{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,3]],"date-time":"2026-11-03T04:09:56Z","timestamp":1793678996445,"version":"4.5.1"},"reference-count":52,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-017"},{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-012"},{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-004"}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Animal Reproduction Science"],"published-print":{"date-parts":[[2026,10]]},"DOI":"10.1016\/j.anireprosci.2026.108304","type":"journal-article","created":{"date-parts":[[2026,8,7]],"date-time":"2026-08-07T06:34:36Z","timestamp":1786084476000},"page":"108304","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":1,"special_numbering":"C","title":["Erythropoietin sustains functional and biochemical integrity of ram spermatozoa during prolonged chilled storage"],"prefix":"10.1016","volume":"293","author":[{"given":"Konstantinos","family":"Feidantsis","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7365-8805","authenticated-orcid":false,"given":"Maria P.","family":"Tsantarliotou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9269-0675","authenticated-orcid":false,"given":"Zacharenia","family":"Kyrana","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4062-2336","authenticated-orcid":false,"given":"Sophia N.","family":"Lavrentiadou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Evaggelia Chrysoula","family":"Chatzikou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7079-4238","authenticated-orcid":false,"given":"Vasiliki G.","family":"Sapanidou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.anireprosci.2026.108304_bib1","doi-asserted-by":"crossref","first-page":"1132","DOI":"10.1182\/blood.V62.5.1132.1132","article-title":"Erythropoietin production by a human testicular germ cell line","volume":"62","author":"Ascensao","year":"1983","journal-title":"Blood"},{"key":"10.1016\/j.anireprosci.2026.108304_bib2","doi-asserted-by":"crossref","first-page":"2669","DOI":"10.21873\/invivo.12550","article-title":"Erythropoietin increases in vitro motility and vitality of human spermatozoa","volume":"35","author":"Asimakopoulos","year":"2021","journal-title":"Vivo"},{"key":"10.1016\/j.anireprosci.2026.108304_bib3","doi-asserted-by":"crossref","first-page":"469","DOI":"10.1038\/35019501","article-title":"Heat-shock protein 70 inhibits apoptosis by preventing recruitment of procaspase-9 to the Apaf-1 apoptosome","volume":"2","author":"Beere","year":"2000","journal-title":"Nat. Cell. Biol."},{"key":"10.1016\/j.anireprosci.2026.108304_bib4","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1159\/000245630","article-title":"The therapeutic potential of erythropoiesis-stimulating agents for tissue protection: a tale of two receptors","volume":"29","author":"Brines","year":"2010","journal-title":"Blood Purif."},{"key":"10.1016\/j.anireprosci.2026.108304_bib5","doi-asserted-by":"crossref","first-page":"246","DOI":"10.1038\/sj.ki.5001546","article-title":"Discovering erythropoietin\u2019s extra-hematopoietic functions: biology and clinical promise","volume":"70","author":"Brines","year":"2006","journal-title":"Kidney Int."},{"issue":"3","key":"10.1016\/j.anireprosci.2026.108304_bib6","first-page":"a011619","article-title":"Erythropoietin","volume":"1","author":"Bunn","year":"2013","journal-title":"Cold Spring Harb. Perspect. Med."},{"key":"10.1016\/j.anireprosci.2026.108304_bib7","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1016\/j.smallrumres.2011.05.010","article-title":"Effects of reduced glutathione and catalase on the kinematics and membrane functionality of sperm during liquid storage of ram semen","volume":"100","author":"C\u00e2mara","year":"2011","journal-title":"Small Rumin. Res."},{"key":"10.1016\/j.anireprosci.2026.108304_bib8","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1002\/j.1939-4640.1999.tb02504.x","article-title":"Modulation of insulin-like growth factor-1 in the seminal plasma of infertile men","volume":"20","author":"Colombo","year":"1999","journal-title":"J. Androl."},{"key":"10.1016\/j.anireprosci.2026.108304_bib9","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1002\/mrd.1120080104","article-title":"Immunocytochemical localization of protamine in the spermatids of the ram","volume":"8","author":"Courtens","year":"1983","journal-title":"Gamete Res."},{"key":"10.1016\/j.anireprosci.2026.108304_bib10","doi-asserted-by":"crossref","first-page":"466","DOI":"10.1016\/0011-2240(77)90008-6","article-title":"Influence of the cholesterol content of mammalian spermatozoa on susceptibility to cold-shock","volume":"14","author":"Darin-Bennett","year":"1977","journal-title":"Cryobiology"},{"key":"10.1016\/j.anireprosci.2026.108304_bib11","doi-asserted-by":"crossref","DOI":"10.1155\/2023\/5514248","article-title":"Erythropoietin suppresses hepatic fibrosis caused by thioacetamide: role of the PI3K\/Akt and TLR4 signaling pathways","volume":"2023","author":"Elbaset","year":"2023","journal-title":"Oxid. Med. Cell. Longev."},{"key":"10.1016\/j.anireprosci.2026.108304_bib12","series-title":"Salamon\u2019s Artificial Insemination of Sheep and Goats","first-page":"122","article-title":"Frozen storage of semen","author":"Evans","year":"1987"},{"key":"10.1016\/j.anireprosci.2026.108304_bib13","first-page":"753","article-title":"Erythropoietin stimulates testosterone production in man","volume":"78","author":"Foresta","year":"1994","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"10.1016\/j.anireprosci.2026.108304_bib14","doi-asserted-by":"crossref","DOI":"10.1155\/2010\/214074","article-title":"Cellular stress responses: cell survival and cell death","volume":"2010","author":"Fulda","year":"2010","journal-title":"Int. J. Cell. Biol."},{"key":"10.1016\/j.anireprosci.2026.108304_bib15","doi-asserted-by":"crossref","DOI":"10.1126\/sciadv.abb7426","article-title":"Circular RNAs from BOULE play conserved roles in protection against stress-induced fertility decline","volume":"6","author":"Gao","year":"2020","journal-title":"Sci. Adv."},{"key":"10.1016\/j.anireprosci.2026.108304_bib16","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1007\/s12020-011-9579-6","article-title":"Erythropoietin produced by the retina: its role in physiology and diabetic retinopathy","volume":"41","author":"Hern\u00e1ndez","year":"2012","journal-title":"Endocrine"},{"key":"10.1016\/j.anireprosci.2026.108304_bib17","doi-asserted-by":"crossref","first-page":"286","DOI":"10.1172\/JCI112796","article-title":"Human recombinant erythropoietin promotes differentiation of murine megakaryocytes in vitro","volume":"79","author":"Ishibashi","year":"1987","journal-title":"J. Clin. Invest."},{"key":"10.1016\/j.anireprosci.2026.108304_bib18","doi-asserted-by":"crossref","first-page":"302","DOI":"10.1159\/000356193","article-title":"Physiology and pharmacology of erythropoietin","volume":"40","author":"Jelkmann","year":"2013","journal-title":"Transfus. Med. Hemother."},{"key":"10.1016\/j.anireprosci.2026.108304_bib19","doi-asserted-by":"crossref","first-page":"208","DOI":"10.1016\/j.smallrumres.2011.03.057","article-title":"Effect of semen extenders on sperm parameters of ram semen during liquid storage at 4\u00b0C","volume":"99","author":"Kasimanickam","year":"2011","journal-title":"Small Rumin. Res."},{"key":"10.1016\/j.anireprosci.2026.108304_bib20","doi-asserted-by":"crossref","DOI":"10.1111\/apha.13826","article-title":"EPO synthesis induced by HIF-PHD inhibition is dependent on myofibroblast transdifferentiation and colocalizes with non-injured nephron segments in murine kidney fibrosis","volume":"235","author":"Kobayashi","year":"2022","journal-title":"Acta Physiol."},{"key":"10.1016\/j.anireprosci.2026.108304_bib21","doi-asserted-by":"crossref","first-page":"673","DOI":"10.1046\/j.1537-2995.1990.30891020321.x","article-title":"Control of red cell production: the roles of programmed cell death (apoptosis) and erythropoietin","volume":"30","author":"Koury","year":"1990","journal-title":"Transfusion"},{"key":"10.1016\/j.anireprosci.2026.108304_bib22","doi-asserted-by":"crossref","first-page":"3219","DOI":"10.3390\/ani13203219","article-title":"Melatonin protects bovine spermatozoa by reinforcing their antioxidant defenses","volume":"13","author":"Lavrentiadou","year":"2023","journal-title":"Animals"},{"key":"10.1016\/j.anireprosci.2026.108304_bib23","doi-asserted-by":"crossref","DOI":"10.18637\/jss.v025.i01","article-title":"FactoMineR: an R package for multivariate analysis","volume":"25","author":"L\u00ea","year":"2008","journal-title":"J. Stat. Softw."},{"key":"10.1016\/j.anireprosci.2026.108304_bib24","doi-asserted-by":"crossref","first-page":"684","DOI":"10.1095\/biolreprod55.3.684","article-title":"Cyclic adenosine 3\u2032,5\u2032-monophosphate-dependent regulation of protein tyrosine phosphorylation in relation to human sperm capacitation and motility","volume":"55","author":"Leclerc","year":"1996","journal-title":"Biol. Reprod."},{"key":"10.1016\/j.anireprosci.2026.108304_bib25","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1159\/000322975","article-title":"Erythropoietin: a hormone with multiple functions","volume":"78","author":"Lombardero","year":"2011","journal-title":"Pathobiology"},{"key":"10.1016\/j.anireprosci.2026.108304_bib26","doi-asserted-by":"crossref","first-page":"321","DOI":"10.1161\/01.ATV.21.3.321","article-title":"Reactive oxygen species regulate heat-shock protein 70 via the JAK\/STAT pathway","volume":"21","author":"Madamanchi","year":"2001","journal-title":"Arterioscler. Thromb. Vasc. Biol."},{"key":"10.1016\/j.anireprosci.2026.108304_bib27","doi-asserted-by":"crossref","first-page":"2872","DOI":"10.1182\/blood.V98.9.2872","article-title":"Erythropoietin expression in primary rat Sertoli and peritubular myoid cells","volume":"98","author":"Magnanti","year":"2001","journal-title":"Blood"},{"key":"10.1016\/j.anireprosci.2026.108304_bib28","doi-asserted-by":"crossref","first-page":"2918","DOI":"10.3390\/jcm13102918","article-title":"NGF, EPO, and IGF-1 in the male reproductive system","volume":"13","author":"Metallinou","year":"2024","journal-title":"J. Clin. Med."},{"key":"10.1016\/j.anireprosci.2026.108304_bib29","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1093\/humupd\/3.5.403","article-title":"A critical review of the physiological importance and analysis of sperm movement in mammals","volume":"3","author":"Mortimer","year":"1997","journal-title":"Hum. Reprod. Update"},{"key":"10.1016\/j.anireprosci.2026.108304_bib30","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1016\/j.theriogenology.2009.10.009","article-title":"Effect of storage duration, storage temperature, and diluent on the viability and fertility of fresh ram sperm","volume":"73","author":"O\u2019Hara","year":"2010","journal-title":"Theriogenology"},{"key":"10.1016\/j.anireprosci.2026.108304_bib31","doi-asserted-by":"crossref","DOI":"10.1016\/j.anireprosci.2025.107853","article-title":"Antioxidant components of ram seminal plasma are influenced by age, season, and the RsaI polymorphism of the MTNR1A gene","volume":"277","author":"Pe\u00f1a-Delgado","year":"2025","journal-title":"Anim. Reprod. Sci."},{"key":"10.1016\/j.anireprosci.2026.108304_bib32","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/j.smallrumres.2016.06.007","article-title":"Addition of antioxidants improves quality of ram spermatozoa during preservation at 4\u00b0C","volume":"141","author":"Rather","year":"2016","journal-title":"Small Rumin. Res."},{"key":"10.1016\/j.anireprosci.2026.108304_bib33","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1111\/j.1439-0531.2007.00920.x","article-title":"CASA assessment of kinematic parameters of ram spermatozoa and their relationship to migration efficiency in ruminant cervical mucus","volume":"43","author":"Robayo","year":"2008","journal-title":"Reprod. Domest. Anim."},{"key":"10.1016\/j.anireprosci.2026.108304_bib34","doi-asserted-by":"crossref","first-page":"4635","DOI":"10.1073\/pnas.95.8.4635","article-title":"vivo evidence that erythropoietin protects neurons from ischemic damage","volume":"95","author":"Sakanaka","year":"1998","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.anireprosci.2026.108304_bib35","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1016\/0378-4320(94)01327-I","article-title":"Frozen storage of ram semen I. Processing, freezing, thawing and fertility after cervical insemination","volume":"37","author":"Salamon","year":"1995","journal-title":"Anim. Reprod. Sci."},{"key":"10.1016\/j.anireprosci.2026.108304_bib36","doi-asserted-by":"crossref","first-page":"2175","DOI":"10.3390\/ani14152175","article-title":"The role of erythropoietin in bovine sperm physiology","volume":"14","author":"Sapanidou","year":"2024","journal-title":"Animals"},{"key":"10.1016\/j.anireprosci.2026.108304_bib37","doi-asserted-by":"crossref","first-page":"1329","DOI":"10.3390\/molecules30061329","article-title":"Supplementing freezing medium with crocin exerts a protective effect on bovine spermatozoa through the modulation of a heat shock-mediated apoptotic pathway","volume":"30","author":"Sapanidou","year":"2025","journal-title":"Molecules"},{"key":"10.1016\/j.anireprosci.2026.108304_bib38","doi-asserted-by":"crossref","first-page":"773","DOI":"10.3390\/ani16050773","article-title":"Crocin supplementation of semen extender preserves chilled ram semen quality during extended storage","volume":"16","author":"Sapanidou","year":"2026","journal-title":"Animals"},{"key":"10.1016\/j.anireprosci.2026.108304_bib39","doi-asserted-by":"crossref","first-page":"32430","DOI":"10.1074\/jbc.271.50.32430","article-title":"Association of JAK2 and STAT5 with erythropoietin receptors. Role of receptor phosphorylation in erythropoietin signal transduction","volume":"13","author":"Sawyer","year":"1996","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.anireprosci.2026.108304_bib40","doi-asserted-by":"crossref","DOI":"10.1016\/j.smallrumres.2020.106139","article-title":"Effects of reduced glutathione on ram sperm parameters, antioxidant status, mitochondrial activity and the abundance of hexose transporters during liquid storage at 5 \u2103","volume":"189","author":"Shi","year":"2020","journal-title":"Small Rumin. Res."},{"key":"10.1016\/j.anireprosci.2026.108304_bib41","doi-asserted-by":"crossref","first-page":"3345","DOI":"10.3390\/biomedicines11123345","article-title":"A possible role for nerve growth factor and its receptors in human sperm pathology","volume":"11","author":"Stabile","year":"2023","journal-title":"Biomedicines"},{"key":"10.1016\/j.anireprosci.2026.108304_bib42","doi-asserted-by":"crossref","first-page":"310","DOI":"10.1111\/j.1600-079X.2010.00843.x","article-title":"Melatonin protects ram spermatozoa from cryopreservation injuries in a dose-dependent manner","volume":"50","author":"Succu","year":"2011","journal-title":"J. Pineal Res."},{"key":"10.1016\/j.anireprosci.2026.108304_bib43","doi-asserted-by":"crossref","first-page":"798","DOI":"10.1016\/j.fertnstert.2003.07.039","article-title":"Detection of erythropoietin in human seminal plasma","volume":"81","author":"Temma","year":"2004","journal-title":"Fertil. Steril."},{"key":"10.1016\/j.anireprosci.2026.108304_bib44","doi-asserted-by":"crossref","first-page":"718","DOI":"10.1016\/j.rbmo.2010.05.022","article-title":"Erythropoietin receptor-like immunostaining on human spermatozoa","volume":"21","author":"Tug","year":"2010","journal-title":"Reprod. Biomed. Online"},{"key":"10.1016\/j.anireprosci.2026.108304_bib45","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1016\/S0093-691X(01)00664-1","article-title":"Computer assisted semen analyzers in andrology research and veterinary practice","volume":"57","author":"Verstegen","year":"2002","journal-title":"Theriogenology"},{"key":"10.1016\/j.anireprosci.2026.108304_bib46","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1016\/j.anireprosci.2014.02.005","article-title":"vitro assessment of sperm quality from rams of high and low field fertility","volume":"146","author":"Vicente-Fiel","year":"2014","journal-title":"Anim. Reprod. Sci."},{"key":"10.1016\/j.anireprosci.2026.108304_bib47","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1017\/S0003356100004384","article-title":"A note on the seasonal oestrous period in three breeds of Greek dairy sheep","volume":"49","author":"Vosniakou","year":"1989","journal-title":"Anim. Sci."},{"key":"10.1016\/j.anireprosci.2026.108304_bib48","doi-asserted-by":"crossref","first-page":"1063","DOI":"10.1152\/physrev.1992.72.4.1063","article-title":"Mammalian stress response: cell physiology, structure\/function of stress proteins, and implications for medicine and disease","volume":"72","author":"Welch","year":"1992","journal-title":"Physiol. Rev."},{"key":"10.1016\/j.anireprosci.2026.108304_bib49","first-page":"161","article-title":"Recombinant human erythropoietin pretreatment attenuates myocardial infarct size: a possible mechanism involves heat shock protein 70 and attenuation of nuclear factor-kappaB","volume":"35","author":"Xu","year":"2005","journal-title":"Ann. Clin. Lab. Sci."},{"key":"10.1016\/j.anireprosci.2026.108304_bib50","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1016\/j.mce.2003.10.031","article-title":"Effect of erythropoietin on Leydig cell is associated with the activation of Stat5 pathway","volume":"213","author":"Yamazaki","year":"2004","journal-title":"Mol. Cell. Endocrinol."},{"key":"10.1016\/j.anireprosci.2026.108304_bib51","doi-asserted-by":"crossref","first-page":"1754","DOI":"10.1096\/fj.02-1191fje","article-title":"Preconditioning with erythropoietin protects against subsequent ischemia-reperfusion injury in rat kidney","volume":"17","author":"Yang","year":"2003","journal-title":"FASEB J."},{"key":"10.1016\/j.anireprosci.2026.108304_bib52","doi-asserted-by":"crossref","first-page":"10296","DOI":"10.3390\/ijms150610296","article-title":"Erythropoietin action in stress response, tissue maintenance and metabolism","volume":"15","author":"Zhang","year":"2014","journal-title":"Int. J. Mol. Sci."}],"container-title":["Animal Reproduction Science"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0378432026002071?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0378432026002071?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T00:49:00Z","timestamp":1788482940000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0378432026002071"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10]]},"references-count":52,"alternative-id":["S0378432026002071"],"URL":"https:\/\/doi.org\/10.1016\/j.anireprosci.2026.108304","relation":{},"ISSN":["0378-4320"],"issn-type":[{"value":"0378-4320","type":"print"}],"subject":[],"published":{"date-parts":[[2026,10]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Erythropoietin sustains functional and biochemical integrity of ram spermatozoa during prolonged chilled storage","name":"articletitle","label":"Article Title"},{"value":"Animal Reproduction Science","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.anireprosci.2026.108304","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 Elsevier B.V. All rights are reserved, including those for text and data mining, AI training, and similar technologies.","name":"copyright","label":"Copyright"}],"article-number":"108304"}}