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Here, we show that N-WASP can trigger the formation of nuclear N-WASP\/F-actin nodules. While a cancer hotspot mutant of N-WASP lacking the VCA domain (V418fs) had a dominant negative function on nuclear F-actin, an even shorter truncation mutant found in melanoma (R128*) strongly promoted nuclear actin polymerization. Nuclear localization of N-WASP was not regulated by the cell cycle and increasing nuclear F-actin formation by N-WASP had no obvious influence on replication. However, nuclear N-WASP\/F-actin nodules colocalized partially with RNA Pol II clusters. N-WASP-dependent actin polymerization promoted the maturation of RNA Pol II clusters, with the short truncation mutant R128* unexpectedly showing the strongest effect. Nuclear N-WASP nodules including V418fs colocalized with WIP and cortactin. Importantly, cortactin binding was essential but not sufficient for F-actin formation, while WIP binding was required for actin polymerization by R128*. These data reveal a cortactin-dependent role for N-WASP in the regulation of nuclear F-actin and indicate contrasting nuclear effects for N-WASP mutants found in cancer.<\/jats:p>","DOI":"10.3390\/cells14010059","type":"journal-article","created":{"date-parts":[[2025,1,6]],"date-time":"2025-01-06T09:17:02Z","timestamp":1736155022000},"page":"59","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":4,"title":["Nuclear N-WASP Induces Actin Polymerization in the Nucleus with Cortactin as an Essential Factor"],"prefix":"10.3390","volume":"14","author":[{"given":"Xin","family":"Jiang","sequence":"first","affiliation":[{"name":"Biotech Research and Innovation Center (BRIC), University of Copenhagen, Ole Maal\u00f8es Vej5, 2200 Copenhagen, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Purusottam","family":"Mohapatra","sequence":"additional","affiliation":[{"name":"Biotech Research and Innovation Center (BRIC), University of Copenhagen, Ole Maal\u00f8es Vej5, 2200 Copenhagen, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria","family":"Rossing","sequence":"additional","affiliation":[{"name":"Center for Genomic Medicine, Rigshospitalet, University of Copenhagen, Blegdamsvej 9, 2100 Copenhagen, Denmark"},{"name":"Department of Clinical Medicine, University of Copenhagen, 2200 Copenhagen, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-9232-4604","authenticated-orcid":false,"given":"Wenqian","family":"Zheng","sequence":"additional","affiliation":[{"name":"Biotech Research and Innovation Center (BRIC), University of Copenhagen, Ole Maal\u00f8es Vej5, 2200 Copenhagen, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Olga","family":"Zbodakova","sequence":"additional","affiliation":[{"name":"Biotech Research and Innovation Center (BRIC), University of Copenhagen, Ole Maal\u00f8es Vej5, 2200 Copenhagen, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jayashree Vijay","family":"Thatte","sequence":"additional","affiliation":[{"name":"Biotech Research and Innovation Center (BRIC), University of Copenhagen, Ole Maal\u00f8es Vej5, 2200 Copenhagen, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0811-9420","authenticated-orcid":false,"given":"Claus Storgaard","family":"S\u00f8rensen","sequence":"additional","affiliation":[{"name":"Biotech Research and Innovation Center (BRIC), University of Copenhagen, Ole Maal\u00f8es Vej5, 2200 Copenhagen, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thu Han","family":"Le Phan","sequence":"additional","affiliation":[{"name":"Biotech Research and Innovation Center (BRIC), University of Copenhagen, Ole Maal\u00f8es Vej5, 2200 Copenhagen, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9342-1634","authenticated-orcid":false,"given":"Cord","family":"Brakebusch","sequence":"additional","affiliation":[{"name":"Biotech Research and Innovation Center (BRIC), University of Copenhagen, Ole Maal\u00f8es Vej5, 2200 Copenhagen, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,1,6]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1038\/nrm2069","article-title":"The WASP-WAVE protein network: Connecting the membrane to the cytoskeleton","volume":"8","author":"Takenawa","year":"2007","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"850","DOI":"10.1093\/embo-reports\/kve197","article-title":"Actin pedestal formation by enteropathogenic Escherichia coli and intracellular motility of Shigella flexneri are abolished in N-WASP-defective cells","volume":"2","author":"Lommel","year":"2001","journal-title":"EMBO Rep."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"897","DOI":"10.1038\/ncb1001-897","article-title":"N-WASP deficiency reveals distinct pathways for cell surface projections and microbial actin-based motility","volume":"3","author":"Snapper","year":"2001","journal-title":"Nat. Cell Biol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"42515","DOI":"10.1074\/jbc.M302177200","article-title":"Translocation of N-WASP by nuclear localization and export signals into the nucleus modulates expression of HSP90","volume":"278","author":"Suetsugu","year":"2003","journal-title":"J. Biol. Chem."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"756","DOI":"10.1038\/ncb1433","article-title":"Regulation of RNA-polymerase-II-dependent transcription by N-WASP and its nuclear-binding partners","volume":"8","author":"Wu","year":"2006","journal-title":"Nat. Cell Biol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1420","DOI":"10.1038\/s41467-018-03704-z","article-title":"Epigenetic control of IL-23 expression in keratinocytes is important for chronic skin inflammation","volume":"9","author":"Li","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"2167","DOI":"10.1158\/0008-5472.CAN-18-1253","article-title":"Negative Regulation of p53-Induced Senescence by N-WASP Is Crucial for DMBA\/TPA-Induced Skin Tumor Formation","volume":"79","author":"Li","year":"2019","journal-title":"Cancer Res."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"3743","DOI":"10.1038\/s41467-022-31415-z","article-title":"WASp modulates RPA function on single-stranded DNA in response to replication stress and DNA damage","volume":"13","author":"Han","year":"2022","journal-title":"Nat. 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Immunol."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Flores, L.R., Keeling, M.C., Zhang, X., Sliogeryte, K., and Gavara, N. (2019). Lifeact-GFP alters F-actin organization, cellular morphology and biophysical behaviour. Sci. Rep., 9.","DOI":"10.1038\/s41598-019-40092-w"},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Gunasekaran, S., Miyagawa, Y., and Miyamoto, K. (2022). Actin nucleoskeleton in embryonic development and cellular differentiation. Curr. Opin. Cell Biol., 76.","DOI":"10.1016\/j.ceb.2022.102100"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"462","DOI":"10.1016\/j.tcb.2019.02.010","article-title":"Nuclear Actin and Actin-Binding Proteins in DNA Repair","volume":"29","author":"Hurst","year":"2019","journal-title":"Trends Cell Biol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"jcs261630","DOI":"10.1242\/jcs.261630","article-title":"Nuclear actin dynamics and functions at a glance","volume":"137","author":"Ulferts","year":"2024","journal-title":"J. Cell Sci."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1389","DOI":"10.1038\/ncb3641","article-title":"A transient pool of nuclear F-actin at mitotic exit controls chromatin organization","volume":"19","author":"Baarlink","year":"2017","journal-title":"Nat. Cell Biol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"354","DOI":"10.18632\/oncoscience.487","article-title":"Senescence regulation by nuclear N-WASP: A role in cancer?","volume":"6","author":"Li","year":"2019","journal-title":"Oncoscience"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"128","DOI":"10.1242\/jcs.053835","article-title":"N-WASP is a novel regulator of hair-follicle cycling that controls antiproliferative TGF{beta} pathways","volume":"123","author":"Lefever","year":"2010","journal-title":"J. Cell Sci."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1007\/978-1-61779-442-1_11","article-title":"Rho GTPase knockout induction in primary keratinocytes from adult mice","volume":"827","author":"Pedersen","year":"2012","journal-title":"Rho GTPases Methods Protoc."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"2281","DOI":"10.1038\/nprot.2013.143","article-title":"Genome engineering using the CRISPR-Cas9 system","volume":"8","author":"Ran","year":"2013","journal-title":"Nat. Protoc."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Carpenter, A.E., Jones, T.R., Lamprecht, M.R., Clarke, C., Kang, I.H., Friman, O., Guertin, D.A., Chang, J.H., Lindquist, R.A., and Moffat, J. (2006). CellProfiler: Image analysis software for identifying and quantifying cell phenotypes. Genome Biol., 7.","DOI":"10.1186\/gb-2006-7-10-r100"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"9565","DOI":"10.1074\/jbc.M310739200","article-title":"Focal adhesion kinase regulation of N-WASP subcellular localization and function","volume":"279","author":"Wu","year":"2004","journal-title":"J. Biol. Chem."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1016\/j.bbrc.2009.04.050","article-title":"Activating mutations of N-WASP alter Shigella pathogenesis","volume":"384","author":"Adamovich","year":"2009","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"5326","DOI":"10.1002\/j.1460-2075.1996.tb00917.x","article-title":"N-WASP, a novel actin-depolymerizing protein, regulates the cortical cytoskeletal rearrangement in a PIP2-dependent manner downstream of tyrosine kinases","volume":"15","author":"Miki","year":"1996","journal-title":"EMBO J."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"C723","DOI":"10.1152\/ajpcell.00462.2010","article-title":"A practical guide to evaluating colocalization in biological microscopy","volume":"300","author":"Dunn","year":"2011","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"3212","DOI":"10.15252\/embj.201796585","article-title":"Initiation of DNA replication requires actin dynamics and formin activity","volume":"36","author":"Parisis","year":"2017","journal-title":"EMBO J."},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"He, X., and Brakebusch, C. (2024). Regulation of Precise DNA Repair by Nuclear Actin Polymerization: A Chance for Improving Gene Therapy?. Cells, 13.","DOI":"10.3390\/cells13131093"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"eaay6515","DOI":"10.1126\/sciadv.aay6515","article-title":"Nuclear actin regulates inducible transcription by enhancing RNA polymerase II clustering","volume":"6","author":"Wei","year":"2020","journal-title":"Sci. Adv."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"713","DOI":"10.1189\/jlb.2RU0314-162R","article-title":"WIP: More than a WASp-interacting protein","volume":"96","author":"Fried","year":"2014","journal-title":"J. Leukoc. Biol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"256","DOI":"10.1080\/21541248.2017.1405773","article-title":"Cortactin function in invadopodia","volume":"11","author":"Jeannot","year":"2020","journal-title":"Small GTPases"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"5269","DOI":"10.1128\/MCB.24.12.5269-5280.2004","article-title":"Erk\/Src phosphorylation of cortactin acts as a switch on-switch off mechanism that controls its ability to activate N-WASP","volume":"24","author":"Ho","year":"2004","journal-title":"Mol. Cell. Biol."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Belyy, A., Merino, F., Sitsel, O., and Raunser, S. (2020). Structure of the Lifeact-F-actin complex. PLoS Biol., 18.","DOI":"10.2210\/pdb7ad9\/pdb"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"484","DOI":"10.1038\/35074551","article-title":"WIP regulates N-WASP-mediated actin polymerization and filopodium formation","volume":"3","author":"Rohatgi","year":"2001","journal-title":"Nat. Cell Biol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1242\/jcs.01586","article-title":"Cortactin regulates cell migration through activation of N-WASP","volume":"118","author":"Kowalski","year":"2005","journal-title":"J. Cell Sci."},{"key":"ref_35","first-page":"669","article-title":"Essential role of neural Wiskott-Aldrich syndrome protein in podosome formation and degradation of extracellular matrix in src-transformed fibroblasts","volume":"62","author":"Mizutani","year":"2002","journal-title":"Cancer Res."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"707","DOI":"10.1146\/annurev.biochem.77.060407.135452","article-title":"Physical mechanisms of signal integration by WASP family proteins","volume":"79","author":"Padrick","year":"2010","journal-title":"Annu. Rev. Biochem."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"e00884","DOI":"10.7554\/eLife.00884","article-title":"Mechanism of synergistic activation of Arp2\/3 complex by cortactin and N-WASP","volume":"2","author":"Helgeson","year":"2013","journal-title":"eLife"},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Latario, C.J., Pickrell, L.E., and Higgs, H.N. (2020). Lysine acetylation of cytoskeletal proteins: Emergence of an actin code. J. Cell Biol., 219.","DOI":"10.1083\/jcb.202006151"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1016\/j.tcb.2017.10.009","article-title":"Cortactin: Cell Functions of A Multifaceted Actin-Binding Protein","volume":"28","author":"Schnoor","year":"2018","journal-title":"Trends Cell Biol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"638","DOI":"10.1002\/dvg.20544","article-title":"Generation of cortactin floxed mice and cellular analysis of motility in fibroblasts","volume":"47","author":"Tanaka","year":"2009","journal-title":"Genesis"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"848","DOI":"10.1016\/j.yexcr.2009.11.018","article-title":"Cortactin is implicated in murine zygotic development","volume":"316","author":"Yu","year":"2010","journal-title":"Exp. Cell Res."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1721","DOI":"10.1084\/jem.20101920","article-title":"Cortactin deficiency is associated with reduced neutrophil recruitment but increased vascular permeability in vivo","volume":"208","author":"Schnoor","year":"2011","journal-title":"J. Exp. Med."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1016\/bs.ircmb.2020.05.006","article-title":"Actin regulators in cancer progression and metastases: From structure and function to cytoskeletal dynamics","volume":"356","author":"Biber","year":"2020","journal-title":"Int. Rev. Cell Mol. Biol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1016\/S0304-3940(03)00506-8","article-title":"Possible involvement of Wiskott-Aldrich syndrome protein family in aberrant neuronal sprouting in Alzheimer\u2019s disease","volume":"346","author":"Kitamura","year":"2003","journal-title":"Neurosci. Lett."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"168","DOI":"10.1016\/j.brainres.2008.07.101","article-title":"Overexpression of N-WASP in the brain of human epilepsy","volume":"1233","author":"Xiao","year":"2008","journal-title":"Brain Res."},{"key":"ref_46","first-page":"1501","article-title":"Mycolactone activation of Wiskott-Aldrich syndrome proteins underpins Buruli ulcer formation","volume":"23","author":"Thoulouze","year":"2013","journal-title":"J. Clin. Investig."}],"container-title":["Cells"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4409\/14\/1\/59\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,8]],"date-time":"2025-10-08T10:23:40Z","timestamp":1759919020000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4409\/14\/1\/59"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,1,6]]},"references-count":46,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,1]]}},"alternative-id":["cells14010059"],"URL":"https:\/\/doi.org\/10.3390\/cells14010059","relation":{},"ISSN":["2073-4409"],"issn-type":[{"value":"2073-4409","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,1,6]]}}}