{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,26]],"date-time":"2026-09-26T08:09:01Z","timestamp":1790410141721,"version":"4.1.0"},"reference-count":95,"publisher":"Cambridge University Press (CUP)","issue":"1","license":[{"start":{"date-parts":[[2019,10,11]],"date-time":"2019-10-11T00:00:00Z","timestamp":1570752000000},"content-version":"unspecified","delay-in-days":0,"URL":"https:\/\/www.cambridge.org\/core\/terms"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Zygote"],"published-print":{"date-parts":[[2020,2]]},"abstract":"<jats:title>Summary<\/jats:title><jats:p>Sperm DNA fragmentation is referred to as one of the main causes of male infertility. Failures in the protamination process, apoptosis and action of reactive oxygen species (ROS) are considered the most important causes of DNA fragmentation. Action of ROS or changes in sperm protamination would increase the susceptibility of sperm DNA to fragmentation. Routine semen analysis is unable to estimate sperm chromatin damage. Sperm DNA integrity influences sperm functional capability, therefore tests that measure sperm DNA fragmentation are important to assess fertility disorders. Actually, there is a considerable number of methods for assessing sperm DNA fragmentation and chromatin integrity, sperm chromatin stability assay (SCSA modified), sperm chromatin dispersion (SCD), comet assay, transferase dUTP nick end labelling (TUNEL); and protamine evaluation in sperm chromatin assay, such as toluidine blue, CMA3, protamine expression and evaluation of cysteine radicals. This review aims to describe the main causes of sperm DNA fragmentation and the tests commonly used to evaluate sperm DNA fragmentation.<\/jats:p>","DOI":"10.1017\/s0967199419000595","type":"journal-article","created":{"date-parts":[[2019,10,11]],"date-time":"2019-10-11T11:53:10Z","timestamp":1570794790000},"page":"1-8","source":"Crossref","is-referenced-by-count":53,"title":["Sperm DNA fragmentation: causes and identification"],"prefix":"10.1017","volume":"28","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-8536-3123","authenticated-orcid":false,"given":"Thais Rose dos Santos","family":"Hamilton","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5967-9316","authenticated-orcid":false,"given":"Mayra Elena Ortiz D\u2019\u00c1vila","family":"Assump\u00e7\u00e3o","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"56","published-online":{"date-parts":[[2019,10,11]]},"reference":[{"key":"S0967199419000595_ref32","doi-asserted-by":"crossref","first-page":"610","DOI":"10.4103\/1008-682X.153853","article-title":"A model for the control of DNA integrity by the sperm nuclear matrix","volume":"17","author":"Gawecka","year":"2015","journal-title":"Asian J Androl"},{"key":"S0967199419000595_ref14","doi-asserted-by":"publisher","DOI":"10.1093\/molehr\/2.3.139"},{"key":"S0967199419000595_ref30","doi-asserted-by":"publisher","DOI":"10.1016\/j.rvsc.2012.09.001"},{"key":"S0967199419000595_ref29","doi-asserted-by":"publisher","DOI":"10.1111\/rda.12817"},{"key":"S0967199419000595_ref22","doi-asserted-by":"publisher","DOI":"10.1002\/cyto.a.10015"},{"key":"S0967199419000595_ref18","doi-asserted-by":"publisher","DOI":"10.1016\/j.aju.2017.11.002"},{"key":"S0967199419000595_ref16","doi-asserted-by":"publisher","DOI":"10.1093\/humupd\/dml057"},{"key":"S0967199419000595_ref15","doi-asserted-by":"publisher","DOI":"10.2527\/jas.2012-5040"},{"key":"S0967199419000595_ref54","first-page":"104","article-title":"Toluidine blue as stain for optimal morphometric evaluation of frozen\u2013thawed bull spermatozoa","volume":"45","author":"Quintero-Moreno","year":"2010","journal-title":"Reprod Dom Anim"},{"key":"S0967199419000595_ref12","doi-asserted-by":"publisher","DOI":"10.1016\/j.theriogenology.2003.10.016"},{"key":"S0967199419000595_ref11","doi-asserted-by":"publisher","DOI":"10.1186\/gb-2007-8-9-227"},{"key":"S0967199419000595_ref6","doi-asserted-by":"crossref","first-page":"232","DOI":"10.1002\/j.1939-4640.1992.tb00306.x","article-title":"Evidence for increased lipid peroxidative damage and loss of superoxide dismutase activity as a mode of sublethal cryodamage to human sperm during cryopreservation","volume":"13","author":"Alvarez","year":"1992","journal-title":"J Androl"},{"key":"S0967199419000595_ref5","doi-asserted-by":"publisher","DOI":"10.1095\/biolreprod59.5.1037"},{"key":"S0967199419000595_ref1","doi-asserted-by":"publisher","DOI":"10.1016\/j.bpc.2018.04.003"},{"key":"S0967199419000595_ref21","doi-asserted-by":"publisher","DOI":"10.1095\/biolreprod.114.124255"},{"key":"S0967199419000595_ref26","doi-asserted-by":"publisher","DOI":"10.2164\/jandrol.04152"},{"key":"S0967199419000595_ref52","doi-asserted-by":"publisher","DOI":"10.1111\/and.12079"},{"key":"S0967199419000595_ref23","doi-asserted-by":"publisher","DOI":"10.1002\/j.1939-4640.2002.tb02599.x"},{"key":"S0967199419000595_ref68","doi-asserted-by":"crossref","first-page":"352","DOI":"10.1159\/000134138","article-title":"The rat PRM3 gene is an intronless member of the protamine gene cluster and is expressed in haploid male germ cells","volume":"71","author":"Schluter","year":"1995","journal-title":"Cytogenet Cell Genet"},{"key":"S0967199419000595_ref17","doi-asserted-by":"publisher","DOI":"10.1016\/j.theriogenology.2017.11.006"},{"key":"S0967199419000595_ref78","doi-asserted-by":"publisher","DOI":"10.1080\/10520290902843595"},{"key":"S0967199419000595_ref81","doi-asserted-by":"publisher","DOI":"10.1093\/humrep\/deu228"},{"key":"S0967199419000595_ref8","doi-asserted-by":"publisher","DOI":"10.2164\/jandrol.106.000703"},{"key":"S0967199419000595_ref28","first-page":"67","article-title":"Variable inter-assay estimation of sperm DNA fragmentation in stallions classified as good and bad semen freezers","volume":"39","author":"Ferreira","year":"2018","journal-title":"Cryo Letters"},{"key":"S0967199419000595_ref80","doi-asserted-by":"publisher","DOI":"10.1093\/humrep\/deu040"},{"key":"S0967199419000595_ref27","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1002\/j.1939-4640.2003.tb02641.x","article-title":"The sperm chromatin dispersion test: a simple method for the determination of sperm DNA fragmentation","volume":"24","author":"Fern\u00e1ndez","year":"2003","journal-title":"J Androl"},{"key":"S0967199419000595_ref93","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1007\/978-1-4419-6857-9","volume-title":"Sperm chromatin. Biological and clinical applications in male infertility and assisted reproduction","author":"Zini","year":"2011"},{"key":"S0967199419000595_ref75","doi-asserted-by":"publisher","DOI":"10.1007\/s10815-015-0635-7"},{"key":"S0967199419000595_ref87","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M312777200"},{"key":"S0967199419000595_ref3","doi-asserted-by":"publisher","DOI":"10.1016\/j.fertnstert.2005.03.080"},{"key":"S0967199419000595_ref20","doi-asserted-by":"publisher","DOI":"10.1016\/j.theriogenology.2005.10.024"},{"key":"S0967199419000595_ref33","doi-asserted-by":"publisher","DOI":"10.1002\/mrd.21394"},{"key":"S0967199419000595_ref4","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1097-010X(19991101)284:6<696::AID-JEZ11>3.0.CO;2-E"},{"key":"S0967199419000595_ref70","doi-asserted-by":"publisher","DOI":"10.1016\/j.anireprosci.2015.05.015"},{"key":"S0967199419000595_ref77","doi-asserted-by":"publisher","DOI":"10.2741\/e415"},{"key":"S0967199419000595_ref10","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.93.2.298"},{"key":"S0967199419000595_ref50","doi-asserted-by":"publisher","DOI":"10.1017\/S0967199410000286"},{"key":"S0967199419000595_ref7","doi-asserted-by":"publisher","DOI":"10.1007\/s10815-007-9177-y"},{"key":"S0967199419000595_ref9","doi-asserted-by":"publisher","DOI":"10.1021\/bi00215a026"},{"key":"S0967199419000595_ref56","doi-asserted-by":"publisher","DOI":"10.1111\/j.2047-2927.2013.00111.x"},{"key":"S0967199419000595_ref62","doi-asserted-by":"publisher","DOI":"10.1111\/rda.13087"},{"key":"S0967199419000595_ref25","doi-asserted-by":"publisher","DOI":"10.1016\/j.anireprosci.2016.01.017"},{"key":"S0967199419000595_ref65","doi-asserted-by":"publisher","DOI":"10.1095\/biolreprod66.4.1061"},{"key":"S0967199419000595_ref13","first-page":"193","article-title":"Fragmentaci\u00f3n Del ADN esperm\u00e1tico","volume":"6","author":"Berenguer","year":"2008","journal-title":"Rev Int Androl"},{"key":"S0967199419000595_ref36","doi-asserted-by":"publisher","DOI":"10.1155\/2018\/5413056"},{"key":"S0967199419000595_ref82","doi-asserted-by":"publisher","DOI":"10.1002\/j.1939-4640.2005.tb01095.x"},{"key":"S0967199419000595_ref73","doi-asserted-by":"publisher","DOI":"10.1007\/s10815-011-9631-8"},{"key":"S0967199419000595_ref44","doi-asserted-by":"publisher","DOI":"10.1007\/s00441-005-1097-5"},{"key":"S0967199419000595_ref91","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1006\/abio.1993.1104","article-title":"Rapid analysis of mammalian sperm nuclear proteins","volume":"209","author":"Yebra","year":"1993","journal-title":"Anal Biochem"},{"key":"S0967199419000595_ref37","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2605.2006.00699.x"},{"key":"S0967199419000595_ref39","doi-asserted-by":"publisher","DOI":"10.1016\/j.theriogenology.2006.04.025"},{"key":"S0967199419000595_ref95","doi-asserted-by":"publisher","DOI":"10.1016\/j.fertnstert.2005.04.044"},{"key":"S0967199419000595_ref41","doi-asserted-by":"publisher","DOI":"10.1530\/jrf.0.0640097"},{"key":"S0967199419000595_ref94","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1007\/978-1-4419-6857-9","volume-title":"Sperm chromatin. Biological and clinical applications in male infertility and assisted reproduction","author":"Zini","year":"2011"},{"key":"S0967199419000595_ref42","doi-asserted-by":"publisher","DOI":"10.1016\/S0015-0282(03)01116-6"},{"key":"S0967199419000595_ref43","doi-asserted-by":"publisher","DOI":"10.1093\/mutage\/gem052"},{"key":"S0967199419000595_ref46","doi-asserted-by":"publisher","DOI":"10.1016\/j.rbmo.2015.05.006"},{"key":"S0967199419000595_ref45","doi-asserted-by":"publisher","DOI":"10.3109\/14647273.2010.528823"},{"key":"S0967199419000595_ref47","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/18.5.1249"},{"key":"S0967199419000595_ref58","doi-asserted-by":"publisher","DOI":"10.1093\/molehr\/gat090"},{"key":"S0967199419000595_ref48","doi-asserted-by":"publisher","DOI":"10.1530\/rep.0.1250625"},{"key":"S0967199419000595_ref79","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-62703-038-0_13"},{"key":"S0967199419000595_ref83","doi-asserted-by":"publisher","DOI":"10.1111\/ahe.12361"},{"key":"S0967199419000595_ref49","first-page":"109","article-title":"Chromatin structure function during mammalian spermatogenesis: DNA nicking repair in elongating spermatids","volume":"37","author":"McPherson","year":"1993","journal-title":"Eur J Histochem"},{"key":"S0967199419000595_ref2","doi-asserted-by":"publisher","DOI":"10.1016\/S0015-0282(02)04948-8"},{"key":"S0967199419000595_ref57","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0044679"},{"key":"S0967199419000595_ref24","doi-asserted-by":"publisher","DOI":"10.1002\/j.1939-4640.2002.tb02599.x"},{"key":"S0967199419000595_ref59","doi-asserted-by":"publisher","DOI":"10.1111\/andr.12334"},{"key":"S0967199419000595_ref60","first-page":"65","article-title":"The use of acridine orange and toluidine blue in the evaluation of male fertility","volume":"18","author":"Rocha","year":"2002","journal-title":"Biosci J"},{"key":"S0967199419000595_ref61","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2605.1985.tb00828.x"},{"key":"S0967199419000595_ref19","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2605.2006.00694.x"},{"key":"S0967199419000595_ref63","doi-asserted-by":"publisher","DOI":"10.1016\/j.theriogenology.2016.11.001"},{"key":"S0967199419000595_ref64","doi-asserted-by":"publisher","DOI":"10.1016\/j.fertnstert.2009.10.046"},{"key":"S0967199419000595_ref66","doi-asserted-by":"publisher","DOI":"10.1016\/S0015-0282(03)00337-6"},{"key":"S0967199419000595_ref31","doi-asserted-by":"publisher","DOI":"10.2164\/jandrol.106.000315"},{"key":"S0967199419000595_ref67","doi-asserted-by":"publisher","DOI":"10.1016\/j.anireprosci.2008.05.126"},{"key":"S0967199419000595_ref69","doi-asserted-by":"publisher","DOI":"10.1530\/jrf.0.1010435"},{"key":"S0967199419000595_ref72","doi-asserted-by":"publisher","DOI":"10.1016\/j.theriogenology.2018.05.023"},{"key":"S0967199419000595_ref76","doi-asserted-by":"publisher","DOI":"10.1016\/j.urology.2010.04.036"},{"key":"S0967199419000595_ref84","doi-asserted-by":"publisher","DOI":"10.1093\/humrep\/dei169"},{"key":"S0967199419000595_ref85","doi-asserted-by":"publisher","DOI":"10.1038\/aja.2011.59"},{"key":"S0967199419000595_ref86","doi-asserted-by":"publisher","DOI":"10.1093\/humrep\/17.1.184"},{"key":"S0967199419000595_ref88","doi-asserted-by":"publisher","DOI":"10.1095\/biolreprod44.4.569"},{"key":"S0967199419000595_ref89","doi-asserted-by":"publisher","DOI":"10.1016\/j.repbio.2015.03.003"},{"key":"S0967199419000595_ref92","doi-asserted-by":"publisher","DOI":"10.1016\/j.theriogenology.2013.01.008"},{"key":"S0967199419000595_ref74","doi-asserted-by":"publisher","DOI":"10.1016\/j.mefs.2013.02.001"},{"key":"S0967199419000595_ref40","doi-asserted-by":"publisher","DOI":"10.1016\/j.theriogenology.2016.08.017"},{"key":"S0967199419000595_ref53","doi-asserted-by":"publisher","DOI":"10.1007\/BF00160507"},{"key":"S0967199419000595_ref71","doi-asserted-by":"publisher","DOI":"10.1016\/j.theriogenology.2017.03.005"},{"key":"S0967199419000595_ref35","doi-asserted-by":"publisher","DOI":"10.1530\/REP-15-0403"},{"key":"S0967199419000595_ref90","doi-asserted-by":"publisher","DOI":"10.1016\/S0300-9084(98)80018-7"},{"key":"S0967199419000595_ref51","doi-asserted-by":"publisher","DOI":"10.1093\/humupd\/dml009"},{"key":"S0967199419000595_ref38","doi-asserted-by":"publisher","DOI":"10.1590\/S1413-70542010000100027"},{"key":"S0967199419000595_ref55","doi-asserted-by":"publisher","DOI":"10.3390\/genes10020105"},{"key":"S0967199419000595_ref34","doi-asserted-by":"publisher","DOI":"10.1111\/andr.12610"}],"container-title":["Zygote"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.cambridge.org\/core\/services\/aop-cambridge-core\/content\/view\/S0967199419000595","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,7,24]],"date-time":"2024-07-24T21:50:36Z","timestamp":1721857836000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.cambridge.org\/core\/product\/identifier\/S0967199419000595\/type\/journal_article"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,10,11]]},"references-count":95,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2020,2]]}},"alternative-id":["S0967199419000595"],"URL":"https:\/\/doi.org\/10.1017\/s0967199419000595","relation":{},"ISSN":["0967-1994","1469-8730"],"issn-type":[{"value":"0967-1994","type":"print"},{"value":"1469-8730","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,10,11]]}}}