{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T20:47:30Z","timestamp":1792010850027,"version":"4.4.0"},"reference-count":141,"publisher":"Springer Science and Business Media LLC","issue":"10","license":[{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T00:00:00Z","timestamp":1791504000000},"content-version":"vor","delay-in-days":8,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat Cell Biol"],"published-print":{"date-parts":[[2026,10]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Genome size varies more than 10,000-fold across eukaryotes, yet how the mitotic spindle adapts to this range remains unknown. Here we show that metaphase plate width scales with genome size following a power law with an exponent of ~1\/3 across eukaryotes. Comparative analysis and a physical model reveal that this scaling arises from the linear relationship between total chromosome volume and genome size, with chromosome number playing a secondary role. The same scaling holds across ploidy levels in cell lines, patient-derived cancer organoids and meiotic systems. Physical model and acute spindle compression demonstrate that chromosome crowding generates interchromosome pushing forces that set metaphase plate width. By contrast, spindle length is regulated by microtubule dynamics, cytoplasmic forces and cell size, indicating that spindle dimensions are independently controlled. The revealed scaling law provides a robust mechanism for accommodating large genomic variation and offers insight into the evolution of open mitosis and mitotic cell rounding, as well as polyploidy tolerance in tumours and during speciation.<\/jats:p>","DOI":"10.1038\/s41556-026-02005-8","type":"journal-article","created":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T09:02:20Z","timestamp":1791536540000},"page":"1996-2007","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Power-law scaling of mitotic spindles with genome sizes across eukaryotes is driven by chromosome crowding"],"prefix":"10.1038","volume":"28","author":[{"ORCID":"https:\/\/orcid.org\/0009-0004-0710-7246","authenticated-orcid":false,"given":"Lovro","family":"Gudlin","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4867-6080","authenticated-orcid":false,"given":"Kruno","family":"Vuku\u0161i\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maja","family":"Novak","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0532-658X","authenticated-orcid":false,"given":"Monika","family":"Trupini\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Matko","family":"Ljulj","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Iva","family":"Dundovi\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5260-8227","authenticated-orcid":false,"given":"Ana","family":"Petelinec","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Loren","family":"Petru\u0161i\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-4457-8646","authenticated-orcid":false,"given":"Anna","family":"Hertel","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thomas","family":"van Ravesteyn","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8749-731X","authenticated-orcid":false,"given":"Marianna","family":"Trakala","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3555-5295","authenticated-orcid":false,"given":"Geert J. P. L.","family":"Kops","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2376-7047","authenticated-orcid":false,"given":"Zuzana","family":"Storchov\u00e1","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1792-6056","authenticated-orcid":false,"given":"Josip","family":"Tamba\u010da","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4313-1081","authenticated-orcid":false,"given":"Nenad","family":"Pavin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1305-7922","authenticated-orcid":false,"given":"Iva M.","family":"Toli\u0107","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,10,9]]},"reference":[{"key":"2005_CR1","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1146\/annurev-cellbio-020520-113246","volume":"36","author":"WF Marshall","year":"2020","unstructured":"Marshall, W. F. Scaling of subcellular structures. Annu. Rev. Cell Dev. Biol. 36, 219\u2013236 (2020).","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"2005_CR2","unstructured":"Alberts, B. et al. Molecular Biology of the Cell (W. W. Norton & Company, 2022)."},{"key":"2005_CR3","doi-asserted-by":"publisher","first-page":"473","DOI":"10.1038\/nrm4025","volume":"16","author":"S Santaguida","year":"2015","unstructured":"Santaguida, S. & Amon, A. Short- and long-term effects of chromosome mis-segregation and aneuploidy. Nat. Rev. Mol. Cell Biol. 16, 473\u2013485 (2015).","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2005_CR4","doi-asserted-by":"crossref","first-page":"D332","DOI":"10.1093\/nar\/gkl828","volume":"35","author":"TR Gregory","year":"2007","unstructured":"Gregory, T. R. et al. Eukaryotic genome size databases. Nucleic Acids Res. 35, D332\u2013D338 (2007).","journal-title":"Nucleic Acids Res."},{"key":"2005_CR5","unstructured":"Helmke, K. J., Heald, R. & Wilbur, J. D. In International Review of Cell and Molecular Biology (ed. Jeon, K. W.) Vol. 306, 83\u2013125 (Academic Press, 2013)."},{"key":"2005_CR6","doi-asserted-by":"crossref","first-page":"1256","DOI":"10.1016\/j.cub.2008.07.092","volume":"18","author":"M W\u00fchr","year":"2008","unstructured":"W\u00fchr, M. et al. Evidence for an upper limit to mitotic spindle length. Curr. Biol. 18, 1256\u20131261 (2008).","journal-title":"Curr. Biol."},{"key":"2005_CR7","doi-asserted-by":"crossref","first-page":"732","DOI":"10.1016\/j.cub.2014.12.060","volume":"25","author":"R Farhadifar","year":"2015","unstructured":"Farhadifar, R. et al. Scaling, selection, and evolutionary dynamics of the mitotic spindle. Curr. Biol. 25, 732\u2013740 (2015).","journal-title":"Curr. Biol."},{"key":"2005_CR8","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.55877","volume":"9","author":"R Farhadifar","year":"2020","unstructured":"Farhadifar, R. et al. Stoichiometric interactions explain spindle dynamics and scaling across 100 million years of nematode evolution. eLife 9, e55877 (2020).","journal-title":"eLife"},{"key":"2005_CR9","doi-asserted-by":"crossref","first-page":"1542","DOI":"10.1016\/j.cub.2015.04.036","volume":"25","author":"ME Crowder","year":"2015","unstructured":"Crowder, M. E. et al. A comparative analysis of spindle morphometrics across metazoans. Curr. Biol. 25, 1542\u20131550 (2015).","journal-title":"Curr. Biol."},{"key":"2005_CR10","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.31149","volume":"7","author":"F Decker","year":"2018","unstructured":"Decker, F., Oriola, D., Dalton, B. & Brugu\u00e9s, J. Autocatalytic microtubule nucleation determines the size and mass of Xenopus laevis egg extract spindles. eLife 7, e31149 (2018).","journal-title":"eLife"},{"key":"2005_CR11","doi-asserted-by":"crossref","first-page":"383","DOI":"10.1042\/EBC20190064","volume":"64","author":"LK Kr\u00fcger","year":"2020","unstructured":"Kr\u00fcger, L. K. & Tran, P. T. Spindle scaling mechanisms. Essays Biochem. 64, 383\u2013396 (2020).","journal-title":"Essays Biochem."},{"key":"2005_CR12","doi-asserted-by":"publisher","first-page":"523","DOI":"10.1038\/s41580-023-00584-0","volume":"24","author":"VA Valdez","year":"2023","unstructured":"Valdez, V. A., Neahring, L., Petry, S. & Dumont, S. Mechanisms underlying spindle assembly and robustness. Nat. Rev. Mol. Cell Biol. 24, 523\u2013542 (2023).","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2005_CR13","doi-asserted-by":"crossref","first-page":"856","DOI":"10.1126\/science.1243147","volume":"342","author":"MC Good","year":"2013","unstructured":"Good, M. C., Vahey, M. D., Skandarajah, A., Fletcher, D. A. & Heald, R. Cytoplasmic volume modulates spindle size during embryogenesis. Science 342, 856\u2013860 (2013).","journal-title":"Science"},{"key":"2005_CR14","doi-asserted-by":"crossref","first-page":"853","DOI":"10.1126\/science.1243110","volume":"342","author":"J Hazel","year":"2013","unstructured":"Hazel, J. et al. Changes in cytoplasmic volume are sufficient to drive spindle scaling. Science 342, 853\u2013856 (2013).","journal-title":"Science"},{"key":"2005_CR15","doi-asserted-by":"crossref","first-page":"4973","DOI":"10.1016\/j.cub.2020.10.093","volume":"30","author":"EM Rieckhoff","year":"2020","unstructured":"Rieckhoff, E. M. et al. Spindle scaling is governed by cell boundary regulation of microtubule nucleation. Curr. Biol. 30, 4973\u20134983 (2020).","journal-title":"Curr. Biol."},{"key":"2005_CR16","doi-asserted-by":"crossref","first-page":"496","DOI":"10.1016\/j.devcel.2018.04.022","volume":"45","author":"B Lacroix","year":"2018","unstructured":"Lacroix, B. et al. Microtubule dynamics scale with cell size to set spindle length and assembly timing. Dev. Cell 45, 496\u2013511 (2018).","journal-title":"Dev. Cell"},{"key":"2005_CR17","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1016\/j.cub.2009.12.050","volume":"20","author":"G Greenan","year":"2010","unstructured":"Greenan, G. et al. Centrosome size sets mitotic spindle length in Caenorhabditis elegans embryos. Curr. Biol. 20, 353\u2013358 (2010).","journal-title":"Curr. Biol."},{"key":"2005_CR18","volume":"12","author":"CY Zhou","year":"2023","unstructured":"Zhou, C. Y. et al. Mitotic chromosomes scale to nuclear-cytoplasmic ratio and cell size in Xenopus. eLife 12, e84360 (2023).","journal-title":"eLife"},{"key":"2005_CR19","doi-asserted-by":"crossref","first-page":"e00290","DOI":"10.7554\/eLife.00290","volume":"2","author":"JD Wilbur","year":"2013","unstructured":"Wilbur, J. D. & Heald, R. Mitotic spindle scaling during Xenopus development by kif2a and importin \u03b1. eLife 2, e00290 (2013).","journal-title":"eLife"},{"key":"2005_CR20","doi-asserted-by":"crossref","first-page":"R749","DOI":"10.1016\/j.cub.2009.07.028","volume":"19","author":"S Dumont","year":"2009","unstructured":"Dumont, S. & Mitchison, T. J. Force and length in the mitotic spindle. Curr. Biol. 19, R749\u2013R761 (2009).","journal-title":"Curr. Biol."},{"key":"2005_CR21","doi-asserted-by":"crossref","first-page":"1411","DOI":"10.1091\/mbc.e12-07-0528","volume":"24","author":"Y Hara","year":"2013","unstructured":"Hara, Y. & Kimura, A. An allometric relationship between mitotic spindle width, spindle length, and ploidy in Caenorhabditis elegans embryos. Mol. Biol. Cell 24, 1411\u20131419 (2013).","journal-title":"Mol. Biol. Cell"},{"key":"2005_CR22","volume":"221","author":"T Kletter","year":"2021","unstructured":"Kletter, T. et al. Volumetric morphometry reveals spindle width as the best predictor of mammalian spindle scaling. J. Cell Biol. 221, e202106170 (2021).","journal-title":"J. Cell Biol."},{"key":"2005_CR23","doi-asserted-by":"crossref","first-page":"1327","DOI":"10.1016\/j.cub.2023.02.021","volume":"33","author":"KE Miller","year":"2023","unstructured":"Miller, K. E., Cadart, C. & Heald, R. Dodecaploid Xenopus longipes provides insight into the emergence of size scaling relationships during development. Curr. Biol. 33, 1327\u20131336 (2023).","journal-title":"Curr. Biol."},{"key":"2005_CR24","doi-asserted-by":"crossref","first-page":"502","DOI":"10.1016\/j.cell.2009.05.027","volume":"138","author":"A Dinarina","year":"2009","unstructured":"Dinarina, A. et al. Chromatin shapes the mitotic spindle. Cell 138, 502\u2013513 (2009).","journal-title":"Cell"},{"key":"2005_CR25","doi-asserted-by":"publisher","first-page":"924","DOI":"10.1038\/ncb1455","volume":"8","author":"J Gaetz","year":"2006","unstructured":"Gaetz, J., Gueroui, Z., Libchaber, A. & Kapoor, T. M. Examining how the spatial organization of chromatin signals influences metaphase spindle assembly. Nat. Cell Biol. 8, 924\u2013932 (2006).","journal-title":"Nat. Cell Biol."},{"key":"2005_CR26","doi-asserted-by":"crossref","first-page":"1578","DOI":"10.1016\/j.cub.2011.08.021","volume":"21","author":"L Gan","year":"2011","unstructured":"Gan, L., Ladinsky, M. S. & Jensen, G. J. Organization of the smallest eukaryotic spindle. Curr. Biol. 21, 1578\u20131583 (2011).","journal-title":"Curr. Biol."},{"key":"2005_CR27","doi-asserted-by":"crossref","first-page":"790","DOI":"10.1016\/j.molcel.2016.10.009","volume":"64","author":"DG Booth","year":"2016","unstructured":"Booth, D. G. et al. 3D-CLEM reveals that a major portion of mitotic chromosomes is not chromatin. Mol. Cell 64, 790\u2013802 (2016).","journal-title":"Mol. Cell"},{"key":"2005_CR28","doi-asserted-by":"crossref","first-page":"684278","DOI":"10.3389\/fcell.2021.684278","volume":"9","author":"EM Kramer","year":"2021","unstructured":"Kramer, E. M., Tayjasanant, P. A. & Cordone, B. Scaling laws for mitotic chromosomes. Front. Cell Dev. Biol. 9, 684278 (2021).","journal-title":"Front. Cell Dev. Biol."},{"key":"2005_CR29","doi-asserted-by":"crossref","DOI":"10.1083\/jcb.202403165","volume":"223","author":"F Cisneros-Soberanis","year":"2024","unstructured":"Cisneros-Soberanis, F. et al. Near millimolar concentration of nucleosomes in mitotic chromosomes from late prometaphase into anaphase. J. Cell Biol. 223, e202403165 (2024).","journal-title":"J. Cell Biol."},{"key":"2005_CR30","doi-asserted-by":"crossref","first-page":"R574","DOI":"10.1016\/j.cub.2021.03.082","volume":"31","author":"P Risteski","year":"2021","unstructured":"Risteski, P., Jagri\u0107, M., Pavin, N. & Toli\u0107, I. M. Biomechanics of chromosome alignment at the spindle midplane. Curr. Biol. 31, R574\u2013R585 (2021).","journal-title":"Curr. Biol."},{"key":"2005_CR31","doi-asserted-by":"crossref","first-page":"13","DOI":"10.3390\/biology6010013","volume":"6","author":"H Maiato","year":"2017","unstructured":"Maiato, H., Gomes, A. M., Sousa, F. & Barisic, M. Mechanisms of chromosome congression during mitosis. Biology 6, 13 (2017).","journal-title":"Biology"},{"key":"2005_CR32","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-34957-4","volume":"13","author":"J Matkovic","year":"2022","unstructured":"Matkovic, J. et al. Kinetochore- and chromosome-driven transition of microtubules into bundles promotes spindle assembly. Nat. Commun. 13, 7307 (2022).","journal-title":"Nat. Commun."},{"key":"2005_CR33","doi-asserted-by":"crossref","DOI":"10.1126\/science.aao6135","volume":"359","author":"JH Gibcus","year":"2018","unstructured":"Gibcus, J. H. et al. A pathway for mitotic chromosome formation. Science 359, eaao6135 (2018).","journal-title":"Science"},{"key":"2005_CR34","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1126\/science.1244508","volume":"343","author":"BJ Wilkins","year":"2014","unstructured":"Wilkins, B. J. et al. A cascade of histone modifications induces chromatin condensation in mitosis. Science 343, 77\u201380 (2014).","journal-title":"Science"},{"key":"2005_CR35","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1038\/s41586-022-05027-y","volume":"609","author":"MWG Schneider","year":"2022","unstructured":"Schneider, M. W. G. et al. A mitotic chromatin phase transition prevents perforation by microtubules. Nature 609, 183\u2013190 (2022).","journal-title":"Nature"},{"key":"2005_CR36","doi-asserted-by":"publisher","first-page":"308","DOI":"10.1038\/nature18610","volume":"535","author":"S Cuylen","year":"2016","unstructured":"Cuylen, S. et al. Ki-67 acts as a biological surfactant to disperse mitotic chromosomes. Nature 535, 308\u2013312 (2016).","journal-title":"Nature"},{"key":"2005_CR37","doi-asserted-by":"crossref","first-page":"eadq7540","DOI":"10.1126\/sciadv.adq7540","volume":"10","author":"CP Kelleher","year":"2024","unstructured":"Kelleher, C. P., Rana, Y. P. & Needleman, D. J. Long-range repulsion between chromosomes in mammalian oocyte spindles. Sci. Adv. 10, eadq7540 (2024).","journal-title":"Sci. Adv."},{"key":"2005_CR38","doi-asserted-by":"crossref","first-page":"2810","DOI":"10.1080\/15384101.2015.1068482","volume":"14","author":"AY Kuznetsova","year":"2015","unstructured":"Kuznetsova, A. Y. et al. Chromosomal instability, tolerance of mitotic errors and multidrug resistance are promoted by tetraploidization in human cells. Cell Cycle 14, 2810\u20132820 (2015).","journal-title":"Cell Cycle"},{"key":"2005_CR39","doi-asserted-by":"publisher","first-page":"486","DOI":"10.1038\/s41586-020-03114-6","volume":"590","author":"Y Cohen-Sharir","year":"2021","unstructured":"Cohen-Sharir, Y. et al. Aneuploidy renders cancer cells vulnerable to mitotic checkpoint inhibition. Nature 590, 486\u2013491 (2021).","journal-title":"Nature"},{"key":"2005_CR40","doi-asserted-by":"crossref","first-page":"1155","DOI":"10.1126\/science.aaa5111","volume":"348","author":"YL Wong","year":"2015","unstructured":"Wong, Y. L. et al. Reversible centriole depletion with an inhibitor of Polo-like kinase 4. Science 348, 1155\u20131160 (2015).","journal-title":"Science"},{"key":"2005_CR41","doi-asserted-by":"publisher","first-page":"146","DOI":"10.1038\/s41586-022-04578-4","volume":"604","author":"S Gemble","year":"2022","unstructured":"Gemble, S. et al. Genetic instability from a single S phase after whole-genome duplication. Nature 604, 146\u2013151 (2022).","journal-title":"Nature"},{"key":"2005_CR42","doi-asserted-by":"publisher","first-page":"278","DOI":"10.1038\/nature08136","volume":"460","author":"NJ Ganem","year":"2009","unstructured":"Ganem, N. J., Godinho, S. A. & Pellman, D. A mechanism linking extra centrosomes to chromosomal instability. Nature 460, 278\u2013282 (2009).","journal-title":"Nature"},{"key":"2005_CR43","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-12772-8","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":"2005_CR44","doi-asserted-by":"crossref","first-page":"3065","DOI":"10.1091\/mbc.e16-05-0268","volume":"27","author":"TA Potapova","year":"2016","unstructured":"Potapova, T. A., Seidel, C. W., Box, A. C., Rancati, G. & Li, R. Transcriptome analysis of tetraploid cells identifies cyclin D2 as a facilitator of adaptation to genome doubling in the presence of p53. Mol. Biol. Cell 27, 3065 (2016).","journal-title":"Mol. Biol. Cell"},{"key":"2005_CR45","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1186\/1471-2164-15-139","volume":"15","author":"M D\u00fcrrbaum","year":"2014","unstructured":"D\u00fcrrbaum, M. et al. Unique features of the transcriptional response to model aneuploidy in human cells. BMC Genomics 15, 139 (2014).","journal-title":"BMC Genomics"},{"key":"2005_CR46","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-33904-7","volume":"13","author":"G Yahya","year":"2022","unstructured":"Yahya, G. et al. Sublinear scaling of the cellular proteome with ploidy. Nat. Commun. 13, 6182 (2022).","journal-title":"Nat. Commun."},{"key":"2005_CR47","doi-asserted-by":"publisher","first-page":"824","DOI":"10.1038\/s41588-019-0399-6","volume":"51","author":"ACF Bolhaqueiro","year":"2019","unstructured":"Bolhaqueiro, A. C. F. et al. Ongoing chromosomal instability and karyotype evolution in human colorectal cancer organoids. Nat. Genet. 51, 824\u2013834 (2019).","journal-title":"Nat. Genet."},{"key":"2005_CR48","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-32465-z","volume":"13","author":"G de Medeiros","year":"2022","unstructured":"de Medeiros, G. et al. Multiscale light-sheet organoid imaging framework. Nat. Commun. 13, 4864 (2022).","journal-title":"Nat. Commun."},{"key":"2005_CR49","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-025-64148-w","volume":"16","author":"K Vuku\u0161i\u0107","year":"2025","unstructured":"Vuku\u0161i\u0107, K. & Toli\u0107, I. M. CENP-E initiates chromosome congression by opposing Aurora kinases to promote end-on attachments. Nat. Commun. 16, 8537 (2025).","journal-title":"Nat. Commun."},{"key":"2005_CR50","doi-asserted-by":"crossref","first-page":"2463","DOI":"10.1083\/jcb.201701151","volume":"217","author":"K Yaguchi","year":"2018","unstructured":"Yaguchi, K. et al. Uncoordinated centrosome cycle underlies the instability of non-diploid somatic cells in mammals. J. Cell Biol. 217, 2463\u20132483 (2018).","journal-title":"J. Cell Biol."},{"key":"2005_CR51","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1002\/(SICI)1097-0169(1997)36:3<291::AID-CM9>3.0.CO;2-A","volume":"36","author":"DA Wise","year":"1997","unstructured":"Wise, D. A. & Brinkley, B. R. Mitosis in cells with unreplicated genomes (MUGs): spindle assembly and behavior of centromere fragments. Cell Motil. Cytoskeleton 36, 291\u2013302 (1997).","journal-title":"Cell Motil. Cytoskeleton"},{"key":"2005_CR52","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. Error-prone chromosome-mediated spindle assembly favors chromosome segregation defects in human oocytes. Science 348, 1143\u20131147 (2015).","journal-title":"Science"},{"key":"2005_CR53","volume":"364","author":"C So","year":"2019","unstructured":"So, C. et al. A liquid-like spindle domain promotes acentrosomal spindle assembly in mammalian oocytes. Science 364, eaat9557 (2019).","journal-title":"Science"},{"key":"2005_CR54","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-12674-9","volume":"10","author":"J Roeles","year":"2019","unstructured":"Roeles, J. & Tsiavaliaris, G. Actin\u2013microtubule interplay coordinates spindle assembly in human oocytes. Nat. Commun. 10, 4651 (2019).","journal-title":"Nat. Commun."},{"key":"2005_CR55","volume":"385","author":"T Wu","year":"2024","unstructured":"Wu, T. et al. Mechanisms of minor pole-mediated spindle bipolarization in human oocytes. Science 385, eado1022 (2024).","journal-title":"Science"},{"key":"2005_CR56","doi-asserted-by":"crossref","first-page":"1251","DOI":"10.1242\/jcs.196188","volume":"130","author":"C Baumann","year":"2017","unstructured":"Baumann, C., Wang, X., Yang, L. & Viveiros, M. M. Error-prone meiotic division and subfertility in mice with oocyte-conditional knockdown of pericentrin. J. Cell Sci. 130, 1251\u20131262 (2017).","journal-title":"J. Cell Sci."},{"key":"2005_CR57","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1016\/j.cub.2023.11.061","volume":"34","author":"BP Mihalas","year":"2024","unstructured":"Mihalas, B. P. et al. Age-dependent loss of cohesion protection in human oocytes. Curr. Biol. 34, 117\u2013131 (2024).","journal-title":"Curr. Biol."},{"key":"2005_CR58","doi-asserted-by":"crossref","first-page":"559","DOI":"10.1530\/rep.1.01216","volume":"132","author":"T Somfai","year":"2006","unstructured":"Somfai, T. et al. Diploid porcine parthenotes produced by inhibition of first polar body extrusion during in vitro maturation of follicular oocytes. Reproduction 132, 559\u2013570 (2006).","journal-title":"Reproduction"},{"key":"2005_CR59","doi-asserted-by":"crossref","first-page":"15393","DOI":"10.1073\/pnas.232442599","volume":"99","author":"MG Poirier","year":"2002","unstructured":"Poirier, M. G. & Marko, J. F. Mitotic chromosomes are chromatin networks without a mechanically contiguous protein scaffold. Proc. Natl Acad. Sci. USA 99, 15393\u201315397 (2002).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2005_CR60","doi-asserted-by":"crossref","first-page":"RP97403","DOI":"10.7554\/eLife.97403","volume":"13","author":"N Liu","year":"2025","unstructured":"Liu, N., Qiang, W., Jordan, P. W., Marko, J. F. & Qiao, H. Cell cycle and age-related modulations of mouse chromosome stiffness. eLife 13, RP97403 (2025).","journal-title":"eLife"},{"key":"2005_CR61","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1083\/jcb.139.1.1","volume":"139","author":"B Houchmandzadeh","year":"1997","unstructured":"Houchmandzadeh, B., Marko, J. F., Chatenay, D. & Libchaber, A. Elasticity and structure of eukaryote chromosomes studied by micromanipulation and micropipette aspiration. J. Cell Biol. 139, 1\u201312 (1997).","journal-title":"J. Cell Biol."},{"key":"2005_CR62","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms10298","volume":"7","author":"J Kajtez","year":"2016","unstructured":"Kajtez, J. et al. Overlap microtubules link sister k-fibres and balance the forces on bi-oriented kinetochores. Nat. Commun. 7, 10298 (2016).","journal-title":"Nat. Commun."},{"key":"2005_CR63","doi-asserted-by":"crossref","first-page":"542","DOI":"10.1083\/jcb.97.2.542","volume":"97","author":"RB Nicklas","year":"1983","unstructured":"Nicklas, R. B. Measurements of the force produced by the mitotic spindle in anaphase. J. Cell Biol. 97, 542\u2013548 (1983).","journal-title":"J. Cell Biol."},{"key":"2005_CR64","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms13221","volume":"7","author":"AA Ye","year":"2016","unstructured":"Ye, A. A., Cane, S. & Maresca, T. J. Chromosome biorientation produces hundreds of piconewtons at a metazoan kinetochore. Nat. Commun. 7, 13221 (2016).","journal-title":"Nat. Commun."},{"key":"2005_CR65","doi-asserted-by":"crossref","first-page":"192","DOI":"10.1016\/j.devcel.2020.11.003","volume":"56","author":"N Pavin","year":"2021","unstructured":"Pavin, N. & Toli\u0107, I. M. Mechanobiology of the mitotic spindle. Dev. Cell 56, 192\u2013201 (2021).","journal-title":"Dev. Cell"},{"key":"2005_CR66","doi-asserted-by":"crossref","first-page":"1086","DOI":"10.1016\/j.cub.2009.05.056","volume":"19","author":"S Dumont","year":"2009","unstructured":"Dumont, S. & Mitchison, T. J. Compression regulates mitotic spindle length by a mechanochemical switch at the poles. Curr. Biol. 19, 1086\u20131095 (2009).","journal-title":"Curr. Biol."},{"key":"2005_CR67","doi-asserted-by":"crossref","first-page":"041101","DOI":"10.1063\/1.4964479","volume":"3","author":"J-W Jiang","year":"2016","unstructured":"Jiang, J.-W., Kim, S. Y. & Park, H. S. Auxetic nanomaterials: recent progress and future development. Appl. Phys. Rev. 3, 041101 (2016).","journal-title":"Appl. Phys. Rev."},{"key":"2005_CR68","doi-asserted-by":"publisher","first-page":"167","DOI":"10.1038\/nmeth.1297","volume":"6","author":"T Itabashi","year":"2009","unstructured":"Itabashi, T. et al. Probing the mechanical architecture of the vertebrate meiotic spindle. Nat. Methods 6, 167\u2013172 (2009).","journal-title":"Nat. Methods"},{"key":"2005_CR69","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1016\/j.cell.2011.07.012","volume":"146","author":"V Magidson","year":"2011","unstructured":"Magidson, V. et al. The spatial arrangement of chromosomes during prometaphase facilitates spindle assembly. Cell 146, 555\u2013567 (2011).","journal-title":"Cell"},{"key":"2005_CR70","doi-asserted-by":"publisher","first-page":"959","DOI":"10.1038\/s41556-025-01678-x","volume":"27","author":"T Kletter","year":"2025","unstructured":"Kletter, T. et al. Cell state-specific cytoplasmic density controls spindle architecture and scaling. Nat. Cell Biol. 27, 959\u2013971 (2025).","journal-title":"Nat. Cell Biol."},{"key":"2005_CR71","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2403153121","volume":"121","author":"S Iida","year":"2024","unstructured":"Iida, S. et al. Orientation-independent-DIC imaging reveals that a transient rise in depletion attraction contributes to mitotic chromosome condensation. Proc. Natl Acad. Sci. USA 121, e2403153121 (2024).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2005_CR72","doi-asserted-by":"crossref","first-page":"2480","DOI":"10.1016\/j.cub.2022.04.035","volume":"32","author":"M Trupini\u0107","year":"2022","unstructured":"Trupini\u0107, M. et al. The chirality of the mitotic spindle provides a mechanical response to forces and depends on microtubule motors and augmin. Curr. Biol. 32, 2480\u20132493 (2022).","journal-title":"Curr. Biol."},{"key":"2005_CR73","doi-asserted-by":"crossref","first-page":"1979","DOI":"10.1016\/j.cub.2005.09.054","volume":"15","author":"G Goshima","year":"2005","unstructured":"Goshima, G., Wollman, R., Stuurman, N., Scholey, J. M. & Vale, R. D. Length control of the metaphase spindle. Curr. Biol. 15, 1979\u20131988 (2005).","journal-title":"Curr. Biol."},{"key":"2005_CR74","doi-asserted-by":"crossref","first-page":"111753","DOI":"10.1016\/j.celrep.2022.111753","volume":"41","author":"Y Kakui","year":"2022","unstructured":"Kakui, Y. et al. Chromosome arm length, and a species-specific determinant, define chromosome arm width. Cell Rep. 41, 111753 (2022).","journal-title":"Cell Rep."},{"key":"2005_CR75","doi-asserted-by":"publisher","first-page":"445","DOI":"10.1038\/s41580-021-00349-7","volume":"22","author":"IF Davidson","year":"2021","unstructured":"Davidson, I. F. & Peters, J.-M. Genome folding through loop extrusion by SMC complexes. Nat. Rev. Mol. Cell Biol. 22, 445\u2013464 (2021).","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2005_CR76","doi-asserted-by":"crossref","first-page":"a018713","DOI":"10.1101\/cshperspect.a018713","volume":"6","author":"E Seto","year":"2014","unstructured":"Seto, E. & Yoshida, M. Erasers of histone acetylation: the histone deacetylase enzymes. Cold Spring Harb. Perspect. Biol. 6, a018713 (2014).","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"2005_CR77","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1016\/j.semcdb.2021.03.014","volume":"117","author":"JR Paulson","year":"2021","unstructured":"Paulson, J. R., Hudson, D. F., Cisneros-Soberanis, F. & Earnshaw, W. C. Mitotic chromosomes. Semin. Cell Dev. Biol. 117, 7\u201329 (2021).","journal-title":"Semin. Cell Dev. Biol."},{"key":"2005_CR78","doi-asserted-by":"crossref","first-page":"270","DOI":"10.1016\/j.devcel.2013.03.014","volume":"25","author":"OM Lancaster","year":"2013","unstructured":"Lancaster, O. M. et al. Mitotic rounding alters cell geometry to ensure efficient bipolar spindle formation. Dev. Cell 25, 270\u2013283 (2013).","journal-title":"Dev. Cell"},{"key":"2005_CR79","doi-asserted-by":"crossref","first-page":"11258","DOI":"10.1073\/pnas.1502029112","volume":"112","author":"CJ Cattin","year":"2015","unstructured":"Cattin, C. J. et al. Mechanical control of mitotic progression in single animal cells. Proc. Natl Acad. Sci. USA 112, 11258\u201311263 (2015).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2005_CR80","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2404551121","volume":"121","author":"G Bastianello","year":"2024","unstructured":"Bastianello, G. et al. Mechanical stress during confined migration causes aberrant mitoses and c-MYC amplification. Proc. Natl Acad. Sci. USA 121, e2404551121 (2024).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2005_CR81","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1016\/j.devcel.2014.04.009","volume":"29","author":"C Cadart","year":"2014","unstructured":"Cadart, C., Zlotek-Zlotkiewicz, E., Le Berre, M., Piel, M. & Matthews, H. K. Exploring the function of cell shape and size during mitosis. Dev. Cell 29, 159\u2013169 (2014).","journal-title":"Dev. Cell"},{"key":"2005_CR82","doi-asserted-by":"crossref","first-page":"R1099","DOI":"10.1016\/j.cub.2014.10.011","volume":"24","author":"S Sazer","year":"2014","unstructured":"Sazer, S., Lynch, M. & Needleman, D. Deciphering the evolutionary history of open and closed mitosis. Curr. Biol. 24, R1099\u2013R1103 (2014).","journal-title":"Curr. Biol."},{"key":"2005_CR83","doi-asserted-by":"publisher","first-page":"226","DOI":"10.1038\/nature09642","volume":"469","author":"MP Stewart","year":"2011","unstructured":"Stewart, M. P. et al. Hydrostatic pressure and the actomyosin cortex drive mitotic cell rounding. Nature 469, 226\u2013230 (2011).","journal-title":"Nature"},{"key":"2005_CR84","doi-asserted-by":"crossref","first-page":"466","DOI":"10.1016\/j.devcel.2022.02.001","volume":"57","author":"AT Molines","year":"2022","unstructured":"Molines, A. T. et al. Physical properties of the cytoplasm modulate the rates of microtubule polymerization and depolymerization. Dev. Cell 57, 466\u2013479 (2022).","journal-title":"Dev. Cell"},{"key":"2005_CR85","doi-asserted-by":"publisher","DOI":"10.1186\/s12915-017-0478-z","volume":"16","author":"N Mchedlishvili","year":"2018","unstructured":"Mchedlishvili, N., Matthews, H. K., Corrigan, A. & Baum, B. Two-step interphase microtubule disassembly aids spindle morphogenesis. BMC Biol. 16, 14 (2018).","journal-title":"BMC Biol."},{"key":"2005_CR86","doi-asserted-by":"publisher","first-page":"82","DOI":"10.1038\/ncb2640","volume":"15","author":"I Kalinina","year":"2013","unstructured":"Kalinina, I. et al. Pivoting of microtubules around the spindle pole accelerates kinetochore capture. Nat. Cell Biol. 15, 82\u201387 (2013).","journal-title":"Nat. Cell Biol."},{"key":"2005_CR87","volume":"9","author":"P Suresh","year":"2020","unstructured":"Suresh, P., Long, A. F. & Dumont, S. Microneedle manipulation of the mammalian spindle reveals specialized, short-lived reinforcement near chromosomes. eLife 9, e53807 (2020).","journal-title":"eLife"},{"key":"2005_CR88","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-026-69830-1","volume":"17","author":"I Koprivec","year":"2026","unstructured":"Koprivec, I. et al. Polar chromosomes are rescued from missegregation by spindle elongation-driven microtubule pivoting. Nat. Commun. 17, 2049 (2026).","journal-title":"Nat. Commun."},{"key":"2005_CR89","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1083\/jcb.201202135","volume":"198","author":"A Courtois","year":"2012","unstructured":"Courtois, A., Schuh, M., Ellenberg, J. & Hiiragi, T. The transition from meiotic to mitotic spindle assembly is gradual during early mammalian development. J. Cell Biol. 198, 357\u2013370 (2012).","journal-title":"J. Cell Biol."},{"key":"2005_CR90","doi-asserted-by":"crossref","first-page":"2442","DOI":"10.1091\/mbc.e13-01-0043","volume":"24","author":"Y Hara","year":"2013","unstructured":"Hara, Y., Iwabuchi, M., Ohsumi, K. & Kimura, A. Intranuclear DNA density affects chromosome condensation in metazoans. Mol. Biol. Cell 24, 2442\u20132453 (2013).","journal-title":"Mol. Biol. Cell"},{"key":"2005_CR91","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-49815-8","volume":"15","author":"Y Ono","year":"2024","unstructured":"Ono, Y. et al. Shape of the first mitotic spindles impacts multinucleation in human embryos. Nat. Commun. 15, 5381 (2024).","journal-title":"Nat. Commun."},{"key":"2005_CR92","doi-asserted-by":"crossref","DOI":"10.1530\/REP-23-0281","volume":"167","author":"M Fluks","year":"2024","unstructured":"Fluks, M., Milewski, R., Tamborski, S., Szkulmowski, M. & Ajduk, A. Spindle shape and volume differ in high- and low-quality metaphase II oocytes. Reproduction 167, e230281 (2024).","journal-title":"Reproduction"},{"key":"2005_CR93","doi-asserted-by":"publisher","first-page":"541","DOI":"10.1038\/nature05178","volume":"443","author":"Z Storchov\u00e1","year":"2006","unstructured":"Storchov\u00e1, Z. et al. Genome-wide genetic analysis of polyploidy in yeast. Nature 443, 541\u2013547 (2006).","journal-title":"Nature"},{"key":"2005_CR94","doi-asserted-by":"crossref","first-page":"3064","DOI":"10.1091\/mbc.e05-02-0174","volume":"16","author":"TJ Mitchison","year":"2005","unstructured":"Mitchison, T. J. et al. Roles of polymerization dynamics, opposed motors, and a tensile element in governing the length of Xenopus extract meiotic spindles. Mol. Biol. Cell 16, 3064\u20133076 (2005).","journal-title":"Mol. Biol. Cell"},{"key":"2005_CR95","doi-asserted-by":"publisher","first-page":"707","DOI":"10.1038\/nature09414","volume":"467","author":"AW Duncan","year":"2010","unstructured":"Duncan, A. W. et al. The ploidy conveyor of mature hepatocytes as a source of genetic variation. Nature 467, 707\u2013710 (2010).","journal-title":"Nature"},{"key":"2005_CR96","doi-asserted-by":"publisher","first-page":"1403","DOI":"10.1038\/cdd.2010.145","volume":"18","author":"I Vitale","year":"2011","unstructured":"Vitale, I. et al. Illicit survival of cancer cells during polyploidization and depolyploidization. Cell Death Differ. 18, 1403\u20131413 (2011).","journal-title":"Cell Death Differ."},{"key":"2005_CR97","doi-asserted-by":"crossref","first-page":"2126","DOI":"10.1158\/0008-5472.CAN-24-3718","volume":"86","author":"M Bloomfield","year":"2026","unstructured":"Bloomfield, M. et al. Cell and nuclear size are associated with chromosomal instability and tumorigenicity in cancer cells that undergo whole genome doubling. Cancer Res. 86, 2126\u20132142 (2026).","journal-title":"Cancer Res."},{"key":"2005_CR98","doi-asserted-by":"publisher","first-page":"411","DOI":"10.1038\/nrg.2017.26","volume":"18","author":"Y Van de Peer","year":"2017","unstructured":"Van de Peer, Y., Mizrachi, E. & Marchal, K. The evolutionary significance of polyploidy. Nat. Rev. Genet. 18, 411\u2013424 (2017).","journal-title":"Nat. Rev. Genet."},{"key":"2005_CR99","doi-asserted-by":"publisher","first-page":"116","DOI":"10.1038\/s41586-024-07430-z","volume":"630","author":"H Shah","year":"2024","unstructured":"Shah, H. et al. Life-cycle-coupled evolution of mitosis in close relatives of animals. Nature 630, 116\u2013122 (2024).","journal-title":"Nature"},{"key":"2005_CR100","doi-asserted-by":"crossref","first-page":"1247","DOI":"10.1016\/j.cub.2022.01.034","volume":"32","author":"KB Velle","year":"2022","unstructured":"Velle, K. B. et al. Naegleria\u2019s mitotic spindles are built from unique tubulins and highlight core spindle features. Curr. Biol. 32, 1247\u20131261 (2022).","journal-title":"Curr. Biol."},{"key":"2005_CR101","doi-asserted-by":"publisher","first-page":"403","DOI":"10.1007\/s10577-012-9284-1","volume":"20","author":"H Zhang","year":"2012","unstructured":"Zhang, H. & Dawe, R. K. Total centromere size and genome size are strongly correlated in ten grass species. Chromosome Res. 20, 403\u2013412 (2012).","journal-title":"Chromosome Res."},{"key":"2005_CR102","doi-asserted-by":"crossref","first-page":"1717","DOI":"10.1242\/jcs.03442","volume":"120","author":"CB O\u2019Connell","year":"2007","unstructured":"O\u2019Connell, C. B. & Khodjakov, A. L. Cooperative mechanisms of mitotic spindle formation. J. Cell Sci. 120, 1717\u20131722 (2007).","journal-title":"J. Cell Sci."},{"key":"2005_CR103","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-018-06005-7","volume":"9","author":"M Novak","year":"2018","unstructured":"Novak, M. et al. The mitotic spindle is chiral due to torques within microtubule bundles. Nat. Commun. 9, 3571 (2018).","journal-title":"Nat. Commun."},{"key":"2005_CR104","unstructured":"Landau, L. D. & Lifshits, E. M. Theory of Elasticity Vol. 7 (Pergamon, 1959)."},{"key":"2005_CR105","doi-asserted-by":"publisher","unstructured":"Johnson, K. L. Contact Mechanics (Cambridge Univ. Press, 1985); https:\/\/doi.org\/10.1017\/CBO9781139171731","DOI":"10.1017\/CBO9781139171731"},{"key":"2005_CR106","doi-asserted-by":"crossref","first-page":"502","DOI":"10.1002\/mma.1670080133","volume":"8","author":"Z Tutek","year":"1986","unstructured":"Tutek, Z. & Aganovi\u0107, I. A justification of the one-dimensional linear model of elastic beam. Math. Methods Appl. Sci. 8, 502\u2013515 (1986).","journal-title":"Math. Methods Appl. Sci."},{"key":"2005_CR107","doi-asserted-by":"crossref","first-page":"e2231","DOI":"10.1002\/nla.2231","volume":"26","author":"L Grubi\u0161i\u0107","year":"2019","unstructured":"Grubi\u0161i\u0107, L. & Tamba\u010da, J. Direct solution method for the equilibrium problem for elastic stents. Numer. Linear Algebra Appl. 26, e2231 (2019).","journal-title":"Numer. Linear Algebra Appl."},{"key":"2005_CR108","doi-asserted-by":"crossref","first-page":"112497","DOI":"10.1016\/j.jcp.2023.112497","volume":"494","author":"S \u010cani\u0107","year":"2023","unstructured":"\u010cani\u0107, S., Grubi\u0161i\u0107, L., Ljulj, M., Mareti\u0107, M. & Tamba\u010da, J. Geometric optimization of vascular stents modeled as networks of 1D rods. J. Comput. Phys. 494, 112497 (2023).","journal-title":"J. Comput. Phys."},{"key":"2005_CR109","doi-asserted-by":"publisher","first-page":"523","DOI":"10.1186\/1029-242X-2014-523","volume":"2014","author":"H Lou","year":"2014","unstructured":"Lou, H. Sharp constants for inequalities of Poincar\u00e9 type: an application of optimal control theory. J. Inequal. Appl. 2014, 523 (2014).","journal-title":"J. Inequal. Appl."},{"key":"2005_CR110","doi-asserted-by":"crossref","first-page":"923","DOI":"10.1083\/jcb.120.4.923","volume":"120","author":"F Gittes","year":"1993","unstructured":"Gittes, F., Mickey, B., Nettleton, J. & Howard, J. Flexural rigidity of microtubules and actin filaments measured from thermal fluctuations in shape. J. Cell Biol. 120, 923\u2013934 (1993).","journal-title":"J. Cell Biol."},{"key":"2005_CR111","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1242\/jcs.104.2.261","volume":"104","author":"KL Wendell","year":"1993","unstructured":"Wendell, K. L., Wilson, L. & Jordan, M. A. Mitotic block in HeLa cells by vinblastine: ultrastructural changes in kinetochore\u2013microtubule attachment and in centrosomes. J. Cell Sci. 104, 261\u2013274 (1993).","journal-title":"J. Cell Sci."},{"key":"2005_CR112","doi-asserted-by":"crossref","first-page":"665","DOI":"10.1083\/jcb.200909005","volume":"188","author":"K Jaqaman","year":"2010","unstructured":"Jaqaman, K. et al. Kinetochore alignment within the metaphase plate is regulated by centromere stiffness and microtubule depolymerases. J. Cell Biol. 188, 665\u2013679 (2010).","journal-title":"J. Cell Biol."},{"key":"2005_CR113","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1023\/B:TRAG.0000026071.41735.8e","volume":"13","author":"H Higginbotham","year":"2004","unstructured":"Higginbotham, H., Bielas, S., Tanaka, T. & Gleeson, J. G. Transgenic mouse line with green-fluorescent protein-labeled centrin 2 allows visualization of the centrosome in living Cells. Transgenic Res. 13, 155\u2013164 (2004).","journal-title":"Transgenic Res."},{"key":"2005_CR114","doi-asserted-by":"crossref","first-page":"783","DOI":"10.1016\/j.neuron.2015.03.027","volume":"86","author":"SP Peron","year":"2015","unstructured":"Peron, S. P., Freeman, J., Iyer, V., Guo, C. & Svoboda, K. A cellular resolution map of barrel cortex activity during tactile behavior. Neuron 86, 783\u2013799 (2015).","journal-title":"Neuron"},{"key":"2005_CR115","doi-asserted-by":"crossref","first-page":"jcs247940","DOI":"10.1242\/jcs.247940","volume":"133","author":"SJ Scott","year":"2020","unstructured":"Scott, S. J., Suvarna, K. S. & D\u2019Avino, P. P. Synchronization of human retinal pigment epithelial-1 cells in mitosis. J. Cell Sci. 133, jcs247940 (2020).","journal-title":"J. Cell Sci."},{"key":"2005_CR116","doi-asserted-by":"crossref","DOI":"10.1098\/rstb.2013.0014","volume":"368","author":"A Fatehullah","year":"2013","unstructured":"Fatehullah, A., Appleton, P. L. & N\u00e4thke, I. S. Cell and tissue polarity in the intestinal tract during tumourigenesis: cells still know the right way up, but tissue organization is lost. Philos. Trans. R. Soc. Lond. B 368, 20130014 (2013).","journal-title":"Philos. Trans. R. Soc. Lond. B"},{"key":"2005_CR117","unstructured":"Lee, I. A. & Preacher, K. J. Calculation for the test of the difference between two dependent correlations with one variable in common [computer software]; https:\/\/quantpsy.org\/corrtest\/corrtest2.htm (2013)."},{"key":"2005_CR118","doi-asserted-by":"crossref","first-page":"e49801","DOI":"10.7554\/eLife.49801","volume":"8","author":"O Dudin","year":"2019","unstructured":"Dudin, O. et al. A unicellular relative of animals generates a layer of polarized cells by actomyosin-dependent cellularization. eLife 8, e49801 (2019).","journal-title":"eLife"},{"key":"2005_CR119","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1083\/jcb.120.1.141","volume":"120","author":"R Ding","year":"1993","unstructured":"Ding, R., McDonald, K. L. & McIntosh, J. R. Three-dimensional reconstruction and analysis of mitotic spindles from the yeast, Schizosaccharomyces pombe. J. Cell Biol. 120, 141\u2013151 (1993).","journal-title":"J. Cell Biol."},{"key":"2005_CR120","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1016\/j.bpj.2021.12.019","volume":"121","author":"P Zareiesfandabadi","year":"2022","unstructured":"Zareiesfandabadi, P. & Elting, M. W. Force by minus-end motors Dhc1 and Klp2 collapses the S. pombe spindle after laser ablation. Biophys. J. 121, 263\u2013276 (2022).","journal-title":"Biophys. J."},{"key":"2005_CR121","doi-asserted-by":"crossref","first-page":"1601","DOI":"10.1083\/jcb.129.6.1601","volume":"129","author":"M Winey","year":"1995","unstructured":"Winey, M. et al. Three-dimensional ultrastructural analysis of the Saccharomyces cerevisiae mitotic spindle. J. Cell Biol. 129, 1601\u20131615 (1995).","journal-title":"J. Cell Biol."},{"key":"2005_CR122","doi-asserted-by":"crossref","first-page":"2560","DOI":"10.1091\/mbc.e11-07-0651","volume":"23","author":"JS Verdaasdonk","year":"2012","unstructured":"Verdaasdonk, J. S., Gardner, R., Stephens, A. D., Yeh, E. & Bloom, K. Tension-dependent nucleosome remodeling at the pericentromere in yeast. Mol. Biol. Cell 23, 2560\u20132570 (2012).","journal-title":"Mol. Biol. Cell"},{"key":"2005_CR123","doi-asserted-by":"crossref","first-page":"e1008584","DOI":"10.1371\/journal.pgen.1008584","volume":"16","author":"A P\u00e9rez-Posada","year":"2020","unstructured":"P\u00e9rez-Posada, A., Dudin, O., Oca\u00f1a-Pallar\u00e8s, E., Ruiz-Trillo, I. & Ondracka, A. Cell cycle transcriptomics of Capsaspora provides insights into the evolution of cyclin-CDK machinery. PLoS Genet. 16, e1008584 (2020).","journal-title":"PLoS Genet."},{"key":"2005_CR124","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1083\/jcb.68.1.113","volume":"68","author":"PB Moens","year":"1976","unstructured":"Moens, P. B. Spindle and kinetochore morphology of Dictyostelium discoideum. J. Cell Biol. 68, 113\u2013122 (1976).","journal-title":"J. Cell Biol."},{"key":"2005_CR125","doi-asserted-by":"publisher","DOI":"10.1186\/1747-1028-8-4","volume":"8","author":"A Catalano","year":"2013","unstructured":"Catalano, A. & O\u2019Day, D. H. Rad53 homologue forkhead-associated kinase A (FhkA) and Ca2+-binding protein 4a (CBP4a) are nucleolar proteins that differentially redistribute during mitosis in Dictyostelium. Cell Div. 8, 4 (2013).","journal-title":"Cell Div."},{"key":"2005_CR126","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1091\/mbc.E19-07-0405","volume":"31","author":"E O\u2019Toole","year":"2020","unstructured":"O\u2019Toole, E., Morphew, M. & McIntosh, J. R. Electron tomography reveals aspects of spindle structure important for mechanical stability at metaphase. Mol. Biol. Cell 31, 184\u2013195 (2020).","journal-title":"Mol. Biol. Cell"},{"key":"2005_CR127","doi-asserted-by":"crossref","first-page":"e53940","DOI":"10.1371\/journal.pone.0053940","volume":"8","author":"JM Esparza","year":"2013","unstructured":"Esparza, J. M. et al. Katanin localization requires triplet microtubules in Chlamydomonas reinhardtii. PLoS ONE 8, e53940 (2013).","journal-title":"PLoS ONE"},{"key":"2005_CR128","doi-asserted-by":"crossref","first-page":"226","DOI":"10.1105\/tpc.106.047613","volume":"19","author":"JC Ambrose","year":"2007","unstructured":"Ambrose, J. C. & Cyr, R. The kinesin ATK5 functions in early spindle assembly in Arabidopsis. Plant Cell 19, 226\u2013236 (2007).","journal-title":"Plant Cell"},{"key":"2005_CR129","doi-asserted-by":"crossref","first-page":"e53995","DOI":"10.15252\/embr.202153995","volume":"23","author":"M Romeiro Motta","year":"2022","unstructured":"Romeiro Motta, M. et al. B1-type cyclins control microtubule organization during cell division in Arabidopsis. EMBO Rep. 23, e53995 (2022).","journal-title":"EMBO Rep."},{"key":"2005_CR130","doi-asserted-by":"crossref","first-page":"2371","DOI":"10.1091\/mbc.e08-11-1127","volume":"20","author":"SA Ribeiro","year":"2009","unstructured":"Ribeiro, S. A. et al. Condensin regulates the stiffness of vertebrate centromeres. Mol. Biol. Cell 20, 2371\u20132380 (2009).","journal-title":"Mol. Biol. Cell"},{"key":"2005_CR131","doi-asserted-by":"crossref","first-page":"1035","DOI":"10.1091\/mbc.e11-09-0767","volume":"23","author":"X Wan","year":"2012","unstructured":"Wan, X., Cimini, D., Cameron, L. A. & Salmon, E. D. The coupling between sister kinetochore directional instability and oscillations in centromere stretch in metaphase PtK1 cells. Mol. Biol. Cell 23, 1035\u20131046 (2012).","journal-title":"Mol. Biol. Cell"},{"key":"2005_CR132","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1126\/science.1082177","volume":"300","author":"CL Rieder","year":"2003","unstructured":"Rieder, C. L. & Khodjakov, A. Mitosis through the microscope: advances in seeing inside live dividing cells. Science 300, 91\u201396 (2003).","journal-title":"Science"},{"key":"2005_CR133","doi-asserted-by":"crossref","first-page":"110610","DOI":"10.1016\/j.celrep.2022.110610","volume":"39","author":"AC Almeida","year":"2022","unstructured":"Almeida, A. C. et al. Augmin-dependent microtubule self-organization drives kinetochore fiber maturation in mammals. Cell Rep. 39, 110610 (2022).","journal-title":"Cell Rep."},{"key":"2005_CR134","doi-asserted-by":"crossref","first-page":"1344","DOI":"10.1016\/j.cub.2018.03.023","volume":"28","author":"D Drpic","year":"2018","unstructured":"Drpic, D. et al. Chromosome segregation is biased by kinetochore size. Curr. Biol. 28, 1344\u20131356 (2018).","journal-title":"Curr. Biol."},{"key":"2005_CR135","doi-asserted-by":"crossref","first-page":"569","DOI":"10.1083\/jcb.119.3.569","volume":"119","author":"TJ Mitchison","year":"1992","unstructured":"Mitchison, T. J. & Salmon, E. D. Poleward kinetochore fiber movement occurs during both metaphase and anaphase-A in newt lung cell mitosis. J. Cell Biol. 119, 569\u2013582 (1992).","journal-title":"J. Cell Biol."},{"key":"2005_CR136","doi-asserted-by":"crossref","first-page":"2823","DOI":"10.1242\/jcs.109.12.2823","volume":"109","author":"JC Waters","year":"1996","unstructured":"Waters, J. C., Skibbens, R. V. & Salmon, E. D. Oscillating mitotic newt lung cell kinetochores are, on average, under tension and rarely push. J. Cell Sci. 109, 2823\u20132831 (1996).","journal-title":"J. Cell Sci."},{"key":"2005_CR137","doi-asserted-by":"crossref","first-page":"e26036","DOI":"10.7554\/eLife.26036","volume":"6","author":"X Grau-Bov\u00e9","year":"2017","unstructured":"Grau-Bov\u00e9, X. et al. Dynamics of genomic innovation in the unicellular ancestry of animals. eLife 6, e26036 (2017).","journal-title":"eLife"},{"key":"2005_CR138","doi-asserted-by":"crossref","first-page":"631","DOI":"10.1016\/j.cell.2010.01.032","volume":"140","author":"LK Fritz-Laylin","year":"2010","unstructured":"Fritz-Laylin, L. K. et al. The genome of Naegleria gruberi illuminates early eukaryotic versatility. Cell 140, 631\u2013642 (2010).","journal-title":"Cell"},{"key":"2005_CR139","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-021-99386-7","volume":"11","author":"K Pla\u010dkov\u00e1","year":"2021","unstructured":"Pla\u010dkov\u00e1, K., Bure\u0161, P. & Zedek, F. Centromere size scales with genome size across eukaryotes. Sci. Rep. 11, 19811 (2021).","journal-title":"Sci. Rep."},{"key":"2005_CR140","doi-asserted-by":"publisher","unstructured":"Gudlin, L. et al. A universal scaling law for mitotic spindles across eukaryotes driven by chromosome crowding. Figshare https:\/\/doi.org\/10.6084\/m9.figshare.32112142 (2026).","DOI":"10.6084\/m9.figshare.32112142"},{"key":"2005_CR141","doi-asserted-by":"publisher","unstructured":"Gudlin, L. et al. MATLAB code for: A universal scaling law for mitotic spindles across eukaryotes driven by chromosome crowding (v1.0.0). Zenodo https:\/\/doi.org\/10.5281\/zenodo.19708772 (2026).","DOI":"10.5281\/zenodo.19708772"}],"container-title":["Nature Cell Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41556-026-02005-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41556-026-02005-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41556-026-02005-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T20:12:06Z","timestamp":1792008726000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41556-026-02005-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10]]},"references-count":141,"journal-issue":{"issue":"10","published-print":{"date-parts":[[2026,10]]}},"alternative-id":["2005"],"URL":"https:\/\/doi.org\/10.1038\/s41556-026-02005-8","relation":{"has-preprint":[{"id-type":"doi","id":"10.21203\/rs.3.rs-6194615\/v1","asserted-by":"object"}]},"ISSN":["1465-7392","1476-4679"],"issn-type":[{"value":"1465-7392","type":"print"},{"value":"1476-4679","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10]]},"assertion":[{"value":"15 April 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"31 May 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"9 October 2026","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","label":"Competing interests","group":{"name":"EthicsHeading","label":"Ethics"}}]}}