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These mutations occur in the terminal exon of the NPM1 gene, affecting the C-terminal DNA-binding domain of the protein and causing its delocalization to the cytoplasm\u2014a hallmark of NPM1-mutated AML. NPM1 shuttling to the nucleoplasm is tightly regulated by posttranslational modifications, such as phosphorylation of Ser254, Ser260, and Tyr271 of the DNA-binding domain. However, the structural mechanisms underlying this process remain unclear. In this work, we show that Ser-to-Asp (S254D\u2013S260D) and Tyr-to-pCMF (para-carboxymethyl phenylalanine) (Y271pCMF) phosphomimetic mutations induce significant structural and dynamical rearrangements, as well as drastic modifications in electrostatic surface potential. These changes compromise recognition of a G-quadruplex sequence from the c-MYC promoter by reducing DNA-binding affinity, reshape histone capturing dynamics, and fade charge segregation in the histone-binding domain. Combination of such substitutions in a triple phosphomimetic variant (S254D\u2013S260D\u2013Y271pCMF) further destabilizes the domain\u2019s structure and triggers protein aggregation. Altogether, these findings suggest that phosphorylation of Ser254, Ser260, and Tyr271 of the C-end DNA-binding domain weakens both DNA affinity and charge block-driven liquid\u2013liquid phase separation, offering a molecular explanation for the delocalization of NPM1 outside of the nucleolus.<\/jats:p>","DOI":"10.1093\/nar\/gkag165","type":"journal-article","created":{"date-parts":[[2026,2,14]],"date-time":"2026-02-14T12:39:05Z","timestamp":1771072745000},"source":"Crossref","is-referenced-by-count":1,"title":["Multisite phosphorylation of the AML-linked C-terminal of nucleophosmin (NPM1) orchestrates protein stability, DNA binding and charge block-driven phase separation"],"prefix":"10.1093","volume":"54","author":[{"ORCID":"https:\/\/orcid.org\/0009-0002-2454-4056","authenticated-orcid":false,"given":"Pablo","family":"Rivero-Garc\u00eda","sequence":"first","affiliation":[{"name":"Institute for Chemical Research\u2014cicCartuja, University of Seville\u2014CSIC , 41092 Seville 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OX11 0DE Didcot ,","place":["United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0122-7859","authenticated-orcid":false,"given":"Elisa","family":"Frezza\u00a0","sequence":"additional","affiliation":[{"name":"Universit\u00e9 Paris Cit\u00e9, CiTCoM, CNRS , F-75006 Paris ,","place":["France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5702-4538","authenticated-orcid":false,"given":"Adri\u00e1n","family":"Vel\u00e1zquez-Campoy","sequence":"additional","affiliation":[{"name":"Institute for Biocomputation and Physics of Complex Systems (BIFI), and Department of Biochemistry and Molecular and Cell Biology, University of Zaragoza , 50018 Zaragoza ,","place":["Spain"]},{"name":"Instituto de Investigaci\u00f3n Sanitaria Arag\u00f3n (IIS Arag\u00f3n) , 50009 Zaragoza ,","place":["Spain"]},{"name":"Centro de Investigaci\u00f3n Biom\u00e9dica en Red en el \u00c1rea Tem\u00e1tica de Enfermedades Hep\u00e1ticas y Digestivas (CIBERehd) , 28029 Madrid ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7616-6515","authenticated-orcid":false,"given":"Sof\u00eda","family":"D\u00edaz-Moreno","sequence":"additional","affiliation":[{"name":"Diamond Light Source\u2014Harwell Science & Innovation Campus , OX11 0DE Didcot ,","place":["United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1187-5737","authenticated-orcid":false,"given":"Miguel\u00a0A\u00a0De","family":"la\u00a0Rosa","sequence":"additional","affiliation":[{"name":"Institute for Chemical Research\u2014cicCartuja, University of Seville\u2014CSIC , 41092 Seville 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