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During late mitosis, spindle microtubules undergo drastic elongation in a process called anaphase B. Two kinesin motors, Kinesin-5 and Kinesin-6, are thought to generate outward forces to drive spindle elongation, and the microtubule crosslinker Ase1\/PRC1 maintains structural integrity of antiparallel microtubules. However, how these three proteins orchestrate this process remains unknown. Here we explore the functional interplay among fission yeast Kinesin-5\/Cut7, Kinesin-6\/Klp9 and Ase1. Using total internal reflection fluorescence microscopy, we show that Klp9 forms homotetramers and that Klp9 is a processive plus end-directed motor.\n                    <jats:italic>klp9\u0394ase1\u0394<\/jats:italic>\n                    is synthetically lethal. Surprisingly, this lethality is not ascribable to the defective motor activity of Klp9; instead, it is dependent upon a nuclear localisation signal and coiled coil domains within the non-motor region. We isolated a\n                    <jats:italic>cut7<\/jats:italic>\n                    mutant (\n                    <jats:italic>cut7-122<\/jats:italic>\n                    ) that displays temperature sensitivity only in the absence of Klp9. Interestingly,\n                    <jats:italic>cut7-122<\/jats:italic>\n                    alone is impaired in spindle elongation during anaphase B, and furthermore,\n                    <jats:italic>cut7-122klp9\u0394<\/jats:italic>\n                    double mutants exhibit additive defects. We propose that Klp9 plays dual roles during anaphase B; one is motor-dependent that collaborates with Cut7 in force generation, while the other is motor-independent that ensures structural integrity of spindle microtubules together with Ase1.\n                  <\/jats:p>","DOI":"10.1038\/s41598-019-43774-7","type":"journal-article","created":{"date-parts":[[2019,5,14]],"date-time":"2019-05-14T06:04:05Z","timestamp":1557813845000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":32,"title":["Kinesin-6 Klp9 plays motor-dependent and -independent roles in collaboration with Kinesin-5 Cut7 and the microtubule crosslinker Ase1 in fission yeast"],"prefix":"10.1038","volume":"9","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1723-890X","authenticated-orcid":false,"given":"Masashi","family":"Yukawa","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Masaki","family":"Okazaki","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yasuhiro","family":"Teratani","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ken\u2019ya","family":"Furuta","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Takashi","family":"Toda","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2019,5,14]]},"reference":[{"issue":"1650","key":"43774_CR1","doi-asserted-by":"publisher","first-page":"20130467","DOI":"10.1098\/rstb.2013.0467","volume":"369","author":"S. A. Godinho","year":"2014","unstructured":"Godinho, S. A. & Pellman, D. Causes and consequences of centrosome abnormalities in cancer. Philos Trans R Soc Lond B Biol Sci 369, https:\/\/doi.org\/10.1098\/rstb.2013.0467 (2014).","journal-title":"Philosophical Transactions of the Royal Society B: Biological Sciences"},{"key":"43774_CR2","doi-asserted-by":"publisher","first-page":"a015784","DOI":"10.1101\/cshperspect.a015784","volume":"7","author":"S Reber","year":"2015","unstructured":"Reber, S. & Hyman, A. A. Emergent properties of the metaphase spindle. Cold Spring Harb Perspect Biol 7, a015784, https:\/\/doi.org\/10.1101\/cshperspect.a015784 (2015).","journal-title":"Cold Spring Harb Perspect Biol"},{"key":"43774_CR3","doi-asserted-by":"publisher","first-page":"1159","DOI":"10.1016\/0092-8674(95)90142-6","volume":"83","author":"A Blangy","year":"1995","unstructured":"Blangy, A. et al. Phosphorylation by p34cdc2 regulates spindle association of human Eg5, a kinesin-related motor essential for bipolar spindle formation in vivo. Cell 83, 1159\u20131169 (1995).","journal-title":"Cell"},{"key":"43774_CR4","doi-asserted-by":"publisher","first-page":"1019","DOI":"10.1016\/0092-8674(90)90350-N","volume":"60","author":"AP Enos","year":"1990","unstructured":"Enos, A. P. & Morris, N. R. Mutation of a gene that encodes a kinesin-like protein blocks nuclear division in A. nidulans. Cell 60, 1019\u20131027 (1990).","journal-title":"Cell"},{"key":"43774_CR5","doi-asserted-by":"publisher","first-page":"563","DOI":"10.1038\/347563a0","volume":"347","author":"I Hagan","year":"1990","unstructured":"Hagan, I. & Yanagida, M. Novel potential mitotic motor protein encoded by the fission yeast cut7\n                           + gene. 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A., He, L., Loo, K. K. & Saunders, W. S. Two Saccharomyces cerevisiae kinesin-related gene products required for mitotic spindle assembly. J Cell Biol 118, 109\u2013120 (1992).","journal-title":"J Cell Biol"},{"key":"43774_CR9","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1083\/jcb.118.1.95","volume":"118","author":"DM Roof","year":"1992","unstructured":"Roof, D. M., Meluh, P. B. & Rose, M. D. Kinesin-related proteins required for assembly of the mitotic spindle. J Cell Biol 118, 95\u2013108 (1992).","journal-title":"J Cell Biol"},{"key":"43774_CR10","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1016\/j.yexcr.2014.12.015","volume":"334","author":"AL Yount","year":"2015","unstructured":"Yount, A. L., Zong, H. & Walczak, C. E. Regulatory mechanisms that control mitotic kinesins. 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Nat Commun 5, 5339, https:\/\/doi.org\/10.1038\/ncomms6339 (2014).","journal-title":"Nat Commun"},{"key":"43774_CR29","doi-asserted-by":"publisher","first-page":"1246","DOI":"10.4161\/cc.7.9.5808","volume":"7","author":"AS Rodriguez","year":"2008","unstructured":"Rodriguez, A. S. et al. Protein complexes at the microtubule organizing center regulate bipolar spindle assembly. Cell Cycle 7, 1246\u20131253, https:\/\/doi.org\/10.4161\/cc.7.9.5808 (2008).","journal-title":"Cell Cycle"},{"key":"43774_CR30","doi-asserted-by":"publisher","first-page":"765","DOI":"10.1038\/nrm2782","volume":"10","author":"KJ Verhey","year":"2009","unstructured":"Verhey, K. J. & Hammond, J. W. Traffic control: regulation of kinesin motors. 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