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PKM2 inhibitor shikonin markedly attenuates KRI1 phase separation and its interaction with NPM1, disrupts nucleolar structure and ribosome biogenesis, thereby inhibiting HCC progression. This study uncovers the crucial role of KRI1 in HCC progression and provides theoretical and experimental evidence for the development of nucleolar protein-targeted therapeutic strategies for HCC treatment.<\/jats:p>","DOI":"10.1073\/pnas.2535921123","type":"journal-article","created":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T19:06:16Z","timestamp":1785783976000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":1,"title":["Co-phase separation of KRI1 and NPM1 sustains nucleolar integrity and ribosome biogenesis to fuel liver cancer progression"],"prefix":"10.1073","volume":"123","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7039-0284","authenticated-orcid":false,"given":"Yu","family":"Lu","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/01y0j0j86","id-type":"ROR","asserted-by":"publisher"}],"name":"School of Life Sciences and Technology, Northwestern Polytechnical University","place":["Xi\u2019an, 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