{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,23]],"date-time":"2026-06-23T02:13:26Z","timestamp":1782180806966,"version":"3.54.5"},"reference-count":74,"publisher":"Springer Science and Business Media LLC","issue":"2","license":[{"start":{"date-parts":[[2025,1,16]],"date-time":"2025-01-16T00:00:00Z","timestamp":1736985600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2025,1,16]],"date-time":"2025-01-16T00:00:00Z","timestamp":1736985600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat Struct Mol Biol"],"published-print":{"date-parts":[[2025,2]]},"DOI":"10.1038\/s41594-024-01471-8","type":"journal-article","created":{"date-parts":[[2025,1,16]],"date-time":"2025-01-16T10:02:13Z","timestamp":1737021733000},"page":"381-392","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":10,"title":["Maternal ELL3 loss-of-function leads to oocyte aneuploidy and early miscarriage"],"prefix":"10.1038","volume":"32","author":[{"given":"Shiqi","family":"Zhu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peng","family":"Xie","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yi","family":"Yang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yan","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chuanxin","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yu","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuhan","family":"Si","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jin","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jingjing","family":"He","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hao","family":"Si","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ke","family":"Fang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Binbin","family":"Ma","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xu","family":"Jiang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lindi","family":"Huang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiamin","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tian","family":"Min","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Beihong","family":"Zheng","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lincui","family":"Da","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dianliang","family":"Lin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kun","family":"Gao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuanyuan","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mingtao","family":"Huang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fengchang","family":"Qiao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Haiqin","family":"Huo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Haoyang","family":"Feng","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9515-7534","authenticated-orcid":false,"given":"Han","family":"Zhao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6637-6631","authenticated-orcid":false,"given":"Zijiang","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhengfeng","family":"Xu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jing","family":"Xie","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hua","family":"Cao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5707-2078","authenticated-orcid":false,"given":"Jin","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8982-5911","authenticated-orcid":false,"given":"Xuebiao","family":"Yao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9179-4787","authenticated-orcid":false,"given":"Wei","family":"Xie","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6909-6018","authenticated-orcid":false,"given":"Yan","family":"Sun","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9552-6096","authenticated-orcid":false,"given":"Keliang","family":"Wu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2456-4559","authenticated-orcid":false,"given":"Bo","family":"Xiong","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7297-8847","authenticated-orcid":false,"given":"Ping","family":"Hu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0715-0648","authenticated-orcid":false,"given":"Zhuojuan","family":"Luo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2121-584X","authenticated-orcid":false,"given":"Chengqi","family":"Lin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,1,16]]},"reference":[{"key":"1471_CR1","doi-asserted-by":"crossref","first-page":"1658","DOI":"10.1016\/S0140-6736(21)00682-6","volume":"397","author":"S Quenby","year":"2021","unstructured":"Quenby, S. et al. Miscarriage matters: the epidemiological, physical, psychological, and economic costs of early pregnancy loss. Lancet 397, 1658\u20131667 (2021).","journal-title":"Lancet"},{"key":"1471_CR2","doi-asserted-by":"crossref","first-page":"3233","DOI":"10.1038\/s41591-023-02645-5","volume":"9","author":"R Essers","year":"2023","unstructured":"Essers, R. et al. Prevalence of chromosomal alterations in first-trimester spontaneous pregnancy loss. Nat. Med. 9, 3233\u20133242 (2023).","journal-title":"Nat. Med."},{"key":"1471_CR3","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1038\/s41580-022-00517-3","volume":"24","author":"C Charalambous","year":"2023","unstructured":"Charalambous, C., Webster, A. & Schuh, M. Aneuploidy in mammalian oocytes and the impact of maternal ageing. Nat. Rev. Mol. Cell Biol. 24, 27\u201344 (2023).","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"1471_CR4","doi-asserted-by":"crossref","first-page":"eabj3944","DOI":"10.1126\/science.abj3944","volume":"375","author":"C So","year":"2022","unstructured":"So, C. et al. Mechanism of spindle pole organization and instability in human oocytes. Science 375, eabj3944 (2022).","journal-title":"Science"},{"key":"1471_CR5","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1042\/BST20200043","volume":"49","author":"C Thomas","year":"2021","unstructured":"Thomas, C., Cavazza, T. & Schuh, M. Aneuploidy in human eggs: contributions of the meiotic spindle. Biochem. Soc. Trans. 49, 107\u2013118 (2021).","journal-title":"Biochem. Soc. Trans."},{"key":"1471_CR6","doi-asserted-by":"crossref","first-page":"1143","DOI":"10.1126\/science.aaa9529","volume":"348","author":"Z Holubcov\u00e1","year":"2015","unstructured":"Holubcov\u00e1, Z., Blayney, M., Elder, K. & Schuh, M. Human oocytes. Error-prone chromosome-mediated spindle assembly favors chromosome segregation defects in human oocytes. Science 348, 1143\u20131147 (2015).","journal-title":"Science"},{"key":"1471_CR7","doi-asserted-by":"crossref","first-page":"a022608","DOI":"10.1101\/cshperspect.a022608","volume":"10","author":"HV Goodson","year":"2018","unstructured":"Goodson, H. V. & Jonasson, E. M. Microtubules and microtubule-associated proteins. Cold Spring Harb. Perspect. Biol. 10, a022608 (2018).","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"1471_CR8","doi-asserted-by":"crossref","first-page":"381","DOI":"10.1146\/annurev-cellbio-100616-060553","volume":"34","author":"B Mogessie","year":"2018","unstructured":"Mogessie, B., Scheffler, K. & Schuh, M. Assembly and positioning of the oocyte meiotic spindle. Annu. Rev. Cell Dev. Biol. 34, 381\u2013403 (2018).","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"1471_CR9","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1016\/j.devcel.2017.04.009","volume":"41","author":"H Kyogoku","year":"2017","unstructured":"Kyogoku, H. & Kitajima, T. S. Large cytoplasm is linked to the error-prone nature of oocytes. Dev. Cell 41, 287\u2013298.e4 (2017).","journal-title":"Dev. Cell"},{"key":"1471_CR10","doi-asserted-by":"crossref","first-page":"158","DOI":"10.1126\/science.adf7760","volume":"380","author":"Q Sang","year":"2023","unstructured":"Sang, Q., Ray, P. F. & Wang, L. Understanding the genetics of human infertility. Science 380, 158\u2013163 (2023).","journal-title":"Science"},{"key":"1471_CR11","doi-asserted-by":"crossref","first-page":"218","DOI":"10.1093\/humupd\/dmac044","volume":"29","author":"A Van Der Kelen","year":"2023","unstructured":"Van Der Kelen, A. et al. A systematic review and evidence assessment of monogenic gene-disease relationships in human female infertility and differences in sex development. Hum. Reprod. Update 29, 218\u2013232 (2023).","journal-title":"Hum. Reprod. Update"},{"key":"1471_CR12","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1056\/NEJMoa1510791","volume":"374","author":"R Feng","year":"2016","unstructured":"Feng, R. et al. Mutations in TUBB8 and human oocyte meiotic arrest. N. Engl. J. Med. 374, 223\u2013232 (2016).","journal-title":"N. Engl. J. Med."},{"key":"1471_CR13","doi-asserted-by":"crossref","first-page":"eabq7361","DOI":"10.1126\/science.abq7361","volume":"378","author":"T Wu","year":"2022","unstructured":"Wu, T. et al. The mechanism of acentrosomal spindle assembly in human oocytes. Science 378, eabq7361 (2022).","journal-title":"Science"},{"key":"1471_CR14","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1093\/humrep\/dew322","volume":"32","author":"B Chen","year":"2017","unstructured":"Chen, B. et al. Novel mutations and structural deletions inTUBB8: expanding mutational and phenotypic spectrum of patients with arrest in oocyte maturation, fertilization or early embryonic development. Hum. Reprod. 32, 457\u2013464 (2017).","journal-title":"Hum. Reprod."},{"key":"1471_CR15","doi-asserted-by":"crossref","first-page":"921","DOI":"10.1007\/s13238-020-00756-0","volume":"11","author":"L Zhao","year":"2020","unstructured":"Zhao, L. et al. Biallelic mutations in CDC20 cause female infertility characterized by abnormalities in oocyte maturation and early embryonic development. Protein Cell 11, 921\u2013927 (2020).","journal-title":"Protein Cell"},{"key":"1471_CR16","doi-asserted-by":"crossref","first-page":"2134","DOI":"10.1093\/humrep\/deaa148","volume":"35","author":"KM Tyc","year":"2020","unstructured":"Tyc, K. M. et al. Exome sequencing links CEP120 mutation to maternally derived aneuploid conception risk. Hum. Reprod. 35, 2134\u20132148 (2020).","journal-title":"Hum. Reprod."},{"key":"1471_CR17","doi-asserted-by":"crossref","first-page":"727","DOI":"10.1038\/ng.3306","volume":"47","author":"CS Ottolini","year":"2015","unstructured":"Ottolini, C. S. et al. Genome-wide maps of recombination and chromosome segregation in human oocytes and embryos show selection for maternal recombination rates. Nat. Genet. 47, 727\u2013735 (2015).","journal-title":"Nat. Genet."},{"key":"1471_CR18","doi-asserted-by":"crossref","first-page":"493","DOI":"10.1038\/nrg3245","volume":"13","author":"SI Nagaoka","year":"2012","unstructured":"Nagaoka, S. I., Hassold, T. J. & Hunt, P. A. Human aneuploidy: mechanisms and new insights into an age-old problem. Nat. Rev. Genet. 13, 493\u2013504 (2012).","journal-title":"Nat. Rev. Genet."},{"key":"1471_CR19","doi-asserted-by":"crossref","first-page":"R895","DOI":"10.1016\/j.cub.2018.06.057","volume":"28","author":"AI Mihajlovi\u0107","year":"2018","unstructured":"Mihajlovi\u0107, A. I. & FitzHarris, G. Segregating chromosomes in the mammalian oocyte. Curr. Biol. 28, R895\u2013R907 (2018).","journal-title":"Curr. Biol."},{"key":"1471_CR20","doi-asserted-by":"crossref","first-page":"1466","DOI":"10.1126\/science.aav7321","volume":"365","author":"JR Gruhn","year":"2019","unstructured":"Gruhn, J. R. et al. Chromosome errors in human eggs shape natural fertility over reproductive life span. Science 365, 1466\u20131469 (2019).","journal-title":"Science"},{"key":"1471_CR21","doi-asserted-by":"crossref","first-page":"32052","DOI":"10.1074\/jbc.M005175200","volume":"275","author":"T Miller","year":"2000","unstructured":"Miller, T., Williams, K., Johnstone, R. W. & Shilatifard, A. Identification, cloning, expression, and biochemical characterization of the testis-specific RNA polymerase II elongation factor ELL3. J. Biol. Chem. 275, 32052\u201332056 (2000).","journal-title":"J. Biol. Chem."},{"key":"1471_CR22","doi-asserted-by":"crossref","first-page":"1873","DOI":"10.1126\/science.271.5257.1873","volume":"271","author":"A Shilatifard","year":"1996","unstructured":"Shilatifard, A., Lane, W. S., Jackson, K. W., Conaway, R. C. & Conaway, J. W. An RNA polymerase II elongation factor encoded by the human ELL gene. Science 271, 1873\u20131876 (1996).","journal-title":"Science"},{"key":"1471_CR23","doi-asserted-by":"crossref","first-page":"429","DOI":"10.1016\/j.molcel.2010.01.026","volume":"37","author":"C Lin","year":"2010","unstructured":"Lin, C. et al. AFF4, a component of the ELL\/P-TEFb elongation complex and a shared subunit of MLL chimeras, can link transcription elongation to leukemia. Mol. Cell 37, 429\u2013437 (2010).","journal-title":"Mol. Cell"},{"key":"1471_CR24","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1016\/j.cell.2012.12.015","volume":"152","author":"C Lin","year":"2013","unstructured":"Lin, C., Garruss, A. S., Luo, Z., Guo, F. & Shilatifard, A. The RNA Pol II elongation factor Ell3 marks enhancers in ES cells and primes future gene activation. Cell 152, 144\u2013156 (2013).","journal-title":"Cell"},{"key":"1471_CR25","doi-asserted-by":"crossref","first-page":"1319","DOI":"10.1038\/s41556-023-01211-y","volume":"25","author":"S Meng","year":"2023","unstructured":"Meng, S. et al. Young LINE-1 transposon 5\u2032 UTRs marked by elongation factor ELL3 function as enhancers to regulate naive pluripotency in embryonic stem cells. Nat. Cell Biol. 25, 1319\u20131331 (2023).","journal-title":"Nat. Cell Biol."},{"key":"1471_CR26","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1016\/j.cub.2017.11.049","volume":"28","author":"A Vallot","year":"2018","unstructured":"Vallot, A. et al. Tension-induced error correction and not kinetochore attachment status activates the SAC in an aurora-B\/C-dependent manner in oocytes. Curr. Biol. 28, 130\u2013139 e3 (2018).","journal-title":"Curr. Biol."},{"key":"1471_CR27","doi-asserted-by":"crossref","first-page":"e1000384","DOI":"10.1371\/journal.pgen.1000384","volume":"5","author":"BE Madsen","year":"2009","unstructured":"Madsen, B. E. & Browning, S. R. A groupwise association test for rare mutations using a weighted sum statistic. PLoS Genet. 5, e1000384 (2009).","journal-title":"PLoS Genet."},{"key":"1471_CR28","doi-asserted-by":"crossref","first-page":"1227","DOI":"10.1007\/s00439-019-02059-9","volume":"138","author":"X Yang","year":"2019","unstructured":"Yang, X. et al. Rare variants in FANCA induce premature ovarian insufficiency. Hum. Genet. 138, 1227\u20131236 (2019).","journal-title":"Hum. Genet."},{"key":"1471_CR29","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1038\/ng817","volume":"30","author":"H Olbrich","year":"2002","unstructured":"Olbrich, H. et al. Mutations in DNAH5 cause primary ciliary dyskinesia and randomization of left-right asymmetry. Nat. Genet. 30, 143\u2013144 (2002).","journal-title":"Nat. Genet."},{"key":"1471_CR30","doi-asserted-by":"crossref","first-page":"1195","DOI":"10.1016\/j.bbamcr.2017.03.005","volume":"1864","author":"CJ Huang","year":"2017","unstructured":"Huang, C. J. et al. Maternal SENP7 programs meiosis architecture and embryo survival in mouse. Biochim. Biophys. Acta, Mol. Cell. Res. 1864, 1195\u20131206 (2017).","journal-title":"Biochim. Biophys. Acta, Mol. Cell. Res."},{"key":"1471_CR31","doi-asserted-by":"crossref","first-page":"2149","DOI":"10.1038\/s41596-018-0028-3","volume":"13","author":"D Clift","year":"2018","unstructured":"Clift, D., So, C., McEwan, W. A., James, L. C. & Schuh, M. Acute and rapid degradation of endogenous proteins by Trim-Away. Nat. Protoc. 13, 2149\u20132175 (2018).","journal-title":"Nat. Protoc."},{"key":"1471_CR32","doi-asserted-by":"crossref","first-page":"991","DOI":"10.1126\/science.1175326","volume":"326","author":"H Homer","year":"2009","unstructured":"Homer, H., Gui, L. & Carroll, J. A spindle assembly checkpoint protein functions in prophase I arrest and prometaphase progression. Science 326, 991\u2013994 (2009).","journal-title":"Science"},{"key":"1471_CR33","doi-asserted-by":"crossref","DOI":"10.1038\/ncomms7946","volume":"6","author":"SA Touati","year":"2015","unstructured":"Touati, S. A. et al. Mouse oocytes depend on BubR1 for proper chromosome segregation but not for prophase i arrest. Nat. Commun. 6, 6946 (2015).","journal-title":"Nat. Commun."},{"key":"1471_CR34","doi-asserted-by":"crossref","first-page":"1469","DOI":"10.1095\/biolreprod.104.031757","volume":"71","author":"ZJ Lan","year":"2004","unstructured":"Lan, Z. J., Xu, X. & Cooney, A. J. Differential oocyte-specific expression of Cre recombinase activity in GDF-9-iCre, Zp3cre, and Msx2Cre transgenic mice. Biol. Reprod. 71, 1469\u20131474 (2004).","journal-title":"Biol. Reprod."},{"key":"1471_CR35","first-page":"109","volume":"2","author":"H Potter","year":"2016","unstructured":"Potter, H. Beyond trisomy 21: phenotypic variability in people with Down syndrome explained by further chromosome mis-segregation and mosaic aneuploidy. J. Syndr. Chromosom. Abnorm. 2, 109 (2016).","journal-title":"J. Syndr. Chromosom. Abnorm."},{"key":"1471_CR36","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1016\/j.cell.2017.03.021","volume":"169","author":"RR Beach","year":"2017","unstructured":"Beach, R. R. et al. Aneuploidy causes non-genetic individuality. Cell 169, 229\u2013242.e21 (2017).","journal-title":"Cell"},{"key":"1471_CR37","doi-asserted-by":"crossref","first-page":"e50","DOI":"10.1371\/journal.pgen.0020050","volume":"2","author":"RJ Roper","year":"2006","unstructured":"Roper, R. J. & Reeves, R. H. Understanding the basis for Down syndrome phenotypes. PLoS Genet. 2, e50 (2006).","journal-title":"PLoS Genet."},{"key":"1471_CR38","doi-asserted-by":"crossref","first-page":"114","DOI":"10.1038\/nature03503","volume":"435","author":"LC Kapitein","year":"2005","unstructured":"Kapitein, L. C. et al. The bipolar mitotic kinesin Eg5 moves on both microtubules that it crosslinks. Nature 435, 114\u2013118 (2005).","journal-title":"Nature"},{"key":"1471_CR39","doi-asserted-by":"crossref","first-page":"682","DOI":"10.1038\/nrm2774","volume":"10","author":"N Hirokawa","year":"2009","unstructured":"Hirokawa, N., Noda, Y., Tanaka, Y. & Niwa, S. Kinesin superfamily motor proteins and intracellular transport. Nat. Rev. Mol. Cell Biol. 10, 682\u2013696 (2009).","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"1471_CR40","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1083\/jcb.201106149","volume":"195","author":"N Ma","year":"2011","unstructured":"Ma, N., Titus, J., Gable, A., Ross, J. L. & Wadsworth, P. TPX2 regulates the localization and activity of Eg5 in the mammalian mitotic spindle. J. Cell Biol. 195, 87\u201398 (2011).","journal-title":"J. Cell Biol."},{"key":"1471_CR41","doi-asserted-by":"crossref","first-page":"17367","DOI":"10.1074\/jbc.M114.612903","volume":"290","author":"SK Balchand","year":"2015","unstructured":"Balchand, S. K., Mann, B. J., Titus, J., Ross, J. L. & Wadsworth, P. TPX2 inhibits Eg5 by interactions with both motor and microtubule. J. Biol. Chem. 290, 17367\u201317379 (2015).","journal-title":"J. Biol. Chem."},{"key":"1471_CR42","doi-asserted-by":"crossref","first-page":"1254","DOI":"10.1091\/mbc.e11-09-0820","volume":"23","author":"A Gable","year":"2012","unstructured":"Gable, A. et al. Dynamic reorganization of Eg5 in the mammalian spindle throughout mitosis requires dynein and TPX2. Mol. Biol. Cell 23, 1254\u20131266 (2012).","journal-title":"Mol. Biol. Cell"},{"key":"1471_CR43","doi-asserted-by":"crossref","first-page":"385","DOI":"10.1083\/jcb.201401014","volume":"206","author":"KJ Helmke","year":"2014","unstructured":"Helmke, K. J. & Heald, R. TPX2 levels modulate meiotic spindle size and architecture in Xenopus egg extracts. J. Cell Biol. 206, 385\u2013393 (2014).","journal-title":"J. Cell Biol."},{"key":"1471_CR44","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1016\/j.cub.2017.11.046","volume":"28","author":"S Eibes","year":"2018","unstructured":"Eibes, S. et al. Nek9 phosphorylation defines a new role for TPX2 in Eg5-dependent centrosome separation before nuclear envelope breakdown. Curr. Biol. 28, 121\u2013129.e4 (2018).","journal-title":"Curr. Biol."},{"key":"1471_CR45","doi-asserted-by":"crossref","first-page":"R1156","DOI":"10.1016\/j.cub.2015.10.003","volume":"25","author":"P Wadsworth","year":"2015","unstructured":"Wadsworth, P. Tpx2. Curr. Biol. 25, R1156\u2013R1158 (2015).","journal-title":"Curr. Biol."},{"key":"1471_CR46","doi-asserted-by":"crossref","first-page":"983","DOI":"10.1083\/jcb.201607060","volume":"216","author":"R Alfaro-Aco","year":"2017","unstructured":"Alfaro-Aco, R., Thawani, A. & Petry, S. Structural analysis of the role of TPX2 in branching microtubule nucleation. J. Cell Biol. 216, 983\u2013997 (2017).","journal-title":"J. Cell Biol."},{"key":"1471_CR47","doi-asserted-by":"crossref","first-page":"e30959","DOI":"10.7554\/eLife.30959","volume":"6","author":"R Zhang","year":"2017","unstructured":"Zhang, R., Roostalu, J., Surrey, T. & Nogales, E. Structural insight into TPX2-stimulated microtubule assembly. eLife 6, e30959 (2017).","journal-title":"eLife"},{"key":"1471_CR48","doi-asserted-by":"crossref","first-page":"568","DOI":"10.1016\/j.cell.2011.07.031","volume":"146","author":"TS Kitajima","year":"2011","unstructured":"Kitajima, T. S., Ohsugi, M. & Ellenberg, J. Complete kinetochore tracking reveals error-prone homologous chromosome biorientation in mammalian oocytes. Cell 146, 568\u2013581 (2011).","journal-title":"Cell"},{"key":"1471_CR49","doi-asserted-by":"crossref","first-page":"971","DOI":"10.1126\/science.286.5441.971","volume":"286","author":"TU Mayer","year":"1999","unstructured":"Mayer, T. U. et al. Small molecule inhibitor of mitotic spindle bipolarity identified in a phenotype-based screen. Science 286, 971\u2013974 (1999).","journal-title":"Science"},{"key":"1471_CR50","doi-asserted-by":"crossref","first-page":"975","DOI":"10.1083\/jcb.150.5.975","volume":"150","author":"TM Kapoor","year":"2000","unstructured":"Kapoor, T. M., Mayer, T. U., Coughlin, M. L. & Mitchison, T. J. Probing spindle assembly mechanisms with monastrol, a small molecule inhibitor of the mitotic kinesin, Eg5. J. Cell Biol. 150, 975\u2013988 (2000).","journal-title":"J. Cell Biol."},{"key":"1471_CR51","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1016\/j.mrgentox.2004.01.001","volume":"559","author":"JB Mailhes","year":"2004","unstructured":"Mailhes, J. B., Mastromatteo, C. & Fuseler, J. W. Transient exposure to the Eg5 kinesin inhibitor monastrol leads to syntelic orientation of chromosomes and aneuploidy in mouse oocytes. Mutat. Res. 559, 153\u2013167 (2004).","journal-title":"Mutat. Res."},{"key":"1471_CR52","doi-asserted-by":"crossref","DOI":"10.1038\/ncomms8217","volume":"6","author":"D Clift","year":"2015","unstructured":"Clift, D. & Schuh, M. A three-step MTOC fragmentation mechanism facilitates bipolar spindle assembly in mouse oocytes. Nat. Commun. 6, 7217 (2015).","journal-title":"Nat. Commun."},{"key":"1471_CR53","doi-asserted-by":"crossref","first-page":"979","DOI":"10.1091\/mbc.e09-07-0601","volume":"21","author":"N Ma","year":"2010","unstructured":"Ma, N. et al. Poleward transport of TPX2 in the mammalian mitotic spindle requires dynein, Eg5, and microtubule flux. Mol. Biol. Cell 21, 979\u2013988 (2010).","journal-title":"Mol. Biol. Cell"},{"key":"1471_CR54","doi-asserted-by":"crossref","first-page":"e3338","DOI":"10.1371\/journal.pone.0003338","volume":"3","author":"S Brunet","year":"2008","unstructured":"Brunet, S. et al. Meiotic regulation of TPX2 protein levels governs cell cycle progression in mouse oocytes. PLoS ONE 3, e3338 (2008).","journal-title":"PLoS ONE"},{"key":"1471_CR55","doi-asserted-by":"crossref","first-page":"543","DOI":"10.1038\/nrm3417","volume":"13","author":"Z Luo","year":"2012","unstructured":"Luo, Z., Lin, C. & Shilatifard, A. The super elongation complex (SEC) family in transcriptional control. Nat. Rev. Mol. Cell Biol. 13, 543\u2013547 (2012).","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"1471_CR56","doi-asserted-by":"crossref","first-page":"954","DOI":"10.1016\/j.molcel.2011.12.008","volume":"44","author":"ER Smith","year":"2011","unstructured":"Smith, E. R. et al. The little elongation complex regulates small nuclear RNA transcription. Mol. Cell 44, 954\u2013965 (2011).","journal-title":"Mol. Cell"},{"key":"1471_CR57","doi-asserted-by":"crossref","first-page":"493","DOI":"10.1016\/j.molcel.2013.07.003","volume":"51","author":"D Hu","year":"2013","unstructured":"Hu, D. et al. The little elongation complex functions at initiation and elongation phases of snRNA gene transcription. Mol. Cell 51, 493\u2013505 (2013).","journal-title":"Mol. Cell"},{"key":"1471_CR58","doi-asserted-by":"crossref","first-page":"1486","DOI":"10.1101\/gad.2059211","volume":"25","author":"C Lin","year":"2011","unstructured":"Lin, C. et al. Dynamic transcriptional events in embryonic stem cells mediated by the super elongation complex (SEC). Genes Dev. 25, 1486\u20131498 (2011).","journal-title":"Genes Dev."},{"key":"1471_CR59","doi-asserted-by":"crossref","first-page":"88","DOI":"10.1126\/science.288.5463.88","volume":"288","author":"RD Vale","year":"2000","unstructured":"Vale, R. D. & Milligan, R. A. The way things move: looking under the hood of molecular motor proteins. Science 288, 88\u201395 (2000).","journal-title":"Science"},{"key":"1471_CR60","doi-asserted-by":"crossref","first-page":"392","DOI":"10.1016\/S0960-9822(02)00504-3","volume":"6","author":"R Simmons","year":"1996","unstructured":"Simmons, R. Molecular motors: single-molecule mechanics. Curr. Biol. 6, 392\u2013394 (1996).","journal-title":"Curr. Biol."},{"key":"1471_CR61","doi-asserted-by":"crossref","first-page":"1896","DOI":"10.1016\/j.bpj.2014.09.009","volume":"107","author":"G Arpa\u011f","year":"2014","unstructured":"Arpa\u011f, G., Shastry, S., Hancock, W. O. & T\u00fczel, E. Transport by populations of fast and slow kinesins uncovers novel family-dependent motor characteristics important for in vivo function. Biophys. J. 107, 1896\u20131904 (2014).","journal-title":"Biophys. J."},{"key":"1471_CR62","doi-asserted-by":"crossref","first-page":"672","DOI":"10.1038\/330672a0","volume":"330","author":"BM Paschal","year":"1987","unstructured":"Paschal, B. M. et al. Isolated flagellar outer arm dynein translocates brain microtubules in vitro. Nature 330, 672\u2013674 (1987).","journal-title":"Nature"},{"key":"1471_CR63","doi-asserted-by":"crossref","first-page":"224","DOI":"10.1016\/j.ajhg.2012.06.007","volume":"91","author":"S Lee","year":"2012","unstructured":"Lee, S. et al. Optimal unified approach for rare-variant association testing with application to small-sample case-control whole-exome sequencing studies. Am. J. Hum. Genet. 91, 224\u2013237 (2012).","journal-title":"Am. J. Hum. Genet."},{"key":"1471_CR64","doi-asserted-by":"crossref","first-page":"593","DOI":"10.1038\/nature12364","volume":"500","author":"Z Xue","year":"2013","unstructured":"Xue, Z. et al. Genetic programs in human and mouse early embryos revealed by single-cell RNA sequencing. Nature 500, 593\u2013597 (2013).","journal-title":"Nature"},{"key":"1471_CR65","doi-asserted-by":"crossref","first-page":"eaax3969","DOI":"10.1126\/sciadv.aax3969","volume":"6","author":"C Zhou","year":"2020","unstructured":"Zhou, C. et al. The cohesin release factor Wapl interacts with Bub3 to govern SAC activity in female meiosis I. Sci. Adv. 6, eaax3969 (2020).","journal-title":"Sci. Adv."},{"key":"1471_CR66","doi-asserted-by":"crossref","first-page":"e51400","DOI":"10.15252\/embr.202051400","volume":"22","author":"A Courtois","year":"2021","unstructured":"Courtois, A., Yoshida, S., Takenouchi, O., Asai, K. & Kitajima, T. S. Stable kinetochore\u2013microtubule attachments restrict MTOC position and spindle elongation in oocytes. EMBO Rep. 22, e51400 (2021).","journal-title":"EMBO Rep."},{"key":"1471_CR67","volume":"10","author":"LMG Paim","year":"2019","unstructured":"Paim, L. M. G. & FitzHarris, G. Tetraploidy causes chromosomal instability in acentriolar mouse embryos. Nat. Commun. 10, 4834 (2019).","journal-title":"Nat. Commun."},{"key":"1471_CR68","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1530\/jrf.0.0170555","volume":"17","author":"T Pedersen","year":"1968","unstructured":"Pedersen, T. & Peters, H. Proposal for a classification of oocytes and follicles in the mouse ovary. J. Reprod. Fertil. 17, 555\u2013557 (1968).","journal-title":"J. Reprod. Fertil."},{"key":"1471_CR69","doi-asserted-by":"crossref","first-page":"e1005582","DOI":"10.1371\/journal.pgen.1005582","volume":"11","author":"M Fuchs","year":"2015","unstructured":"Fuchs, M. et al. A role for the chaperone complex BAG3-HSPB8 in actin dynamics, spindle orientation and proper chromosome segregation during mitosis. PLoS Genet. 11, e1005582 (2015).","journal-title":"PLoS Genet."},{"key":"1471_CR70","doi-asserted-by":"crossref","first-page":"eaay4858","DOI":"10.1126\/sciadv.aay4858","volume":"6","author":"C Guo","year":"2020","unstructured":"Guo, C. et al. ENL initiates multivalent phase separation of the super elongation complex (SEC) in controlling rapid transcriptional activation. Sci. Adv. 6, eaay4858 (2020).","journal-title":"Sci. Adv."},{"key":"1471_CR71","doi-asserted-by":"crossref","first-page":"4884","DOI":"10.1073\/pnas.89.11.4884","volume":"89","author":"MR Webb","year":"1992","unstructured":"Webb, M. R. A continuous spectrophotometric assay for inorganic phosphate and for measuring phosphate release kinetics in biological systems. Proc. Natl Acad. Sci. USA 89, 4884\u20134887 (1992).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1471_CR72","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1038\/nprot.2014.006","volume":"9","author":"S Picelli","year":"2014","unstructured":"Picelli, S. et al. Full-length RNA-seq from single cells using Smart-seq2. Nat. Protoc. 9, 171\u2013181 (2014).","journal-title":"Nat. Protoc."},{"key":"1471_CR73","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1038\/nmeth.3690","volume":"13","author":"P Strnad","year":"2016","unstructured":"Strnad, P. et al. Inverted light-sheet microscope for imaging mouse pre-implantation development. Nat. Methods 13, 139\u2013142 (2016).","journal-title":"Nat. Methods"},{"key":"1471_CR74","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1038\/nprot.2014.011","volume":"9","author":"H Peng","year":"2014","unstructured":"Peng, H., Bria, A., Zhou, Z., Iannello, G. & Long, F. Extensible visualization and analysis for multidimensional images using Vaa3D. Nat. Protoc. 9, 193\u2013208 (2014).","journal-title":"Nat. Protoc."}],"container-title":["Nature Structural &amp; Molecular Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41594-024-01471-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41594-024-01471-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41594-024-01471-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,2,17]],"date-time":"2025-02-17T23:04:30Z","timestamp":1739833470000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41594-024-01471-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,1,16]]},"references-count":74,"journal-issue":{"issue":"2","published-print":{"date-parts":[[2025,2]]}},"alternative-id":["1471"],"URL":"https:\/\/doi.org\/10.1038\/s41594-024-01471-8","relation":{},"ISSN":["1545-9993","1545-9985"],"issn-type":[{"value":"1545-9993","type":"print"},{"value":"1545-9985","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,1,16]]},"assertion":[{"value":"5 February 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"9 December 2024","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"16 January 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}]}}