{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,13]],"date-time":"2026-02-13T17:27:50Z","timestamp":1771003670972,"version":"3.50.1"},"reference-count":82,"publisher":"MDPI AG","issue":"2","license":[{"start":{"date-parts":[[2026,2,13]],"date-time":"2026-02-13T00:00:00Z","timestamp":1770940800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"JSPS KAKENHI","award":["19J21096"],"award-info":[{"award-number":["19J21096"]}]},{"name":"JSPS KAKENHI","award":["20K21259"],"award-info":[{"award-number":["20K21259"]}]},{"name":"JSPS KAKENHI","award":["21H02062"],"award-info":[{"award-number":["21H02062"]}]},{"name":"JSPS KAKENHI","award":["24K21801"],"award-info":[{"award-number":["24K21801"]}]},{"name":"JSPS KAKENHI","award":["21K06158"],"award-info":[{"award-number":["21K06158"]}]},{"name":"JST SPRING","award":["JPMJSP2117"],"award-info":[{"award-number":["JPMJSP2117"]}]},{"name":"the Public Interest Incorporated Foundation of the Hyogo Science and Technology Association and Terumo Life Science Foundation"},{"name":"the Junzo Tateno Foundation and the Hirao Taro Foundation of Konan Gakuen for Academic Research"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biomolecules"],"abstract":"<jats:p>Chemotherapeutic agents targeting ribosome biogenesis induce profound reorganization of nucleolar architecture, yet how the tumor suppressor p53 governs these structural responses remains unclear. Here, we show that loss of p53 leads to NF-\u03baB-dependent disappearance of nucleolar caps induced by doxorubicin (DOXO). Under these conditions, fibrillarin (FBL), which is normally confined to the nucleolus, relocates to the nucleoplasm and forms foci that partially associate with G-quadruplex (G4) structures, non-canonical nucleic acid secondary structures enriched at transcriptionally active genomic regions. To examine whether this redistribution is linked to transcriptional changes, we integrated publicly available transcriptomic datasets and identified genes that were upregulated in p53-deficient cells under DOXO treatment and downregulated upon FBL depletion. Given that casein kinase 2 alpha (CK2\u03b1) is a nuclear binding partner of FBL, we further analyzed CK2\u03b1-dependent gene programs. This analysis revealed that a fraction of FBL-responsive genes overlapped with CK2\u03b1-dependent signatures and were enriched for promoter-proximal G4 structures. Among candidate regulators, the G4-binding transcription factor MAZ emerged as a potential mediator linking nucleoplasmic FBL and CK2\u03b1 to G4-associated transcriptional regulation. Together, our findings identify a mechanism linking loss of p53 to G4-associated transcriptional reprogramming through nucleolar architectural disruption mediated by an FBL\u2013CK2\u03b1\u2013MAZ axis during DOXO treatment.<\/jats:p>","DOI":"10.3390\/biom16020296","type":"journal-article","created":{"date-parts":[[2026,2,13]],"date-time":"2026-02-13T16:09:32Z","timestamp":1770998972000},"page":"296","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Loss of p53 Provokes NF-\u03baB-Dependent Disruption of Nucleolar Cap and Nucleoplasmic Redistribution of Fibrillarin During Nucleolar Stress"],"prefix":"10.3390","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6278-7617","authenticated-orcid":false,"given":"Takeru","family":"Torii","sequence":"first","affiliation":[{"name":"Faculty of Frontiers of Innovative Research in Science and Technology (FIRST), Konan University, Kobe 650-0047, Hy\u014dgo, Japan"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mako","family":"Sumida","sequence":"additional","affiliation":[{"name":"Faculty of Frontiers of Innovative Research in Science and Technology (FIRST), Konan University, Kobe 650-0047, Hy\u014dgo, Japan"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Atsushi","family":"Kobayashi","sequence":"additional","affiliation":[{"name":"Faculty of Frontiers of Innovative Research in Science and Technology (FIRST), Konan University, Kobe 650-0047, Hy\u014dgo, Japan"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2827-4199","authenticated-orcid":false,"given":"Toshiyuki","family":"Goto","sequence":"additional","affiliation":[{"name":"Faculty of Frontiers of Innovative Research in Science and Technology (FIRST), Konan University, Kobe 650-0047, Hy\u014dgo, Japan"},{"name":"Graduate School of Science, Technology and Innovation, Kobe University, Kobe 650-0047, Hy\u014dgo, Japan"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ryosuke","family":"Suzuki","sequence":"additional","affiliation":[{"name":"Faculty of Frontiers of Innovative Research in Science and Technology (FIRST), Konan University, Kobe 650-0047, Hy\u014dgo, Japan"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2513-5222","authenticated-orcid":false,"given":"Shin","family":"Kuwamoto","sequence":"additional","affiliation":[{"name":"Faculty of Frontiers of Innovative Research in Science and Technology (FIRST), Konan University, Kobe 650-0047, Hy\u014dgo, Japan"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Wataru","family":"Nakajima","sequence":"additional","affiliation":[{"name":"Department of Molecular Oncology, Institute for Advanced Medical Sciences, Nippon Medical School, 1-1-5 Sendagi, Bunkyo-ku 113-8602, Tokyo, 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Japan"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4430-579X","authenticated-orcid":false,"given":"Masayuki","family":"Tera","sequence":"additional","affiliation":[{"name":"Department of Biotechnology and Life Science, Tokyo University of Agriculture and Technology, 2-24-16 Naka-cho, Koganei 184-8588, Tokyo, Japan"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6373-2220","authenticated-orcid":false,"given":"Nobuyuki","family":"Tanaka","sequence":"additional","affiliation":[{"name":"Department of Molecular Oncology, Institute for Advanced Medical Sciences, Nippon Medical School, 1-1-5 Sendagi, Bunkyo-ku 113-8602, Tokyo, Japan"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2604-9158","authenticated-orcid":false,"given":"Hiroaki","family":"Hirata","sequence":"additional","affiliation":[{"name":"Department of Life Science and Biotechnology, Kanazawa Institute of Technology, Hakusan 924-0838, Ishikawa, Japan"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6161-9817","authenticated-orcid":false,"given":"Hisae","family":"Tateishi-Karimata","sequence":"additional","affiliation":[{"name":"Faculty of Frontiers of Innovative Research in Science and Technology (FIRST), Konan University, Kobe 650-0047, Hy\u014dgo, Japan"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2965-1793","authenticated-orcid":false,"given":"Takahito","family":"Nishikata","sequence":"additional","affiliation":[{"name":"Faculty of Frontiers of Innovative Research in Science and Technology (FIRST), Konan University, Kobe 650-0047, Hy\u014dgo, Japan"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Miwako Kato","family":"Homma","sequence":"additional","affiliation":[{"name":"Department of Biomolecular Sciences, Fukushima Medical University School of 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