{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,21]],"date-time":"2026-04-21T07:24:03Z","timestamp":1776756243281,"version":"3.51.2"},"reference-count":24,"publisher":"Oxford University Press (OUP)","issue":"3","license":[{"start":{"date-parts":[[2020,9,1]],"date-time":"2020-09-01T00:00:00Z","timestamp":1598918400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"content-domain":{"domain":["rep.bioscientifica.com"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2020,9,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>In this study, we investigated a gene-edited (Zp2MT\/MT) rat model of infertility caused by the failure to express the zona pellucida glycoprotein 2 (ZP2) due to the significant reduction of mRNA amount. We examined the defects in the zona pellucida (ZP) caused by ZP2 nullification and the influence of these defects on aspects of oocyte development, including apoptosis and fertilization ability. To investigate the cause of the influence to the oocytes\u2019 development, we evaluated the morphology of follicular transzonal projections (TZPs), known as \u2018bridges\u2019, which mediate the bidirectional signaling between the oocyte and surrounding granulosa cells and the level of reactive oxygen species (ROS) in ovulated eggs. Our results showed that two types of ZP defects were generated in the Zp2MT\/MT rat,that is, ZP intact but thinned and ZP cracked (or even absent). The fertilization rate of the ovulated eggs reduced in both types, while increased oocyte apoptosis was observed only in the latter type. Moreover, the increased oocyte apoptosis rate correlated closely with the reduction in follicular TZPs and increased ROS levels in ovulated egg. In conclusion, nullification of rat ZP2 destroyed the integrity of the ZP, impaired the bidirectional signaling between the oocyte and surrounding granulosa cells. Therefore, the resulting infertility likely occurs via elevation of oxidative stress and oocytes apoptosis.<\/jats:p>","DOI":"10.1530\/rep-20-0037","type":"journal-article","created":{"date-parts":[[2020,6,9]],"date-time":"2020-06-09T12:29:41Z","timestamp":1591705781000},"page":"353-365","update-policy":"https:\/\/doi.org\/10.1530\/crossmarkpolicy-11","source":"Crossref","is-referenced-by-count":9,"title":["Mutation of rat\n                    <i>Zp2<\/i>\n                    causes ROS-mediated oocyte apoptosis"],"prefix":"10.1093","volume":"160","author":[{"given":"Yan","family":"Wang","sequence":"first","affiliation":[{"name":"Institute of Reproductive & Stem Cell Engineering , School of Basic Medical Science, Central South University, Changsha, China"},{"name":"Center of Reproductive Health , School of Basic Medical Science, Central South University, Changsha, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hualin","family":"Huang","sequence":"additional","affiliation":[{"name":"Institute of Reproductive & Stem Cell Engineering , School of Basic Medical Science, Central South University, Changsha, China"},{"name":"Center of Reproductive Health , School of Basic Medical Science, Central South University, Changsha, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Minghua","family":"Zeng","sequence":"additional","affiliation":[{"name":"Institute of Reproductive & Stem Cell Engineering , School of Basic Medical Science, Central South University, Changsha, China"},{"name":"Center of Reproductive Health , School of Basic Medical Science, Central South University, Changsha, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ru-Ping","family":"Quan","sequence":"additional","affiliation":[{"name":"Institute of Reproductive & Stem Cell Engineering , School of Basic Medical Science, Central South University, Changsha, China"},{"name":"Center of Reproductive Health , School of Basic Medical Science, Central South University, Changsha, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jun-Ting","family":"Yang","sequence":"additional","affiliation":[{"name":"Institute of Reproductive & Stem Cell Engineering , School of Basic Medical Science, Central South University, Changsha, China"},{"name":"Center of Reproductive Health , School of Basic Medical Science, Central South University, Changsha, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Dan","family":"Guo","sequence":"additional","affiliation":[{"name":"Institute of Reproductive & Stem Cell Engineering , School of Basic Medical Science, Central South University, Changsha, China"},{"name":"Center of Reproductive Health , School of Basic Medical Science, Central South University, Changsha, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ying","family":"Sun","sequence":"additional","affiliation":[{"name":"Institute of Reproductive & Stem Cell Engineering , School of Basic Medical Science, Central South University, Changsha, China"},{"name":"Center of Reproductive Health , School of Basic Medical Science, Central South University, Changsha, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hongwen","family":"Deng","sequence":"additional","affiliation":[{"name":"Center for Bioinformatics and Genomics , Department of Global Biostatistics and Data Science, School of Public Health and Tropical Medicine, Tulane University, New Orleans, Louisiana, USA"},{"name":"Center of System Biology and Data Information , School of Basic Medical Science, Central South University, Changsha, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8121-9498","authenticated-orcid":false,"given":"Hongmei","family":"Xiao","sequence":"additional","affiliation":[{"name":"Institute of Reproductive & Stem Cell Engineering , School of Basic Medical Science, Central South University, Changsha, China"},{"name":"Center of Reproductive Health , School of Basic Medical Science, Central South University, Changsha, China"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"286","published-online":{"date-parts":[[2020,9,1]]},"reference":[{"key":"2026010712240998800_bib1","doi-asserted-by":"publisher","first-page":"801","DOI":"10.1083\/jcb.201404025","article-title":"A single domain of the ZP2 zona pellucida protein mediates gamete recognition in mice and humans","volume":"205","author":"Avella","year":"2014","journal-title":"Journal of Cell Biology"},{"key":"2026010712240998800_bib2","doi-asserted-by":"publisher","first-page":"459","DOI":"10.1016\/j.ajhg.2017.08.001","article-title":"A recurrent missense mutation in zp3 causes empty follicle syndrome and female infertility","volume":"101","author":"Chen","year":"2017","journal-title":"American Journal of Human Genetics"},{"key":"2026010712240998800_bib3","doi-asserted-by":"publisher","first-page":"2201","DOI":"10.1093\/humrep\/dez174","article-title":"ZP1 mutations are associated with empty follicle syndrome: evidence for the existence of an intact oocyte and a zona pellucida in follicles up to the early antral stage. A case report","volume":"34","author":"Dai","year":"2019","journal-title":"Human Reproduction"},{"key":"2026010712240998800_bib4","doi-asserted-by":"publisher","first-page":"431","DOI":"10.1038\/s41436-018-0064-y","article-title":"ZP2 pathogenic variants cause in vitro fertilization failure and female infertility","volume":"21","author":"Dai","year":"2019","journal-title":"Genetics in Medicine"},{"key":"2026010712240998800_bib5","doi-asserted-by":"publisher","first-page":"e17","DOI":"10.4081\/ejh.2012.17","article-title":"Immunohistochemical and ultrastructural visualization of different routes of oocyte elimination in adult rats","volume":"56","author":"Escobar","year":"2012","journal-title":"European Journal of Histochemistry"},{"key":"2026010712240998800_bib6","doi-asserted-by":"publisher","first-page":"394","DOI":"10.1093\/humrep\/dem373","article-title":"Detection of zona pellucida proteins during human folliculogenesis","volume":"23","author":"Gook","year":"2008","journal-title":"Human Reproduction and Embryology"},{"key":"2026010712240998800_bib7","doi-asserted-by":"publisher","first-page":"665","DOI":"10.1007\/s00441-011-1319-y","article-title":"Mammalian zona pellucida glycoproteins: structure and function during fertilization","volume":"349","author":"Gupta","year":"2012","journal-title":"Cell and Tissue Research"},{"key":"2026010712240998800_bib8","doi-asserted-by":"publisher","first-page":"1220","DOI":"10.1056\/NEJMoa1308851","article-title":"Mutant ZP1 in familial infertility","volume":"370","author":"Huang","year":"2014","journal-title":"New England Journal of Medicine"},{"key":"2026010712240998800_bib9","doi-asserted-by":"publisher","first-page":"1335","DOI":"10.3390\/ijms21041335","article-title":"Mechanisms and regulation of nonsense-mediated mRNA decay and nonsense-associated altered splicing in lymphocytes","volume":"21","author":"Lambert","year":"2020","journal-title":"International Journal of Molecular Sciences"},{"key":"2026010712240998800_bib10","doi-asserted-by":"publisher","first-page":"e48904","DOI":"10.7554\/eLife.48904","article-title":"ZP4 confers structural properties to the zona pellucida essential for embryo development","volume":"8","author":"Lamas-Toranzo","year":"2019","journal-title":"eLife"},{"key":"2026010712240998800_bib11","doi-asserted-by":"publisher","first-page":"5431","DOI":"10.1073\/pnas.93.11.5431","article-title":"Targeted disruption of the mZP3 gene results in production of eggs lacking a zona pellucida and infertility in female mice","volume":"93","author":"Liu","year":"1996","journal-title":"PNAS"},{"key":"2026010712240998800_bib12","doi-asserted-by":"publisher","first-page":"975","DOI":"10.1007\/s00439-017-1822-7","article-title":"Dosage effects of ZP2 and ZP3 heterozygous mutations cause human infertility","volume":"136","author":"Liu","year":"2017","journal-title":"Human Genetics"},{"key":"2026010712240998800_bib13","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1089\/biores.2014.0013","article-title":"A moderate increase of hydrogen peroxide level is beneficial for spontaneous resumption of meiosis from diplotene arrest in rat oocytes cultured in vitro","volume":"3","author":"Pandey","year":"2014","journal-title":"BioResearch Open Access"},{"key":"2026010712240998800_bib14","doi-asserted-by":"crossref","first-page":"3847","DOI":"10.1242\/dev.126.17.3847","article-title":"Abnormal zonae pellucidae in mice lacking ZP1 result in early embryonic loss","volume":"126","author":"Rankin","year":"1999","journal-title":"Development"},{"key":"2026010712240998800_bib15","doi-asserted-by":"crossref","first-page":"1119","DOI":"10.1242\/dev.128.7.1119","article-title":"Defective zonae pellucidae in Zp2-null mice disrupt folliculogenesis, fertility and development","volume":"128","author":"Rankin","year":"2001","journal-title":"Development"},{"key":"2026010712240998800_bib16","doi-asserted-by":"publisher","first-page":"525","DOI":"10.1038\/385525a0","article-title":"Female infertility in mice lacking connexin 37","volume":"385","author":"Simon","year":"1997","journal-title":"Nature"},{"key":"2026010712240998800_bib17","doi-asserted-by":"publisher","first-page":"2001","DOI":"10.1002\/humu.23864","article-title":"Compound heterozygous ZP1 mutations cause empty follicle syndrome in infertile sisters","volume":"40","author":"Sun","year":"2019","journal-title":"Human Mutation"},{"key":"2026010712240998800_bib18","doi-asserted-by":"publisher","first-page":"78","DOI":"10.1095\/biolreprod53.1.78","article-title":"Developmental ability of zona-free mouse embryos is influenced by cell association at the 4-cell stage","volume":"53","author":"Suzuki","year":"1995","journal-title":"Biology of Reproduction"},{"key":"2026010712240998800_bib19","doi-asserted-by":"publisher","first-page":"1019","DOI":"10.1007\/s10495-015-1136-y","article-title":"Apoptosis in mammalian oocytes: a review","volume":"20","author":"Tiwari","year":"2015","journal-title":"Apoptosis"},{"key":"2026010712240998800_bib20","doi-asserted-by":"publisher","first-page":"457","DOI":"10.1093\/biolre\/ioz093","article-title":"Influence of mouse defective zona pellucida in folliculogenesis on apoptosis of granulosa cells and developmental competence of oocytesdagger","volume":"101","author":"Wang","year":"2019","journal-title":"Biology of Reproduction"},{"key":"2026010712240998800_bib21","doi-asserted-by":"publisher","first-page":"24285","DOI":"10.1074\/jbc.R800027200","article-title":"Zona pellucida glycoproteins","volume":"283","author":"Wassarman","year":"2008","journal-title":"Journal of Biological Chemistry"},{"key":"2026010712240998800_bib22","doi-asserted-by":"publisher","first-page":"833","DOI":"10.1387\/ijdb.120136pw","article-title":"Influence of the zona pellucida of the mouse egg on folliculogenesis and fertility","volume":"56","author":"Wassarman","year":"2012","journal-title":"International Journal of Developmental Biology"},{"key":"2026010712240998800_bib23","doi-asserted-by":"publisher","first-page":"323","DOI":"10.1098\/rspb.1997.0046","article-title":"Mutant female mice carrying a single mZP3 allele produce eggs with a thin zona pellucida, but reproduce normally","volume":"264","author":"Wassarman","year":"1997","journal-title":"Proceedings Biological Sciences"},{"key":"2026010712240998800_bib24","doi-asserted-by":"publisher","first-page":"327","DOI":"10.1007\/s00439-019-01990-1","article-title":"Novel mutations in ZP1, ZP2, and ZP3 cause female infertility due to abnormal zona pellucida formation","volume":"138","author":"Zhou","year":"2019","journal-title":"Human Genetics"}],"container-title":["Reproduction"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/rep.bioscientifica.com\/view\/journals\/rep\/160\/3\/REP-20-0037.xml","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/rep.bioscientifica.com\/downloadpdf\/journals\/rep\/160\/3\/REP-20-0037.xml","content-type":"text\/html","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/reproduction\/article-pdf\/160\/3\/353\/64127212\/rep-20-0037.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/reproduction\/article-pdf\/160\/3\/353\/64127212\/rep-20-0037.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,7]],"date-time":"2026-01-07T17:24:15Z","timestamp":1767806655000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/reproduction\/article\/160\/3\/353\/8235605"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,9,1]]},"references-count":24,"journal-issue":{"issue":"3","published-print":{"date-parts":[[2020,9,1]]}},"URL":"https:\/\/doi.org\/10.1530\/rep-20-0037","relation":{},"ISSN":["1470-1626","1741-7899"],"issn-type":[{"value":"1470-1626","type":"print"},{"value":"1741-7899","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2020,9,1]]},"published":{"date-parts":[[2020,9,1]]}}}