{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T09:28:40Z","timestamp":1785403720512,"version":"3.56.0"},"reference-count":114,"publisher":"MDPI AG","issue":"3","license":[{"start":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:00:00Z","timestamp":1782864000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000057","name":"National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["R35GM142524"],"award-info":[{"award-number":["R35GM142524"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000057","name":"National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["R15GM147865"],"award-info":[{"award-number":["R15GM147865"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]},{"award":["R35GM142524"],"award-info":[{"award-number":["R35GM142524"]}],"id":[{"id":"https:\/\/ror.org\/04q48ey07","id-type":"ROR","asserted-by":"publisher"}]},{"award":["R15GM147865"],"award-info":[{"award-number":["R15GM147865"]}],"id":[{"id":"https:\/\/ror.org\/04q48ey07","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000005","name":"Department of Defense","doi-asserted-by":"publisher","award":["W911NF1810465"],"award-info":[{"award-number":["W911NF1810465"]}],"id":[{"id":"10.13039\/100000005","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["JDB"],"abstract":"<jats:p>Regulation of the cell cycle is critical for maintaining genomic integrity. Therefore, cells have adapted several mechanisms to ensure that cell cycle events occur in a precise order. Some mechanisms regulate cell cycle progression by inhibiting cell cycle drivers, cyclin-dependent kinases (CDKs). The Wee1\/Myt1 family of kinases regulate the G2-to-M phase transition by phosphorylating and inactivating Cdk1. Investigations of Wee1\/Myt1 have mainly focused on its regulation of mitosis; the role of Wee1\/Myt1 kinases in the meiotic cell cycle is less well understood. However, misregulation of Wee1\/Myt1 during meiosis can have a range of fertility consequences from mild to severe, including human fertilization failure and infertility. Studies from several organisms reveal that the meiotic functions of Wee1\/Myt1 kinases differ from mitosis depending on the species and sex. Here, we review how Wee1\/Myt1 kinases regulate cell cycle progression in meiosis across species. We highlight current knowledge of Wee1\/Myt1 in meiosis and discuss unanswered questions and new directions to advance the field of meiosis and reproduction. Understanding the molecular and cellular functions of Wee1\/Myt1 homologs in these various systems may contribute to the discovery of the mechanisms underlying human infertility cases, better diagnoses, and clinical treatments.<\/jats:p>","DOI":"10.3390\/jdb14030029","type":"journal-article","created":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T10:41:01Z","timestamp":1782902461000},"page":"29","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Wee Kinases, Big Impact: Wee and Myt Kinases as Critical Regulators of Meiotic Progression"],"prefix":"10.3390","volume":"14","author":[{"ORCID":"https:\/\/orcid.org\/0009-0009-4662-7144","authenticated-orcid":false,"given":"Shannon","family":"Pfeiffer","sequence":"first","affiliation":[{"name":"Department of Biological Sciences, University of Delaware, Newark, DE 19716, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lourds M.","family":"Fernando","sequence":"additional","affiliation":[{"name":"Department of Developmental Neurobiology, St. Jude\u2019s Children\u2019s Research Hospital, Memphis, TN 38105, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8952-7233","authenticated-orcid":false,"given":"Anna K.","family":"Allen","sequence":"additional","affiliation":[{"name":"Department of Biology, Howard University, Washington, DC 20059, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7697-8580","authenticated-orcid":false,"given":"Aimee","family":"Jaramillo-Lambert","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, University of Delaware, Newark, DE 19716, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,7,1]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"746","DOI":"10.1128\/jb.146.2.746-754.1981","article-title":"Isolation of Cell Size Mutants of a Fission Yeast by a New Selective Method: Characterization of Mutants and Implications for Division Control Mechanisms","volume":"146","author":"Fantes","year":"1981","journal-title":"J. Bacteriol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1038\/342039a0","article-title":"Tyrosine Phosphorylation of the Fission Yeast Cdc2+ Protein Kinase Regulates Entry into Mitosis","volume":"342","author":"Gould","year":"1989","journal-title":"Nature"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"877","DOI":"10.1016\/0955-0674(94)90060-4","article-title":"Cdc2 Regulatory Factors","volume":"6","author":"Coleman","year":"1994","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"5527","DOI":"10.1073\/pnas.082618399","article-title":"Cdk-7 Is Required for mRNA Transcription and Cell Cycle Progression in Caenorhabditis elegans Embryos","volume":"99","author":"Wallenfang","year":"2002","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"872","DOI":"10.1016\/j.tips.2016.06.006","article-title":"Targeting WEE1 Kinase in Cancer","volume":"37","author":"Matheson","year":"2016","journal-title":"Trends Pharmacol. Sci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1016\/j.tips.2025.09.005","article-title":"WEE-Family Kinases in Cancer: Synthetic Lethal Interactions and Drug Discovery","volume":"47","author":"Wang","year":"2025","journal-title":"Trends Pharmacol. Sci."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Zhang, Z., Harish, R., Elahi, N., Saini, S., Telia, A., Kundlas, M., Koroleva, A., Umoh, I.N., Lota, M., and Bilkhu, M. (2025). Targeting WEE1 Kinase for Breast Cancer Therapeutics: An Update. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26125701"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"17997","DOI":"10.1021\/acs.jmedchem.4c01619","article-title":"Targeting Protein Kinase, Membrane-Associated Tyrosine\/Threonine 1 (PKMYT1) for Precision Cancer Therapy: From Discovery to Clinical Trial","volume":"67","author":"Yang","year":"2024","journal-title":"J. Med. Chem."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"428","DOI":"10.1038\/279428a0","article-title":"Epistatic Gene Interactions in the Control of Division in Fission Yeast","volume":"279","author":"Fantes","year":"1979","journal-title":"Nature"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1016\/0014-4827(78)90286-0","article-title":"Control of the Timing of Cell Division in Fission Yeast. Cell Size Mutants Reveal a Second Control Pathway","volume":"115","author":"Fantes","year":"1978","journal-title":"Exp. Cell Res."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"365","DOI":"10.1016\/0014-4827(77)90358-5","article-title":"Controls over the Timing of DNA Replication during the Cell Cycle of Fission Yeast","volume":"107","author":"Nurse","year":"1977","journal-title":"Exp. Cell Res."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"558","DOI":"10.1038\/292558a0","article-title":"Gene Required in G1 for Commitment to Cell Cycle and in G2 for Control of Mitosis in Fission Yeast","volume":"292","author":"Nurse","year":"1981","journal-title":"Nature"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"559","DOI":"10.1016\/0092-8674(87)90458-2","article-title":"Negative Regulation of Mitosis by Wee1+, a Gene Encoding a Protein Kinase Homolog","volume":"49","author":"Russell","year":"1987","journal-title":"Cell"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"3417","DOI":"10.1002\/j.1460-2075.1993.tb06016.x","article-title":"Properties of Saccharomyces Cerevisiae Wee1 and Its Differential Regulation of p34CDC28 in Response to G1 and G2 Cyclins","volume":"12","author":"Booher","year":"1993","journal-title":"EMBO J."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1111","DOI":"10.1016\/0092-8674(91)90266-2","article-title":"Mik1 and Wee1 Cooperate in the Inhibitory Tyrosine Phosphorylation of Cdc2","volume":"64","author":"Lundgren","year":"1991","journal-title":"Cell"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"27847","DOI":"10.1074\/jbc.271.44.27847","article-title":"Sequential Dephosphorylation of P34Cdc2 on Thr-14 and Tyr-15 at the Prophase\/Metaphase Transition","volume":"271","author":"Borgne","year":"1996","journal-title":"J. Biol. Chem."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"2760","DOI":"10.1002\/j.1460-2075.1995.tb07276.x","article-title":"A Pre-Start Checkpoint Preventing Mitosis in Fission Yeast Acts Independently of P34cdc2 Tyrosine Phosphorylation","volume":"14","author":"Hayles","year":"1995","journal-title":"EMBO J."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1016\/0092-8674(92)90177-E","article-title":"The Cdc25 M-Phase Inducer: An Unconventional Protein Phosphatase","volume":"68","author":"Millar","year":"1992","journal-title":"Cell"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1088","DOI":"10.1139\/o92-154","article-title":"Regulation of P105wee1 and P34cdc2 during Meiosis in Schizosaccharomyces pombe","volume":"70","author":"Szankasi","year":"1992","journal-title":"Biochem. Cell Biol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"2853","DOI":"10.4161\/15384101.2014.946807","article-title":"Regulation of Wee1+ Expression during Meiosis in Fission Yeast","volume":"13","author":"Ohtsuka","year":"2014","journal-title":"Cell Cycle"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1091\/mbc.e10-07-0644","article-title":"Spatiotemporal Regulations of Wee1 at the G2\/M Transition","volume":"22","author":"Masuda","year":"2011","journal-title":"Mol. Biol. Cell"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"805","DOI":"10.1016\/S1097-2765(00)80390-1","article-title":"The Pachytene Checkpoint in S. Cerevisiae Depends on Swe1-Mediated Phosphorylation of the Cyclin-Dependent Kinase Cdc28","volume":"4","author":"Leu","year":"1999","journal-title":"Mol. Cell"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"4124","DOI":"10.1073\/pnas.0400641101","article-title":"Coupling Morphogenesis to Mitotic Entry","volume":"101","author":"Sakchaisri","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1007\/BF00326233","article-title":"Common Genes and Pathways in the Regulation of the Mitotic and Meiotic Cell Cycles of Schizosaccharomyces pombe","volume":"20","author":"Grallert","year":"1991","journal-title":"Curr. Genet."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"2581","DOI":"10.1101\/gad.13.19.2581","article-title":"Meiotic DNA Replication Checkpoint Control in Fission Yeast","volume":"13","author":"Murakami","year":"1999","journal-title":"Genes Dev."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1042\/bj3490001","article-title":"DNA Replication and Damage Checkpoints and Meiotic Cell Cycle Controls in the Fission and Budding Yeasts","volume":"349","author":"Murakami","year":"2000","journal-title":"Biochem. J."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"2600","DOI":"10.1101\/gad.11.20.2600","article-title":"Meiotic Chromosomes: It Takes Two to Tango","volume":"11","author":"Roeder","year":"1997","journal-title":"Genes Dev."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"2051","DOI":"10.1091\/mbc.E16-02-0104","article-title":"Myt1 Inhibition of Cyclin A\/Cdk1 Is Essential for Fusome Integrity and Premeiotic Centriole Engagement in Drosophila Spermatocytes","volume":"27","author":"Varadarajan","year":"2016","journal-title":"Mol. Biol. Cell"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"2329","DOI":"10.1534\/g3.114.013649","article-title":"An RNAi-Based Suppressor Screen Identifies Interactors of the Myt1 Ortholog of Caenorhabditis elegans","volume":"4","author":"Allen","year":"2014","journal-title":"G3 Genes Genomes Genet."},{"key":"ref_30","first-page":"10","article-title":"Comparison of N- and C-Terminally Endogenously GFP-Tagged WEE-1.3 Strains in C. elegans","volume":"2021","author":"Fernando","year":"2021","journal-title":"microPubl. Biol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"evab060","DOI":"10.1093\/gbe\/evab060","article-title":"Divergent Gene Expression Following Duplication of Meiotic Genes in the Stick Insect Clitarchus Hookeri","volume":"13","author":"Wu","year":"2021","journal-title":"Genome Biol. Evol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"3597","DOI":"10.1083\/jcb.201812122","article-title":"SGK Phosphorylates Cdc25 and Myt1 to Trigger Cyclin B\u2013Cdk1 Activation at the Meiotic G2\/M Transition","volume":"218","author":"Hiraoka","year":"2019","journal-title":"J. Cell Biol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1126\/science.270.5233.86","article-title":"Myt1: A Membrane-Associated Inhibitory Kinase That Phosphorylates Cdc2 on Both Threonine-14 and Tyrosine-15","volume":"270","author":"Mueller","year":"1995","journal-title":"Science"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"5037","DOI":"10.1093\/emboj\/17.17.5037","article-title":"A Link between MAP Kinase and P34Cdc2\/Cyclin B during Oocyte Maturation: P90Rsk Phosphorylates and Inactivates the P34Cdc2 Inhibitory Kinase Myt1","volume":"17","author":"Palmer","year":"1998","journal-title":"EMBO J."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1002\/mrd.20952","article-title":"Activation of Both Mos and Cdc25 Is Required for G2-M Transition in Perch Oocyte","volume":"76","author":"Priyadarshini","year":"2009","journal-title":"Mol. Reprod. Dev."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1083\/jcb.200907161","article-title":"Wee1B, Myt1, and Cdc25 Function in Distinct Compartments of the Mouse Oocyte to Control Meiotic Resumption","volume":"188","author":"Oh","year":"2010","journal-title":"J. Cell Biol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"107858","DOI":"10.1016\/j.celrep.2020.107858","article-title":"Cyclin N-Terminal Domain-Containing-1 Coordinates Meiotic Crossover Formation with Cell-Cycle Progression in a Cyclin-Independent Manner","volume":"32","author":"Gray","year":"2020","journal-title":"Cell Rep."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"2375","DOI":"10.4161\/cc.8.15.9073","article-title":"Critical Effect of pigWee1B on the Regulation of Meiotic Resumption in Porcine Immature Oocytes","volume":"8","author":"Shimaoka","year":"2009","journal-title":"Cell Cycle"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1093\/humrep\/dei285","article-title":"Decreased mRNA Transcripts of M-Phase Promoting Factor and Its Regulators in the Testes of Infertile Men","volume":"21","author":"Lin","year":"2006","journal-title":"Hum. Reprod."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"649","DOI":"10.1016\/j.ajhg.2018.02.015","article-title":"Homozygous Mutations in WEE2 Cause Fertilization Failure and Female Infertility","volume":"102","author":"Sang","year":"2018","journal-title":"Am. J. Hum. Genet."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/S0167-4781(99)00027-5","article-title":"A Caenorhabditis elegans Wee1 Homolog Is Expressed in a Temporally and Spatially Restricted Pattern during Embryonic Development","volume":"1445","author":"Wilson","year":"1999","journal-title":"Biochim. Biophys. Acta BBA\u2014Gene Struct. Expr."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1186\/1747-1028-1-6","article-title":"Cyclin-Dependent Kinases in C. elegans","volume":"1","author":"Boxem","year":"2006","journal-title":"Cell Div."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"5009","DOI":"10.1242\/dev.129.21.5009","article-title":"Dominant Mutations in the Caenorhabditis elegans Myt1 Orthologwee-1.3 Reveal a Novel Domain That Controls M-Phase Entry during Spermatogenesis","volume":"129","author":"Lamitina","year":"2002","journal-title":"Development"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"3411","DOI":"10.1038\/s44319-025-00485-7","article-title":"Dynamic SAS-6 Phosphorylation Aids Centrosome Duplication and Elimination in C. elegans Oogenesis","volume":"26","author":"Qi","year":"2025","journal-title":"EMBO Rep."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"381","DOI":"10.1146\/annurev-cellbio-100616-060553","article-title":"Assembly and Positioning of the Oocyte Meiotic Spindle","volume":"34","author":"Mogessie","year":"2018","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"research0070.1","DOI":"10.1186\/gb-2002-3-12-research0070","article-title":"Whole-Genome Analysis of Animal A- and B-Type Cyclins","volume":"3","author":"Nieduszynski","year":"2002","journal-title":"Genome Biol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"4091","DOI":"10.4161\/cc.8.24.10171","article-title":"Elegans Mitotic Cyclins Have Distinct as Well as Overlapping Functions in Chromosome Segregation","volume":"8","author":"Lorson","year":"2009","journal-title":"Cell Cycle"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"2722","DOI":"10.1242\/jcs.204578","article-title":"The B-Type Cyclin CYB-1 Maintains the Proper Position and Number of Centrosomes during Spermatogenesis in Caenorhabditis elegans","volume":"130","author":"Yoon","year":"2017","journal-title":"J. Cell Sci."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"993","DOI":"10.1242\/jcs.060442","article-title":"Cdc-25.2, a C. elegans Ortholog of Cdc25, Is Required to Promote Oocyte Maturation","volume":"123","author":"Kim","year":"2010","journal-title":"J. Cell Sci."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"2144","DOI":"10.1126\/science.1057586","article-title":"A Sperm Cytoskeletal Protein That Signals Oocyte Meiotic Maturation and Ovulation","volume":"291","author":"Miller","year":"2001","journal-title":"Science"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"697","DOI":"10.1242\/dev.02241","article-title":"The C. elegans Myt1 Ortholog Is Required for the Proper Timing of Oocyte Maturation","volume":"133","author":"Burrows","year":"2006","journal-title":"Development"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1002\/dvdy.235","article-title":"The Caenorhabditis elegans Proteasome Subunit RPN-12 Is Required for Hermaphrodite Germline Sex Determination and Oocyte Quality","volume":"250","author":"Fernando","year":"2021","journal-title":"Dev. Dyn."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"717","DOI":"10.1016\/j.cub.2010.02.050","article-title":"A Two-Step Inactivation Mechanism of Myt1 Ensures CDK1\/Cyclin B Activation and Meiosis I Entry","volume":"20","author":"Ruiz","year":"2010","journal-title":"Curr. Biol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1534\/genetics.113.156281","article-title":"Dual Phosphorylation of Cdk1 Coordinates Cell Proliferation with Key Developmental Processes in Drosophila","volume":"196","author":"Ayeni","year":"2014","journal-title":"Genetics"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"4075","DOI":"10.1242\/dev.01965","article-title":"Drosophila Myt1 Is a Cdk1 Inhibitory Kinase That Regulates Multiple Aspects of Cell Cycle Behavior during Gametogenesis","volume":"132","author":"Jin","year":"2005","journal-title":"Development"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"666","DOI":"10.1016\/S0955-0674(00)00150-2","article-title":"Regulation of the Meiotic Cell Cycle in Oocytes","volume":"12","author":"Nebreda","year":"2000","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1333","DOI":"10.1091\/mbc.6.10.1333","article-title":"Drosophila Wee1 Kinase Rescues Fission Yeast from Mitotic Catastrophe and Phosphorylates Drosophila Cdc2 in Vitro","volume":"6","author":"Campbell","year":"1995","journal-title":"Mol. Biol. Cell"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"467","DOI":"10.1016\/S0898-6568(01)00276-5","article-title":"Identification of Drosophila Myt1 Kinase and Its Role in Golgi during Mitosis","volume":"14","author":"Cornwell","year":"2002","journal-title":"Cell. Signal."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"233","DOI":"10.1007\/978-1-4615-5371-7_18","article-title":"Myt1: A Wee1-Type Kinase That Phosphorylates Cdc2 on Residue Thr14","volume":"3","author":"Fattaey","year":"1997","journal-title":"Prog. Cell Cycle Res."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1093\/genetics\/155.1.159","article-title":"Drosophila Wee1 Has an Essential Role in the Nuclear Divisions of Early Embryogenesis","volume":"155","author":"Price","year":"2000","journal-title":"Genetics"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"721","DOI":"10.1093\/genetics\/161.2.721","article-title":"Ectopic Expression of the Drosophila Cdk1 Inhibitory Kinases, Wee1 and Myt1, Interferes with the Second Mitotic Wave and Disrupts Pattern Formation during Eye Development","volume":"161","author":"Price","year":"2002","journal-title":"Genetics"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"a005850","DOI":"10.1101\/cshperspect.a005850","article-title":"Germ Cell Intercellular Bridges","volume":"3","author":"Greenbaum","year":"2011","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1006\/scdb.1998.0227","article-title":"Genetic Control of Cell Proliferation and Differentiation in Drosophila Spermatogenesis","volume":"9","author":"Fuller","year":"1998","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1016\/j.ymeth.2014.04.020","article-title":"Investigating Spermatogenesis in Drosophila melanogaster","volume":"68","author":"Demarco","year":"2014","journal-title":"Methods"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1016\/S0074-7696(08)62542-5","article-title":"The Meiotic Behavior of the Drosophila Oocyte","volume":"28","author":"King","year":"1970","journal-title":"Int. Rev. Cytol."},{"key":"ref_66","first-page":"a005553","article-title":"Developmental Control of Oocyte Maturation and Egg Activation in Metazoan Models","volume":"3","year":"2011","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"649","DOI":"10.1016\/0092-8674(93)90393-5","article-title":"Germline Cysts: Communes That Work","volume":"72","author":"Spradling","year":"1993","journal-title":"Cell"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"R787","DOI":"10.1016\/j.cub.2004.09.020","article-title":"APC\/C and SCF: Controlling Each Other and the Cell Cycle","volume":"14","author":"Vodermaier","year":"2004","journal-title":"Curr. Biol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1091\/mbc.6.1.119","article-title":"Cell Cycle Regulation of a Xenopus Wee1-like Kinase","volume":"6","author":"Mueller","year":"1995","journal-title":"Mol. Biol. Cell"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"2472","DOI":"10.1093\/emboj\/21.10.2472","article-title":"The Existence of Two Distinct Wee1 Isoforms in Xenopus: Implications for the Developmental Regulation of the Cell Cycle","volume":"21","author":"Okamoto","year":"2002","journal-title":"EMBO J."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"3735","DOI":"10.1242\/dev.063974","article-title":"A Critical Balance between Cyclin B Synthesis and Myt1 Activity Controls Meiosis Entry in Xenopus Oocytes","volume":"138","author":"Martoriati","year":"2011","journal-title":"Development"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"328","DOI":"10.1101\/gad.14.3.328","article-title":"Absence of Wee1 Ensures the Meiotic Cell Cycle in Xenopus Oocytes","volume":"14","author":"Nakajo","year":"2000","journal-title":"Genes Dev."},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Tsai, T.Y.-C., Theriot, J.A., and Ferrell, J.E. (2014). Changes in Oscillatory Dynamics in the Cell Cycle of Early Xenopus Laevis Embryos. PLoS Biol., 12.","DOI":"10.1371\/journal.pbio.1001788"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"203754","DOI":"10.1016\/j.cdev.2021.203754","article-title":"Regulation of Myt1 Kinase Activity via Its N-Terminal Region in Xenopus Meiosis and Mitosis","volume":"169","author":"Aiba","year":"2022","journal-title":"Cells Dev."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"571","DOI":"10.1128\/MCB.17.2.571","article-title":"The Human Myt1 Kinase Preferentially Phosphorylates Cdc2 on Threonine 14 and Localizes to the Endoplasmic Reticulum and Golgi Complex","volume":"17","author":"Liu","year":"1997","journal-title":"Mol. Cell. Biol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1242\/dev.125.2.237","article-title":"Analysis of the Early Embryonic Cell Cycles of Xenopus; Regulation of Cell Cycle Length by Xe-Wee1 and Mos","volume":"125","author":"Murakami","year":"1998","journal-title":"Development"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1002\/jez.1401770202","article-title":"Cytoplasmic Control of Nuclear Behavior during Meiotic Maturation of Frog Oocytes","volume":"177","author":"Masui","year":"1971","journal-title":"J. Exp. Zool."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"2129","DOI":"10.1242\/dev.129.9.2129","article-title":"A New Role for Mos in Xenopus Oocyte Maturation: Targeting Myt1 Independently of MAPK","volume":"129","author":"Peter","year":"2002","journal-title":"Development"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1016\/0962-8924(96)81034-8","article-title":"Meiotic Metaphase Arrest in Animal Oocytes: Its Mechanisms and Biological Significance","volume":"6","author":"Sagata","year":"1996","journal-title":"Trends Cell Biol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"3279","DOI":"10.1091\/mbc.10.10.3279","article-title":"Two Distinct Mechanisms Control the Accumulation of Cyclin B1 and Mos in Xenopus Oocytes in Response to Progesterone","volume":"10","author":"Jessus","year":"1999","journal-title":"Mol. Biol. Cell"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1083\/jcb.114.4.755","article-title":"On the Synthesis and Destruction of A- and B-Type Cyclins during Oogenesis and Meiotic Maturation in Xenopus Laevis","volume":"114","author":"Kobayashi","year":"1991","journal-title":"J. Cell Biol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"1057","DOI":"10.1038\/sj.emboj.7600567","article-title":"The Polo-like Kinase Plx1 Interacts with and Inhibits Myt1 after Fertilization of Xenopus Eggs","volume":"24","author":"Inoue","year":"2005","journal-title":"EMBO J."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"421","DOI":"10.4161\/cc.5.4.2492","article-title":"Autophosphorylation of Ser66 on Xenopus Myt1 Is a Prerequisite for Meiotic Inactivation of Myt1","volume":"5","author":"Safi","year":"2006","journal-title":"Cell Cycle"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"1190","DOI":"10.1095\/biolreprod.109.081984","article-title":"WEE2 Is an Oocyte-Specific Meiosis Inhibitor in Rhesus Macaque Monkeys","volume":"82","author":"Hanna","year":"2010","journal-title":"Biol. Reprod."},{"key":"ref_85","doi-asserted-by":"crossref","unstructured":"Latorraca, L.B., Galv\u00e3o, A., Rabaglino, M.B., D\u2019Augero, J.M., Kelsey, G., and Fair, T. (2024). Single-Cell Profiling Reveals Transcriptome Dynamics during Bovine Oocyte Growth. BMC Genom., 25.","DOI":"10.1186\/s12864-024-10234-0"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"4515","DOI":"10.1091\/mbc.e12-04-0260","article-title":"Human Cdc14A Regulates Wee1 Stability by Counteracting CDK-Mediated Phosphorylation","volume":"23","author":"Ovejero","year":"2012","journal-title":"Mol. Biol. Cell"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"10431","DOI":"10.1073\/pnas.1506974112","article-title":"The Genetics of Human Infertility by Functional Interrogation of SNPs in Mice","volume":"112","author":"Singh","year":"2015","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"1670","DOI":"10.1016\/j.cub.2005.07.056","article-title":"Wee1B Is an Oocyte-Specific Kinase Involved in the Control of Meiotic Arrest in the Mouse","volume":"15","author":"Han","year":"2005","journal-title":"Curr. Biol."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1262\/jrd.10-122H","article-title":"Analyses of the Regulatory Mechanism of Porcine WEE1B: The Phosphorylation Sites of Porcine WEE1B and Mouse WEE1B Are Different","volume":"57","author":"Shimaoka","year":"2011","journal-title":"J. Reprod. Dev."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1530\/rep.1.00192","article-title":"The Dynamics of Cyclin B1 Distribution during Meiosis I in Mouse Oocytes","volume":"128","author":"Marangos","year":"2004","journal-title":"Reproduction"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"501","DOI":"10.1093\/biolre\/iox168","article-title":"Effects of Exportin 1 on Nuclear Transport and Meiotic Resumption in Porcine Full-Grown and Growing Oocytes","volume":"98","author":"Onuma","year":"2018","journal-title":"Biol. Reprod."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"1929","DOI":"10.3892\/mmr.2013.1437","article-title":"Ser 15 of WEE1B Is a Potential PKA Phosphorylation Target in G2\/M Transition in One-Cell Stage Mouse Embryos","volume":"7","author":"Liu","year":"2013","journal-title":"Mol. Med. Rep."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"462","DOI":"10.1126\/science.1199211","article-title":"Protein Tyrosine Kinase Wee1B Is Essential for Metaphase II Exit in Mouse Oocytes","volume":"332","author":"Oh","year":"2011","journal-title":"Science"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1016\/j.fertnstert.2018.11.018","article-title":"Novel WEE2 Gene Variants Identified in Patients with Fertilization Failure and Female Infertility","volume":"111","author":"Zhao","year":"2019","journal-title":"Fertil. Steril."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"1096","DOI":"10.1080\/09513590.2021.1916458","article-title":"Clinical Exome Sequencing Identifies Novel Compound Heterozygous Mutations of the WEE2 Gene in Primary Infertile Women with Fertilization Failure","volume":"37","author":"Wang","year":"2021","journal-title":"Gynecol. Endocrinol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"2861","DOI":"10.1007\/s10815-021-02285-z","article-title":"Novel WEE2 Compound Heterozygous Mutations Identified in Patients with Fertilization Failure or Poor Fertilization","volume":"38","author":"Jin","year":"2021","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"700","DOI":"10.1111\/cge.14429","article-title":"Novel WEE2 Homozygous Mutations c.1346C>T and c.949A>T Identified in Primary Infertile Women Due to Unexplained Fertilization Failure","volume":"104","author":"Guo","year":"2023","journal-title":"Clin. Genet."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"3337","DOI":"10.1007\/s10815-024-03260-0","article-title":"Novel Splicing Mutations in PATL2 and WEE2 Cause Oocyte Degradation and Fertilization Failure","volume":"41","author":"Liu","year":"2024","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"3045","DOI":"10.2147\/IJWH.S545145","article-title":"Association of WEE2 Gene Polymorphism with Fertilization Failure in Women Undergoing Intracytoplasmic Sperm Injection","volume":"17","author":"Abdulhussein","year":"2025","journal-title":"Int. J. Women\u2019s Health"},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"3817","DOI":"10.1007\/s10815-025-03626-y","article-title":"Novel Homozygous Variants in ASTL and WEE2 Responsible for Female Infertility Characterized by Abnormal Fertilization","volume":"42","author":"Xie","year":"2025","journal-title":"J. Assist. Reprod. Genet."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"13363","DOI":"10.1002\/slct.201903696","article-title":"Identification and Screening of Selective WEE2 Inhibitors to Develop Non-Hormonal Contraceptives That Specifically Target Meiosis","volume":"4","author":"Hanna","year":"2019","journal-title":"ChemistrySelect"},{"key":"ref_102","doi-asserted-by":"crossref","unstructured":"Ali, S., Elasbali, A.M., Alzahrani, W., Mohammad, T., Hassan, M.I., and Zhou, T. (2026). Identification of FDA-Approved Drugs as Potential Inhibitors of WEE2: Structure-Based Virtual Screening and Molecular Dynamics with Perspectives for Machine Learning-Assisted Prioritization. Life, 16.","DOI":"10.3390\/life16020185"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"112178","DOI":"10.1016\/j.yexcr.2020.112178","article-title":"Cdc14a Has a Role in Spermatogenesis, Sperm Maturation and Male Fertility","volume":"395","author":"Wen","year":"2020","journal-title":"Exp. Cell Res."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1038\/ncb741","article-title":"Akt Inhibits Myt1 in the Signalling Pathway That Leads to Meiotic G2\/M-Phase Transition","volume":"4","author":"Okumura","year":"2002","journal-title":"Nat. Cell Biol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1093\/jhered\/est011","article-title":"Inventory and Phylogenetic Analysis of Meiotic Genes in Monogonont Rotifers","volume":"104","author":"Hanson","year":"2013","journal-title":"J. Hered."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"MSB201175","DOI":"10.1038\/msb.2011.75","article-title":"Fast, Scalable Generation of High-quality Protein Multiple Sequence Alignments Using Clustal Omega","volume":"7","author":"Sievers","year":"2011","journal-title":"Mol. Syst. Biol."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"1189","DOI":"10.1093\/bioinformatics\/btp033","article-title":"Jalview Version 2\u2014A Multiple Sequence Alignment Editor and Analysis Workbench","volume":"25","author":"Waterhouse","year":"2009","journal-title":"Bioinformatics"},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"105806","DOI":"10.1016\/j.isci.2022.105806","article-title":"Temporal Phosphoproteomics Reveals WEE1-Dependent Control of 53BP1 Pathway","volume":"26","author":"Petrosius","year":"2023","journal-title":"iScience"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"941","DOI":"10.1038\/ncb3576","article-title":"Budding Yeast Wee1 Distinguishes Spindle Pole Bodies to Guide Their Pattern of Age-Dependent Segregation","volume":"19","author":"Lengefeld","year":"2017","journal-title":"Nat. Cell Biol."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"4127","DOI":"10.1534\/g3.116.036129","article-title":"Cdc-25.4, a Caenorhabditis elegans Ortholog of Cdc25, Is Required for Male Mating Behavior","volume":"6","author":"Oh","year":"2016","journal-title":"G3 Genes Genomes Genet."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"2131","DOI":"10.1002\/1873-3468.12717","article-title":"Depletion of Cdc-25.3, a Caenorhabditis elegans Orthologue of Cdc25, Increases Physiological Germline Apoptosis","volume":"591","author":"Sung","year":"2017","journal-title":"FEBS Lett."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"1354","DOI":"10.4161\/cc.19755","article-title":"CDC-25.1 Controls the Rate of Germline Mitotic Cell Cycle by Counteracting WEE-1.3 and by Positively Regulating CDK-1 in Caenorhabditis elegans","volume":"11","author":"Yoon","year":"2012","journal-title":"Cell Cycle"},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"e2503192","DOI":"10.1002\/advs.202503192","article-title":"Macrophage WEE1 Directly Binds to and Phosphorylates NF-\u03baB P65 Subunit to Induce Inflammatory Response and Drive Atherosclerosis","volume":"12","author":"Huang","year":"2025","journal-title":"Adv. Sci."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"1702","DOI":"10.1093\/eurheartj\/ehaf349","article-title":"Checkpoint Kinase Wee1 Activation Drives Inflammation and Hypertrophy through the Protein Kinase B\/Phosphoinositide 3-Kinases-Nuclear Factor \u03baB Pathway in Cardiomyocytes","volume":"47","author":"Wang","year":"2025","journal-title":"Eur. Heart J."}],"container-title":["Journal of Developmental Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2221-3759\/14\/3\/29\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T12:24:09Z","timestamp":1782908649000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2221-3759\/14\/3\/29"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,1]]},"references-count":114,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["jdb14030029"],"URL":"https:\/\/doi.org\/10.3390\/jdb14030029","relation":{},"ISSN":["2221-3759"],"issn-type":[{"value":"2221-3759","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,1]]}}}