{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T19:20:06Z","timestamp":1786994406056,"version":"build-2736575974"},"reference-count":94,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2025,7,1]],"date-time":"2025-07-01T00:00:00Z","timestamp":1751328000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2025,7,1]],"date-time":"2025-07-01T00:00:00Z","timestamp":1751328000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2025,7,1]],"date-time":"2025-07-01T00:00:00Z","timestamp":1751328000000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-017"},{"start":{"date-parts":[[2025,7,1]],"date-time":"2025-07-01T00:00:00Z","timestamp":1751328000000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2025,7,1]],"date-time":"2025-07-01T00:00:00Z","timestamp":1751328000000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-012"},{"start":{"date-parts":[[2025,7,1]],"date-time":"2025-07-01T00:00:00Z","timestamp":1751328000000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2025,7,1]],"date-time":"2025-07-01T00:00:00Z","timestamp":1751328000000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-004"}],"funder":[{"DOI":"10.13039\/501100001409","name":"Department of Science and Technology","doi-asserted-by":"publisher","award":["SPG\/2021\/002635"],"award-info":[{"award-number":["SPG\/2021\/002635"]}],"id":[{"id":"10.13039\/501100001409","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001412","name":"CSIR","doi-asserted-by":"publisher","award":["09\/254(0274)\/2017-EMR-I"],"award-info":[{"award-number":["09\/254(0274)\/2017-EMR-I"]}],"id":[{"id":"10.13039\/501100001412","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Mutation Research - Reviews in Mutation Research"],"published-print":{"date-parts":[[2025,7]]},"DOI":"10.1016\/j.mrrev.2025.108560","type":"journal-article","created":{"date-parts":[[2025,9,2]],"date-time":"2025-09-02T16:09:24Z","timestamp":1756829364000},"page":"108560","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":2,"special_numbering":"C","title":["Genetic variations in zona pellucida glycoproteins: Implications for fertility and ART outcomes"],"prefix":"10.1016","volume":"796","author":[{"given":"Neha","family":"Rajput","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0058-2827","authenticated-orcid":false,"given":"Gagandeep Kaur","family":"Gahlay","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.mrrev.2025.108560_bib1","doi-asserted-by":"crossref","first-page":"904","DOI":"10.1007\/s43032-020-00306-3","article-title":"Can oocyte diameter predict embryo quality?","volume":"28","author":"Bassil","year":"2021","journal-title":"Reprod. Sci."},{"key":"10.1016\/j.mrrev.2025.108560_bib2","doi-asserted-by":"crossref","first-page":"1596","DOI":"10.1093\/humrep\/deh811","article-title":"High magnitude of light retardation by the zona pellucida is associated with conception cycles","volume":"20","author":"Shen","year":"2005","journal-title":"Hum. Reprod."},{"key":"10.1016\/j.mrrev.2025.108560_bib3","doi-asserted-by":"crossref","first-page":"695","DOI":"10.1146\/annurev-biochem-011520-105310","article-title":"Zona pellucida proteins, fibrils, and matrix","volume":"89","author":"Litscher","year":"2020","journal-title":"Annu Rev. Biochem."},{"key":"10.1016\/j.mrrev.2025.108560_bib4","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1023\/A:1012744617241","article-title":"The zona pellucida in folliculogenesis, fertilization and early development","volume":"3","author":"Zhao","year":"2002","journal-title":"Rev. Endocr. Metab. Disord."},{"key":"10.1016\/j.mrrev.2025.108560_bib5","doi-asserted-by":"crossref","first-page":"413","DOI":"10.1016\/bs.ctdb.2018.02.007","article-title":"Structure of zona pellucida module proteins","volume":"130","author":"Bokhove","year":"2018","journal-title":"Curr. Top. Dev. Biol."},{"key":"10.1016\/j.mrrev.2025.108560_bib6","doi-asserted-by":"crossref","first-page":"1580","DOI":"10.1093\/humrep\/deh301","article-title":"Four zona pellucida glycoproteins are expressed in the human","volume":"19","author":"Lefi\u00e8vre","year":"2004","journal-title":"Hum. Reprod."},{"key":"10.1016\/j.mrrev.2025.108560_bib7","doi-asserted-by":"crossref","first-page":"1440","DOI":"10.1016\/j.cell.2024.02.013","article-title":"ZP2 cleavage blocks polyspermy by modulating the architecture of the egg coat","volume":"187","author":"Nishio","year":"2024","journal-title":"Cell"},{"key":"10.1016\/j.mrrev.2025.108560_bib8","doi-asserted-by":"crossref","first-page":"2903","DOI":"10.1242\/dev.122.9.2903","article-title":"Mice homozygous for an insertional mutation in the Zp3 gene lack a zona pellucida and are infertile","volume":"122","author":"Rankin","year":"1996","journal-title":"Development"},{"key":"10.1016\/j.mrrev.2025.108560_bib9","doi-asserted-by":"crossref","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":"Lui","year":"1996","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.mrrev.2025.108560_bib10","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":"10.1016\/j.mrrev.2025.108560_bib11","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":"10.1016\/j.mrrev.2025.108560_bib12","doi-asserted-by":"crossref","first-page":"1282","DOI":"10.1093\/biolre\/ioab047","article-title":"Zp4 is completely dispensable for fertility in female rats","volume":"104","author":"Zeng","year":"2021","journal-title":"Biol. Reprod."},{"key":"10.1016\/j.mrrev.2025.108560_bib13","doi-asserted-by":"crossref","first-page":"1871","DOI":"10.1093\/bioinformatics\/btm265","article-title":"Isolated ZP-N domains constitute the N-terminal extensions of zona pellucida proteins","volume":"23","author":"Callebaut","year":"2007","journal-title":"Bioinformatics"},{"key":"10.1016\/j.mrrev.2025.108560_bib14","doi-asserted-by":"crossref","first-page":"24285","DOI":"10.1074\/jbc.R800027200","article-title":"Zona pellucida glycoproteins","volume":"283","author":"Wassarman","year":"2008","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.mrrev.2025.108560_bib15","doi-asserted-by":"crossref","first-page":"3111","DOI":"10.1128\/MCB.22.9.3111-3120.2002","article-title":"Conserved furin cleavage site not essential for secretion and integration of ZP3 into the extracellular egg coat of transgenic mice","volume":"22","author":"Zhao","year":"2002","journal-title":"Mol. Cell Biol."},{"key":"10.1016\/j.mrrev.2025.108560_bib16","doi-asserted-by":"crossref","first-page":"8982","DOI":"10.1128\/MCB.23.24.8982-8991.2003","article-title":"Mutation of a conserved hydrophobic patch prevents incorporation of ZP3 into the zona pellucida surrounding mouse eggs","volume":"23","author":"Zhao","year":"2003","journal-title":"Mol. Cell Biol."},{"key":"10.1016\/j.mrrev.2025.108560_bib17","doi-asserted-by":"crossref","first-page":"886","DOI":"10.1039\/c6ib00044d","article-title":"Investigating the mechanical properties of zona pellucida of whole human oocytes by atomic force spectroscopy","volume":"8","author":"Andolfi","year":"2016","journal-title":"Integr. Biol. (Camb. )"},{"key":"10.1016\/j.mrrev.2025.108560_bib18","doi-asserted-by":"crossref","first-page":"20130066","DOI":"10.1098\/rsfs.2013.0066","article-title":"A hybrid characterization framework to determine the visco-hyperelastic properties of a porcine zona pellucida","volume":"4","author":"Boccaccio","year":"2014","journal-title":"Interface Focus"},{"key":"10.1016\/j.mrrev.2025.108560_bib19","doi-asserted-by":"crossref","first-page":"569","DOI":"10.1109\/TBME.2012.2230444","article-title":"Viscoelastic characterization of mouse zona pellucida","volume":"60","author":"Kiim","year":"2013","journal-title":"IEEE Trans. Biomed. Eng."},{"key":"10.1016\/j.mrrev.2025.108560_bib20","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1016\/S0021-9290(03)00242-2","article-title":"Micro-mechanical sensing platform for the characterization of the elastic properties of the ovum via uniaxial measurement","volume":"37","author":"Murayama","year":"2004","journal-title":"J. Biomech."},{"key":"10.1016\/j.mrrev.2025.108560_bib21","doi-asserted-by":"crossref","first-page":"987","DOI":"10.1007\/s00249-009-0468-3","article-title":"Mechanical properties of zona pellucida hardening","volume":"39","author":"Papi","year":"2010","journal-title":"Eur. Biophys. J."},{"key":"10.1016\/j.mrrev.2025.108560_bib22","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/0012-1606(81)90329-8","article-title":"Mammalian sperm-egg interaction: fertilization of mouse eggs triggers modification of the major zona pellucida glycoprotein, ZP2","volume":"86","author":"Bleil","year":"1981","journal-title":"Dev. Biol."},{"issue":"1979","key":"10.1016\/j.mrrev.2025.108560_bib23","doi-asserted-by":"crossref","first-page":"216","DOI":"10.1126\/science.1188178","article-title":"Gamete recognition in mice depends on the cleavage status of an egg\u2019s zona pellucida protein","volume":"329","author":"Gahlay","year":"2010","journal-title":"Science"},{"key":"10.1016\/j.mrrev.2025.108560_bib24","doi-asserted-by":"crossref","first-page":"897","DOI":"10.1083\/jcb.201203062","article-title":"Human sperm bind to the N-terminal domain of ZP2 in humanized zonae pellucidae in transgenic mice","volume":"197","author":"Baibakov","year":"2012","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.mrrev.2025.108560_bib25","doi-asserted-by":"crossref","first-page":"404","DOI":"10.1016\/j.cell.2010.09.041","article-title":"Insights into egg coat assembly and egg-sperm interaction from the X-ray structure of full-length ZP3","volume":"143","author":"Han","year":"2010","journal-title":"Cell"},{"key":"10.1016\/j.mrrev.2025.108560_bib26","doi-asserted-by":"crossref","first-page":"762","DOI":"10.3390\/ijms23020762","article-title":"Identification of Sperm-Binding sites in the N-Terminal domain of bovine egg coat glycoprotein ZP4","volume":"23","author":"Dilimulati","year":"2022","journal-title":"Int J. Mol. Sci."},{"key":"10.1016\/j.mrrev.2025.108560_bib27","doi-asserted-by":"crossref","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":"10.1016\/j.mrrev.2025.108560_bib28","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1111\/j.1600-0897.2012.01177.x","article-title":"Mapping of epitopes relevant for induction of acrosome reaction on human zona pellucida Glycoprotein-4 using monoclonal antibodies","volume":"68","author":"Xu","year":"2012","journal-title":"Am. J. Reprod. Immunol."},{"key":"10.1016\/j.mrrev.2025.108560_bib29","doi-asserted-by":"crossref","DOI":"10.3390\/genes12081266","article-title":"Zona pellucida genes and proteins: essential players in mammalian oogenesis and fertility","volume":"12","author":"Wassarman","year":"2021","journal-title":"Genes (Basel)"},{"key":"10.1016\/j.mrrev.2025.108560_bib30","doi-asserted-by":"crossref","first-page":"847","DOI":"10.1016\/j.rbmo.2023.01.010","article-title":"Novel variants in ZP1, ZP2 and ZP3 associated with empty follicle syndrome and abnormal zona pellucida","volume":"46","author":"Sun","year":"2023","journal-title":"Reprod. Biomed. Online"},{"key":"10.1016\/j.mrrev.2025.108560_bib31","doi-asserted-by":"crossref","DOI":"10.1186\/s12958-022-01046-6","article-title":"Can successful pregnancy be achieved and predicted from patients with identified ZP mutations? A literature review","volume":"20","author":"Zhou","year":"2022","journal-title":"Reprod. Biol. Endocrinol."},{"key":"10.1016\/j.mrrev.2025.108560_bib32","first-page":"522","article-title":"The human zona pellucida","volume":"39","author":"Trocka","year":"2023","journal-title":"N. Genet. Find. Med. \/Sci."},{"key":"10.1016\/j.mrrev.2025.108560_bib33","doi-asserted-by":"crossref","DOI":"10.3389\/fcell.2021.763166","article-title":"Identification and in silico characterization of deleterious single nucleotide variations in human ZP2 gene","volume":"9","author":"Rajput","year":"2021","journal-title":"Front Cell Dev. Biol."},{"key":"10.1016\/j.mrrev.2025.108560_bib34","doi-asserted-by":"crossref","DOI":"10.1002\/0471142905.hg0720s76","article-title":"Predicting functional effect of human missense mutations using PolyPhen-2","author":"Adzhubei","year":"2013","journal-title":"Curr. Protoc. Hum. Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib35","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1038\/nmeth0410-248","article-title":"A method and server for predicting damaging missense mutations","volume":"7","author":"Adzhubei","year":"2010","journal-title":"Nat. Methods"},{"issue":"8","key":"10.1016\/j.mrrev.2025.108560_bib36","article-title":"Better prediction of functional effects for sequence variants","volume":"16","author":"Hecht","year":"2015","journal-title":"BMC Genom."},{"key":"10.1016\/j.mrrev.2025.108560_bib37","doi-asserted-by":"crossref","first-page":"2729","DOI":"10.1093\/bioinformatics\/btl423","article-title":"Predicting the insurgence of human genetic diseases associated to single point protein mutations with support vector machines and evolutionary information","volume":"22","author":"Capriotti","year":"2006","journal-title":"Bioinformatics"},{"key":"10.1016\/j.mrrev.2025.108560_bib38","doi-asserted-by":"crossref","first-page":"1237","DOI":"10.1002\/humu.21047","article-title":"Functional annotations improve the predictive score of human disease-related mutations in proteins","volume":"30","author":"Calabrese","year":"2009","journal-title":"Hum. Mutat."},{"issue":"3","key":"10.1016\/j.mrrev.2025.108560_bib39","doi-asserted-by":"crossref","first-page":"S6","DOI":"10.1186\/1471-2164-14-S3-S6","article-title":"WS-SNPs&GO: a web server for predicting the deleterious effect of human protein variants using functional annotation","volume":"14","author":"Capriotti","year":"2013","journal-title":"BMC Genom."},{"key":"10.1016\/j.mrrev.2025.108560_bib40","doi-asserted-by":"crossref","first-page":"1551","DOI":"10.1038\/nprot.2013.092","article-title":"Large-scale gene function analysis with the PANTHER classification system","volume":"8","author":"Mi","year":"2013","journal-title":"Nat. Protoc."},{"key":"10.1016\/j.mrrev.2025.108560_bib41","doi-asserted-by":"crossref","first-page":"5918","DOI":"10.1038\/s41467-020-19669-x","article-title":"Inferring the molecular and phenotypic impact of amino acid variants with MutPred2","volume":"11","author":"Pejaver","year":"2020","journal-title":"Nat. Commun."},{"key":"10.1016\/j.mrrev.2025.108560_bib42","doi-asserted-by":"crossref","DOI":"10.1093\/nar\/gki375","article-title":"I-Mutant2.0: predicting stability changes upon mutation from the protein sequence or structure","volume":"33","author":"Capriotti","year":"2005","journal-title":"Nucleic Acids Res"},{"key":"10.1016\/j.mrrev.2025.108560_bib43","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1038\/s41586-021-03819-2","article-title":"Highly accurate protein structure prediction with AlphaFold","volume":"596","author":"Jumper","year":"2021","journal-title":"Nature"},{"key":"10.1016\/j.mrrev.2025.108560_bib44","doi-asserted-by":"crossref","first-page":"1605","DOI":"10.1002\/jcc.20084","article-title":"UCSF Chimera\u2014A visualization system for exploratory research and analysis","volume":"25","author":"Pettersen","year":"2004","journal-title":"J. Comput. Chem."},{"key":"10.1016\/j.mrrev.2025.108560_bib45","doi-asserted-by":"crossref","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":"Hum. Mutat."},{"key":"10.1016\/j.mrrev.2025.108560_bib46","doi-asserted-by":"crossref","first-page":"741","DOI":"10.1007\/s10815-019-01404-1","article-title":"Novel mutation in the ZP1 gene and clinical implications","volume":"36","author":"Yuan","year":"2019","journal-title":"J. Assist Reprod. Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib47","doi-asserted-by":"crossref","first-page":"1259","DOI":"10.1016\/j.fertnstert.2020.11.003","article-title":"The critical role of ZP genes in female infertility characterized by empty follicle syndrome and oocyte degeneration","volume":"115","author":"Yang","year":"2021","journal-title":"Fertil. Steril."},{"key":"10.1016\/j.mrrev.2025.108560_bib48","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1111\/cge.13921","article-title":"Novel mutations in ZP1: expanding the mutational spectrum associated with empty follicle syndrome in infertile women","volume":"99","author":"Wu","year":"2021","journal-title":"Clin. Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib49","doi-asserted-by":"crossref","first-page":"1220","DOI":"10.1056\/NEJMoa1308851","article-title":"Mutant ZP1 in familial infertility","volume":"370","author":"Huang","year":"2014","journal-title":"N. Engl. J. Med."},{"key":"10.1016\/j.mrrev.2025.108560_bib50","doi-asserted-by":"crossref","first-page":"3086","DOI":"10.1038\/s41467-019-10931-5","article-title":"Molecular basis of egg coat cross-linking sheds light on ZP1-associated female infertility","volume":"10","author":"Nishimura","year":"2019","journal-title":"Nat. Commun."},{"key":"10.1016\/j.mrrev.2025.108560_bib51","doi-asserted-by":"crossref","first-page":"1459","DOI":"10.1007\/s10815-021-02136-x","article-title":"A novel homozygous nonsense mutation in zona pellucida 1 (ZP1) causes human female empty follicle syndrome","volume":"38","author":"Wang","year":"2021","journal-title":"J. Assist Reprod. Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib52","doi-asserted-by":"crossref","DOI":"10.3389\/fgene.2022.772143","article-title":"Gene spectrum and clinical traits of nine patients with oocyte maturation arrest","volume":"13","author":"Huo","year":"2022","journal-title":"Front Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib53","doi-asserted-by":"crossref","first-page":"183","DOI":"10.4103\/2096-2924.248486","article-title":"Infertility due to lack of zona pellucida caused by a compound heterozygous mutation in ZP1 gene","volume":"2","author":"Zhang","year":"2018","journal-title":"Reprod. Dev. Med."},{"key":"10.1016\/j.mrrev.2025.108560_bib54","doi-asserted-by":"crossref","first-page":"2151","DOI":"10.1007\/s10815-020-01855-x","article-title":"Novel biallelic loss-of-function variants in ZP1 identified in an infertile female with empty follicle syndrome","volume":"37","author":"Liu","year":"2020","journal-title":"J. Assist Reprod. Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib55","doi-asserted-by":"crossref","first-page":"1364","DOI":"10.1016\/j.fertnstert.2017.03.029","article-title":"Novel zona pellucida gene variants identified in patients with oocyte anomalies","volume":"107","author":"Yang","year":"2017","journal-title":"Fertil. Steril."},{"key":"10.1016\/j.mrrev.2025.108560_bib56","doi-asserted-by":"crossref","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":"Hum. Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib57","doi-asserted-by":"crossref","first-page":"2853","DOI":"10.1007\/s10815-020-01926-z","article-title":"Novel mutations in ZP1 and ZP2 cause primary infertility due to empty follicle syndrome and abnormal zona pellucida","volume":"37","author":"Luo","year":"2020","journal-title":"J. Assist Reprod. Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib58","first-page":"2201","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":"Hum. Reprod."},{"key":"10.1016\/j.mrrev.2025.108560_bib59","article-title":"A novel homozygous nonsense ZP1 variant causes human female infertility associated with empty follicle syndrome (EFS)","volume":"8","author":"Xu","year":"2020","journal-title":"Mol. Genet Genom. Med"},{"key":"10.1016\/j.mrrev.2025.108560_bib60","doi-asserted-by":"crossref","first-page":"74","DOI":"10.1016\/j.tibs.2009.10.001","article-title":"N-glycan structures: recognition and processing in the ER","volume":"35","author":"Aebi","year":"2010","journal-title":"Trends Biochem Sci."},{"key":"10.1016\/j.mrrev.2025.108560_bib61","doi-asserted-by":"crossref","first-page":"1019","DOI":"10.1146\/annurev.biochem.73.011303.073752","article-title":"Roles of N-Linked glycans in the endoplasmic reticulum","volume":"73","author":"Helenius","year":"2004","journal-title":"Annu Rev. Biochem"},{"key":"10.1016\/j.mrrev.2025.108560_bib62","doi-asserted-by":"crossref","first-page":"716","DOI":"10.1016\/j.semcdb.2007.09.003","article-title":"The endoplasmic reticulum and the unfolded protein response","volume":"18","author":"Malhotra","year":"2007","journal-title":"Semin Cell Dev. Biol."},{"key":"10.1016\/j.mrrev.2025.108560_bib63","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1038\/ncb802","article-title":"The ZP domain is a conserved module for polymerization of extracellular proteins","volume":"4","author":"Jovine","year":"2002","journal-title":"Nat. Cell Biol."},{"key":"10.1016\/j.mrrev.2025.108560_bib64","doi-asserted-by":"crossref","first-page":"234","DOI":"10.1111\/j.1439-0531.2009.01370.x","article-title":"Functional role of feline zona pellucida protein 4 trefoil domain: a sperm receptor or structural component of the domestic cat zona pellucida?","volume":"44","author":"Braun","year":"2009","journal-title":"Reprod. Domest. Anim."},{"key":"10.1016\/j.mrrev.2025.108560_bib65","doi-asserted-by":"crossref","first-page":"64","DOI":"10.1016\/j.cbi.2010.01.003","article-title":"Alanine-to-threonine substitutions and amyloid diseases: butyrylcholinesterase as a case study","volume":"187","author":"Podoly","year":"2010","journal-title":"Chem. Biol. Inter."},{"key":"10.1016\/j.mrrev.2025.108560_bib66","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1136\/jclinpath-2020-207170","article-title":"Mutations in ZP4 are associated with abnormal zona pellucida and female infertility","volume":"75","author":"Wei","year":"2022","journal-title":"J. Clin. Pathol."},{"key":"10.1016\/j.mrrev.2025.108560_bib67","doi-asserted-by":"crossref","first-page":"1385","DOI":"10.1093\/humrep\/den047","article-title":"Native human zona pellucida glycoproteins: purification and binding properties","volume":"23","author":"Chiu","year":"2008","journal-title":"Hum. Reprod."},{"key":"10.1016\/j.mrrev.2025.108560_bib68","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1002\/mrd.20726","article-title":"Relevance of glycosylation of human zona pellucida glycoproteins for their binding to capacitated human spermatozoa and subsequent induction of acrosomal exocytosis","volume":"75","author":"Chakravarty","year":"2008","journal-title":"Mol. Reprod. Dev."},{"key":"10.1016\/j.mrrev.2025.108560_bib69","doi-asserted-by":"crossref","first-page":"876","DOI":"10.1177\/1933719111398146","article-title":"Characterization of human sperm binding to homologous recombinant zona pellucida proteins","volume":"18","author":"Chirinos","year":"2011","journal-title":"Reprod. Sci."},{"key":"10.1016\/j.mrrev.2025.108560_bib70","doi-asserted-by":"crossref","first-page":"869","DOI":"10.1095\/biolreprod.108.069344","article-title":"Effects of native human zona pellucida glycoproteins 3 and 4 on acrosome reaction and zona pellucida binding of human spermatozoa","volume":"79","author":"Chiu","year":"2008","journal-title":"Biol. Reprod."},{"key":"10.1016\/j.mrrev.2025.108560_bib71","doi-asserted-by":"crossref","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":"10.1016\/j.mrrev.2025.108560_bib72","doi-asserted-by":"crossref","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":"Hum. Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib73","doi-asserted-by":"crossref","DOI":"10.1002\/wdev.294","article-title":"Regulation of germ cell development by intercellular signaling in the mammalian ovarian follicle","volume":"7","author":"Clarke","year":"2018","journal-title":"WIREs Dev. Biol."},{"key":"10.1016\/j.mrrev.2025.108560_bib74","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1038\/nrm3531","article-title":"The road to maturation: somatic cell interaction and self-organization of the mammalian oocyte","volume":"14","author":"Li","year":"2013","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"10.1016\/j.mrrev.2025.108560_bib75","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1006\/dbio.1998.9087","article-title":"Characterization of oocyte and follicle development in growth differentiation Factor-9-Deficient mice","volume":"204","author":"Carabatsos","year":"1998","journal-title":"Dev. Biol."},{"key":"10.1016\/j.mrrev.2025.108560_bib76","doi-asserted-by":"crossref","first-page":"1262","DOI":"10.1038\/s41598-018-37766-2","article-title":"Three-dimensional organization of transzonal projections and other cytoplasmic extensions in the mouse ovarian follicle","volume":"9","author":"Baena","year":"2019","journal-title":"Sci. Rep."},{"key":"10.1016\/j.mrrev.2025.108560_bib77","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1002\/mrd.23645","article-title":"Transzonal projections: essential structures mediating intercellular communication in the mammalian ovarian follicle","volume":"89","author":"Clarke","year":"2022","journal-title":"Mol. Reprod. Dev."},{"key":"10.1016\/j.mrrev.2025.108560_bib78","doi-asserted-by":"crossref","DOI":"10.3389\/fendo.2023.1169378","article-title":"Identification of zona pellucida defects revealed a novel loss-of-function mutation in ZP2 in humans and rats","volume":"14","author":"Zeng","year":"2023","journal-title":"Front Endocrinol. (Lausanne)"},{"key":"10.1016\/j.mrrev.2025.108560_bib79","doi-asserted-by":"crossref","first-page":"8557","DOI":"10.1111\/jcmm.15482","article-title":"Heterozygous mutations in ZP1 and ZP3 cause formation disorder of ZP and female infertility in human","volume":"24","author":"Cao","year":"2020","journal-title":"J. Cell Mol. Med"},{"key":"10.1016\/j.mrrev.2025.108560_bib80","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1017\/S0967199420000441","article-title":"Role of diagnostic intracytoplasmic sperm injection (ICSI) in the management of genetically determined zona pellucida-free oocytes during in vitro fertilization: a case report","volume":"28","author":"Metwalley","year":"2020","journal-title":"Zygote"},{"key":"10.1016\/j.mrrev.2025.108560_bib81","doi-asserted-by":"crossref","first-page":"787","DOI":"10.1111\/cge.13693","article-title":"Novel ZP1 pathogenic variants identified in an infertile patient and a successful live birth following ICSI treatment","volume":"97","author":"Chu","year":"2020","journal-title":"Clin. Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib82","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1111\/cge.14144","article-title":"A recurrent zp1 variant is responsible for oocyte maturation defect with degenerated oocytes in infertile females","volume":"102","author":"Loeuillet","year":"2022","journal-title":"Clin. Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib83","doi-asserted-by":"crossref","DOI":"10.3389\/fgene.2024.1407202","article-title":"ZP1-Y262C mutation causes abnormal zona pellucida formation and female infertility in humans","volume":"15","author":"Cao","year":"2024","journal-title":"Front Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib84","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1016\/j.ejogrb.2022.12.011","article-title":"A ZP1 gene mutation in a patient with empty follicle syndrome: a case report and literature review","volume":"280","author":"Pujalte","year":"2023","journal-title":"Eur. J. Obstet. Gynecol. Reprod. Biol."},{"key":"10.1016\/j.mrrev.2025.108560_bib85","doi-asserted-by":"crossref","first-page":"1239","DOI":"10.1007\/s10815-021-02107-2","article-title":"A novel homozygous variant in ZP2 causes abnormal zona pellucida formation and female infertility","volume":"38","author":"Sun","year":"2021","journal-title":"J. Assist Reprod. Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib86","doi-asserted-by":"crossref","first-page":"3047","DOI":"10.1007\/s43032-022-00958-3","article-title":"Novel heterozygous mutations in ZP2 cause abnormal zona pellucida and female infertility","volume":"29","author":"Hou","year":"2022","journal-title":"Reprod. Sci."},{"key":"10.1016\/j.mrrev.2025.108560_bib87","article-title":"A heterozygous ZP2 mutation causes zona pellucida defects and female infertility in mouse and human","volume":"26","author":"Liu","year":"2023","journal-title":"IScience"},{"key":"10.1016\/j.mrrev.2025.108560_bib88","doi-asserted-by":"crossref","first-page":"1205","DOI":"10.1007\/s10815-022-02466-4","article-title":"Novel mutations in ZP2 and ZP3 cause female infertility in three patients","volume":"39","author":"Jia","year":"2022","journal-title":"J. Assist Reprod. Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib89","doi-asserted-by":"crossref","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":"Genet Med"},{"key":"10.1016\/j.mrrev.2025.108560_bib90","doi-asserted-by":"crossref","DOI":"10.3389\/fgene.2023.1256549","article-title":"Case report: a novel homozygous variant in ZP3 is associated with human empty follicle syndrome","volume":"14","author":"Kong","year":"2023","journal-title":"Front Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib91","doi-asserted-by":"crossref","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":"Am. J. Hum. Genet."},{"key":"10.1016\/j.mrrev.2025.108560_bib92","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1002\/humu.24297","article-title":"A novel gene mutation in ZP3 loop region identified in patients with empty follicle syndrome","volume":"43","author":"Zhang","year":"2022","journal-title":"Hum. Mutat."},{"key":"10.1016\/j.mrrev.2025.108560_bib93","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1007\/s10815-020-01995-0","article-title":"A novel mutation in ZP3 causes empty follicle syndrome and abnormal zona pellucida formation","volume":"38","author":"Zhang","year":"2021","journal-title":"J. Assist Reprod. Genet"},{"key":"10.1016\/j.mrrev.2025.108560_bib94","article-title":"Case report: a novel heterozygous ZP3 deletion associated with empty follicle syndrome and abnormal follicular development","volume":"12","author":"Chen","year":"2021","journal-title":"Front Genet"}],"container-title":["Mutation Research - Reviews in Mutation Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1383574225000316?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1383574225000316?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,3,7]],"date-time":"2026-03-07T12:28:32Z","timestamp":1772886512000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S1383574225000316"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,7]]},"references-count":94,"alternative-id":["S1383574225000316"],"URL":"https:\/\/doi.org\/10.1016\/j.mrrev.2025.108560","relation":{},"ISSN":["1383-5742"],"issn-type":[{"value":"1383-5742","type":"print"}],"subject":[],"published":{"date-parts":[[2025,7]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Genetic variations in zona pellucida glycoproteins: Implications for fertility and ART outcomes","name":"articletitle","label":"Article Title"},{"value":"Mutation Research - Reviews in Mutation Research","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.mrrev.2025.108560","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2025 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":"108560"}}