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Important mitotic events occur prior to NEBD, including condensation of replicated chromosomes and assembly of kinetochores to rapidly engage spindle microtubules. Here, we show that nuclear-enriched protein phosphatase 4 (PP4) ensures robust assembly of the microtubule-coupling outer kinetochore prior to NEBD. In the absence of PP4, chromosomes exhibit extended monopolar orientation after NEBD and subsequently mis-segregate. A secondary consequence of diminished outer kinetochore assembly is defective sister chromatid resolution. After NEBD, a cytoplasmic activity compensates for PP4 loss, leading to outer kinetochore assembly and recovery of chromosomes from monopolar orientation to significant bi-orientation. The Ndc80-Ska microtubule-binding module of the outer kinetochore is required for this recovery. PP4 associates with the inner kinetochore protein CENP-C; however, disrupting the PP4\u2013CENP-C interaction does not perturb chromosome segregation. These results establish that PP4-dependent outer kinetochore assembly prior to NEBD is critical for timely and proper engagement of chromosomes with spindle microtubules.<\/jats:p>","DOI":"10.1083\/jcb.202208154","type":"journal-article","created":{"date-parts":[[2023,1,6]],"date-time":"2023-01-06T02:21:14Z","timestamp":1672971674000},"update-policy":"https:\/\/doi.org\/10.1083\/jcb.crossmarkpolicy","source":"Crossref","is-referenced-by-count":6,"title":["Nuclear-enriched protein phosphatase 4 ensures outer kinetochore assembly prior to nuclear dissolution"],"prefix":"10.1083","volume":"222","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1062-2498","authenticated-orcid":true,"given":"Helder","family":"Rocha","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/05qpmg879","id-type":"ROR","asserted-by":"publisher"}],"name":"Instituto de Investiga\u00e7\u00e3o e Inova\u00e7\u00e3o em Sa\u00fade \u2013 i3S, Universidade do Porto 1 , Porto, Portugal"},{"name":"Instituto de 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