Chem."},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Pizzi, A. (2008). Tannins: Major Sources, Properties and Applications. Monomers, Polymers and Composites from Renewable Resources, Elsevier.","DOI":"10.1016\/B978-0-08-045316-3.00008-9"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"2191","DOI":"10.3390\/molecules16032191","article-title":"Tannins of constant structure in medicinal and food plants\u2014Hydrolyzable tannins and polyphenols related to tannins","volume":"16","author":"Okuda","year":"2011","journal-title":"Molecules"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1039\/C4NP00085D","article-title":"Antiviral drug discovery: Broad-spectrum drugs from nature","volume":"32","author":"Martinez","year":"2015","journal-title":"Nat. Prod. Rep."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"100481","DOI":"10.1016\/j.scp.2021.100481","article-title":"Tannins medical\/pharmacological and related applications: A critical review","volume":"22","author":"Pizzi","year":"2021","journal-title":"Sustain. Chem. Pharm."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"268","DOI":"10.1002\/ijch.201400134","article-title":"Naturally occurring polyphenolic glucosidase inhibitors","volume":"55","author":"Yang","year":"2015","journal-title":"Isr. J. Chem."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1016\/S0377-8401(01)00234-6","article-title":"Use of tannin-binding chemicals to assay for tannins and their negative postingestive effects in ruminants","volume":"91","author":"Silanikove","year":"2001","journal-title":"Anim. Feed Sci. Technol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1021\/jf049144d","article-title":"Metabolism of antioxidant and chemopreventive ellagitannins from strawberries, raspberries, walnuts, and oak-aged wine in humans: Identification of biomarkers and individual variability","volume":"53","year":"2005","journal-title":"J. Agric. Food Chem."},{"key":"ref_51","first-page":"2502","article-title":"Tannins: An antinutrient with positive effect to manage diabetes","volume":"2277","author":"Kumari","year":"2012","journal-title":"Res. J. Recent Sci. ISSN"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1801","DOI":"10.1016\/S0093-691X(02)00653-2","article-title":"Cryopreservation and post-thawed fertility of ram semen frozen in different trehalose concentrations","volume":"57","author":"Aisen","year":"2002","journal-title":"Theriogenology"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1016\/j.smallrumres.2016.03.017","article-title":"Influence of lycopene and cysteamine on sperm and oxidative stress parameters during liquid storage of ram semen at 5 C","volume":"137","author":"Akalin","year":"2016","journal-title":"Small Rumin. Res."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1002\/bies.950160409","article-title":"Reactive oxygen species generation and human spermatozoa: The balance of benefit and risk","volume":"16","author":"Aitken","year":"1994","journal-title":"Bioessays"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1111\/j.1365-2605.2005.00531.x","article-title":"Impact of radio frequency electromagnetic radiation on DNA integrity in the male germline","volume":"28","author":"Aitken","year":"2005","journal-title":"Int. J. Androl."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.foodchem.2016.07.136","article-title":"\u03b1-Glucosidase inhibition and antioxidant activity of an oenological commercial tannin. Extraction, fractionation and analysis by HPLC\/ESI-MS\/MS and 1H NMR","volume":"215","author":"Muccilli","year":"2017","journal-title":"Food Chem."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"568","DOI":"10.1016\/j.foodres.2017.12.054","article-title":"The effect of prefermentative addition of gallic acid and ellagic acid on the red wine color, copigmentation and phenolic profiles during wine aging","volume":"106","author":"Zhang","year":"2018","journal-title":"Food Res. Int."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"6978","DOI":"10.3390\/ijms16046978","article-title":"A review of polyphenolics in oak woods","volume":"16","author":"Zhang","year":"2015","journal-title":"Int. J. Mol. Sci."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"113702","DOI":"10.1016\/j.jep.2020.113702","article-title":"In vitro effects of Capparis spinosa L. extract on human sperm function, DNA fragmentation, and oxidative stress","volume":"269","author":"Rad","year":"2021","journal-title":"J. Ethnopharmacol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1016\/j.theriogenology.2017.12.015","article-title":"Biological effects of polyphenol-rich extract and fractions from an oenological oak-derived tannin on in vitro swine sperm capacitation and fertilizing ability","volume":"108","author":"Spinaci","year":"2018","journal-title":"Theriogenology"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"19783","DOI":"10.1007\/s11356-020-11509-1","article-title":"Zingiber officinale ethanolic extract attenuates oxidative stress, steroidogenic gene expression alterations, and testicular histopathology induced by sodium arsenite in male rats","volume":"28","author":"Seif","year":"2021","journal-title":"Environ. Sci. Pollut. Res."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"56","DOI":"10.4103\/2305-0500.311609","article-title":"Antifertility effects of 60-day oral gavage of ethanol extract of Spondias mombin leaves in guinea pigs: A biochemical, reproductive and histological study","volume":"10","author":"Ogunro","year":"2021","journal-title":"Asian Pac. J. Reprod."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"2205","DOI":"10.1007\/s12011-021-02818-8","article-title":"Wild Garlic Allium triquetrum L. Alleviates Lead Acetate-Induced Testicular Injuries in Rats","volume":"200","author":"Kahalerras","year":"2022","journal-title":"Biol. Trace Elem. Res."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"400","DOI":"10.2174\/1389201021666200730145621","article-title":"Leaf and Fruit Methanolic Extracts of Azadirachta indica Exhibit Antifertility Activity on Rats\u2019 Sperm Quality and Testicular Histology","volume":"22","author":"Irais","year":"2021","journal-title":"Curr. Pharm. Biotechnol."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Ahmed, O., Lehloenya, K., Mphaphathi, M., and Hassen, A. (2021). Effect of Acacia mearnsii Tannin Extract Supplementation on Reproductive Performance and Oxidative Status of South African Mutton Merino Rams. Animals, 11.","DOI":"10.3390\/ani11113266"},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Shehzad, M., Rasheed, H., Naqvi, S.A., Al-Khayri, J.M., Lorenzo, J.M., Alaghbari, M.A., Manzoor, M.F., and Aadil, R.M. (2021). Therapeutic Potential of Date Palm against Human Infertility: A Review. Metabolites, 11.","DOI":"10.3390\/metabo11060408"},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Sobeh, M., Mahmoud, M.F., Sabry, O.M., Adel, R., Dmirieh, M., El-Shazly, A.M., and Wink, M. (2017). HPLC-PDA-MS\/MS characterization of bioactive secondary metabolites from Turraea fischeri bark extract and its antioxidant and hepatoprotective activities in vivo. Molecules, 22.","DOI":"10.3390\/molecules22122089"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1016\/j.joim.2020.03.003","article-title":"Evaluation of antioxidant capacity and reproductive toxicity of aqueous extract of Thai Mucuna pruriens seeds","volume":"18","author":"Iamsaard","year":"2020","journal-title":"J. Integr. Med."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"160","DOI":"10.1016\/j.theriogenology.2021.06.015","article-title":"Dietary supplementation of grape seed tannin extract stimulated testis development, changed fatty acid profiles and increased testis antioxidant capacity in pre-puberty hu lambs","volume":"172","author":"Li","year":"2021","journal-title":"Theriogenology"},{"key":"ref_70","first-page":"393","article-title":"Evaluation of Anti Fertility Activities of Bark Extracts of Caesalpinia pulcherrima Linn (Caesalpiniaceae) in Rats","volume":"83","author":"Shaik","year":"2021","journal-title":"Indian J. Pharm. Sci."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"530","DOI":"10.14202\/vetworld.2020.530-537","article-title":"The effect of crude guava leaf tannins on motility, viability, and intact plasma membrane of stored spermatozoa of Etawa crossbred goats","volume":"13","author":"Wurlina","year":"2020","journal-title":"Vet. World"},{"key":"ref_72","doi-asserted-by":"crossref","unstructured":"Ros-Santaella, J.L., and Pintus, E. (2017). Rooibos (Aspalathus linearis) extract enhances boar sperm velocity up to 96 h of semen storage. PLoS ONE, 12.","DOI":"10.1371\/journal.pone.0183682"},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Galeati, G., Bucci, D., Nerozzi, C., Gadani, B., Tamanini, C., Mislei, B., and Spinaci, M. (2020). Improvement of in vitro fertilization by a tannin rich vegetal extract addition to frozen thawed boar sperm. Anim. Reprod., 17.","DOI":"10.1590\/1984-3143-ar2019-0130"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"604477","DOI":"10.3389\/fvets.2020.604477","article-title":"A polyphenol-rich extract from Entada abyssinica reduces oxidative damage in cryopreserved ram semen","volume":"956","author":"Sobeh","year":"2020","journal-title":"Front. Vet. Sci."},{"key":"ref_75","first-page":"1209","article-title":"Effect of adding n-acetylcystiene and avena sativa extract to tris extender on post-cryopreservative semen characteristics of holstein bulls","volume":"20","author":"Ali","year":"2020","journal-title":"Plant Arch."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Fitriyah, A., Said, D.O., and Harianto, H. (2017). Improvement of Sperm Quality of Bali Cattle by Supplementation of Crude Tannin in the Semen. Proceeding of the 1st International Conference on Tropical Agriculture, Springer.","DOI":"10.1007\/978-3-319-60363-6_30"}],"container-title":["Animals"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2615\/12\/9\/1130\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T23:02:33Z","timestamp":1760137353000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2615\/12\/9\/1130"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,4,28]]},"references-count":76,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2022,5]]}},"alternative-id":["ani12091130"],"URL":"https:\/\/doi.org\/10.3390\/ani12091130","relation":{},"ISSN":["2076-2615"],"issn-type":[{"value":"2076-2615","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,4,28]]}}}