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However, the molecular basis of nucleolar dynamics and their relationship to ribosome assembly remain incompletely understood. We present a platform for high-throughput FRAP (HiT-FRAP) and use it to screen hundreds of genes for their impact on dynamics of the nucleolar scaffold nucleophosmin (NPM1). We find that NPM1 dynamics and nucleolar morphology are sensitive to ribosome assembly state: accumulation of early pre-ribosomal intermediates slows NPM1 dynamics and compacts the condensate, while accumulation of abortive late precursors accelerates dynamics and disrupts condensate integrity. These opposing biophysical states correlate with the strength of NPM1\u2013pre-ribosome interactions. Importantly, mutations in the NPM1 intrinsically disordered region that alter pre-ribosome binding directly tune nucleolar dynamics. These results establish that ribosomal precursor assembly state determines nucleolar material properties through the strength of scaffold\u2013pre-ribosome interactions and introduce HiT-FRAP as a platform for interrogating condensate dynamics broadly.<\/jats:p>","DOI":"10.1083\/jcb.202605096","type":"journal-article","created":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T06:21:17Z","timestamp":1786688477000},"update-policy":"https:\/\/doi.org\/10.1083\/jcb.crossmarkpolicy","source":"Crossref","is-referenced-by-count":1,"title":["Nucleolar dynamics are determined by the ordered assembly of the ribosome"],"prefix":"10.1083","volume":"225","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6806-4532","authenticated-orcid":true,"given":"Jessica","family":"Sheu-Gruttadauria","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/043mz5j54","id-type":"ROR","asserted-by":"publisher"}],"name":"University of California, San Francisco 1 Department of Cellular and Molecular Pharmacology, , San Francisco, CA,","place":["USA"]},{"id":[{"id":"https:\/\/ror.org\/043mz5j54","id-type":"ROR","asserted-by":"publisher"}],"name":"University of California, San Francisco 2 Department of Cell and Tissue Biology, , San Francisco, CA,","place":["USA"]},{"id":[{"id":"https:\/\/ror.org\/02yrq0923","id-type":"ROR","asserted-by":"publisher"}],"name":"Molecular Biology Program, Sloan Kettering Institute 4 , New York, NY,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4846-8812","authenticated-orcid":true,"given":"Xiaowei","family":"Yan","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/043mz5j54","id-type":"ROR","asserted-by":"publisher"}],"name":"University of California, San Francisco 1 Department of Cellular and Molecular Pharmacology, , San Francisco, CA,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6179-8613","authenticated-orcid":true,"given":"Nico","family":"Stuurman","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/043mz5j54","id-type":"ROR","asserted-by":"publisher"}],"name":"University of California, San Francisco 1 Department of Cellular and Molecular Pharmacology, , San Francisco, CA,","place":["USA"]},{"name":"Howard Hughes Medical Institute 3 , San Francisco, CA,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-0907-0455","authenticated-orcid":true,"given":"Riley O.","family":"Ogrean","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/02yrq0923","id-type":"ROR","asserted-by":"publisher"}],"name":"Molecular Biology Program, Sloan Kettering Institute 4 , New York, NY,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-0676-7690","authenticated-orcid":true,"given":"Sabrina","family":"Lin","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/02yrq0923","id-type":"ROR","asserted-by":"publisher"}],"name":"Molecular Biology Program, Sloan Kettering Institute 4 , New York, NY,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9965-9694","authenticated-orcid":true,"given":"Stephen N.","family":"Floor","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/043mz5j54","id-type":"ROR","asserted-by":"publisher"}],"name":"University of California, San Francisco 2 Department of Cell and Tissue Biology, , San Francisco, CA,","place":["USA"]},{"id":[{"id":"https:\/\/ror.org\/043mz5j54","id-type":"ROR","asserted-by":"publisher"}],"name":"Helen Diller Family Comprehensive Cancer Center, University of California, San Francisco 5 , San Francisco, CA,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3460-2758","authenticated-orcid":true,"given":"Ronald D.","family":"Vale","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/043mz5j54","id-type":"ROR","asserted-by":"publisher"}],"name":"University of California, San Francisco 1 Department of Cellular and Molecular Pharmacology, , San Francisco, CA,","place":["USA"]},{"name":"Howard Hughes Medical Institute 3 , San Francisco, CA,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"291","published-online":{"date-parts":[[2026,9,9]]},"reference":[{"key":"2026090909170885200_bib1","doi-asserted-by":"publisher","first-page":"196","DOI":"10.1038\/s41580-020-00326-6","article-title":"Biomolecular condensates at the nexus of cellular stress, protein aggregation disease and ageing","volume":"22","author":"Alberti","year":"2021","journal-title":"Nat. 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