{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,13]],"date-time":"2026-04-13T12:50:08Z","timestamp":1776084608513,"version":"3.50.1"},"reference-count":48,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2018,7,5]],"date-time":"2018-07-05T00:00:00Z","timestamp":1530748800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2018,7,5]],"date-time":"2018-07-05T00:00:00Z","timestamp":1530748800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Hypoxia is a common feature of solid tumours that promotes invasion and metastatic dissemination. Invadopodia are actin-rich membrane protrusions that direct extracellular matrix proteolysis and facilitate tumour cell invasion. Here, we show that CSRP2, an invadopodial actin bundling protein, is upregulated by hypoxia in various breast cancer cell lines, as well as in pre-clinical and clinical breast tumour specimens. We functionally characterized two hypoxia responsive elements within the proximal promoter of CSRP2 gene which are targeted by hypoxia-inducible factor-1 (HIF-1) and required for promoter transactivation in response to hypoxia. Remarkably, CSRP2 knockdown significantly inhibits hypoxia-stimulated invadopodium formation, ECM degradation and invasion in MDA-MB-231 cells, while CSRP2 forced expression was sufficient to enhance the invasive capacity of HIF-1\u03b1-depleted cells under hypoxia. In MCF-7 cells, CSRP2 upregulation was required for hypoxia-induced formation of invadopodium precursors that were unable to promote ECM degradation. Collectively, our data support that CSRP2 is a novel and direct cytoskeletal target of HIF-1 which facilitates hypoxia-induced breast cancer cell invasion by promoting invadopodia formation.<\/jats:p>","DOI":"10.1038\/s41598-018-28637-x","type":"journal-article","created":{"date-parts":[[2018,6,29]],"date-time":"2018-06-29T11:32:20Z","timestamp":1530271940000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":74,"title":["Hypoxia promotes breast cancer cell invasion through HIF-1\u03b1-mediated up-regulation of the invadopodial actin bundling protein CSRP2"],"prefix":"10.1038","volume":"8","author":[{"given":"C\u00e9line","family":"Hoffmann","sequence":"first","affiliation":[]},{"given":"Xianqing","family":"Mao","sequence":"additional","affiliation":[]},{"given":"Joshua","family":"Brown-Clay","sequence":"additional","affiliation":[]},{"given":"Flora","family":"Moreau","sequence":"additional","affiliation":[]},{"given":"Antoun","family":"Al Absi","sequence":"additional","affiliation":[]},{"given":"Hannah","family":"Wurzer","sequence":"additional","affiliation":[]},{"given":"Barbara","family":"Sousa","sequence":"additional","affiliation":[]},{"given":"Fernando","family":"Schmitt","sequence":"additional","affiliation":[]},{"given":"Guy","family":"Berchem","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9763-0943","authenticated-orcid":false,"given":"Bassam","family":"Janji","sequence":"additional","affiliation":[]},{"given":"Cl\u00e9ment","family":"Thomas","sequence":"additional","affiliation":[]}],"member":"297","published-online":{"date-parts":[[2018,7,5]]},"reference":[{"key":"28637_CR1","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1038\/nrc.2016.25","volume":"16","author":"PS Steeg","year":"2016","unstructured":"Steeg, P. S. Targeting metastasis. Nat Rev Cancer 16, 201\u2013218, https:\/\/doi.org\/10.1038\/nrc.2016.25 (2016).","journal-title":"Nat Rev Cancer"},{"key":"28637_CR2","doi-asserted-by":"publisher","first-page":"430","DOI":"10.1038\/nrc3726","volume":"14","author":"DM Gilkes","year":"2014","unstructured":"Gilkes, D. M., Semenza, G. L. & Wirtz, D. Hypoxia and the extracellular matrix: drivers of tumour metastasis. Nat Rev Cancer 14, 430\u2013439, https:\/\/doi.org\/10.1038\/nrc3726 (2014).","journal-title":"Nat Rev Cancer"},{"key":"28637_CR3","doi-asserted-by":"publisher","first-page":"382","DOI":"10.1016\/j.bbamcr.2015.05.036","volume":"1863","author":"GL Semenza","year":"2016","unstructured":"Semenza, G. L. The hypoxic tumor microenvironment: A driving force for breast cancer progression. Biochim Biophys Acta 1863, 382\u2013391, https:\/\/doi.org\/10.1016\/j.bbamcr.2015.05.036 (2016).","journal-title":"Biochim Biophys Acta"},{"key":"28637_CR4","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1126\/science.aaf4405","volume":"352","author":"EB Rankin","year":"2016","unstructured":"Rankin, E. B. & Giaccia, A. J. Hypoxic control of metastasis. Science 352, 175\u2013180, https:\/\/doi.org\/10.1126\/science.aaf4405 (2016).","journal-title":"Science"},{"key":"28637_CR5","doi-asserted-by":"publisher","first-page":"226","DOI":"10.4161\/cam.28346","volume":"8","author":"CM Gould","year":"2014","unstructured":"Gould, C. M. & Courtneidge, S. A. Regulation of invadopodia by the tumor microenvironment. Cell Adh Migr 8, 226\u2013235 (2014).","journal-title":"Cell Adh Migr"},{"key":"28637_CR6","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1016\/j.tcb.2007.01.002","volume":"17","author":"S Linder","year":"2007","unstructured":"Linder, S. The matrix corroded: podosomes and invadopodia in extracellular matrix degradation. Trends in cell biology 17, 107\u2013117, https:\/\/doi.org\/10.1016\/j.tcb.2007.01.002 (2007).","journal-title":"Trends in cell biology"},{"key":"28637_CR7","doi-asserted-by":"publisher","unstructured":"Eddy, R. J., Weidmann, M. D., Sharma, V. P. & Condeelis, J. S. Tumor Cell Invadopodia: Invasive Protrusions that Orchestrate Metastasis. Trends in cell biology, https:\/\/doi.org\/10.1016\/j.tcb.2017.03.003 (2017).","DOI":"10.1016\/j.tcb.2017.03.003"},{"key":"28637_CR8","doi-asserted-by":"publisher","first-page":"1558","DOI":"10.1016\/j.celrep.2014.07.050","volume":"8","author":"HS Leong","year":"2014","unstructured":"Leong, H. S. et al. Invadopodia are required for cancer cell extravasation and are a therapeutic target for metastasis. Cell reports 8, 1558\u20131570, https:\/\/doi.org\/10.1016\/j.celrep.2014.07.050 (2014).","journal-title":"Cell reports"},{"key":"28637_CR9","doi-asserted-by":"publisher","first-page":"733","DOI":"10.1586\/14737140.2015.1058711","volume":"15","author":"K Stoletov","year":"2015","unstructured":"Stoletov, K. & Lewis, J. D. Invadopodia: a new therapeutic target to block cancer metastasis. Expert Rev Anticancer Ther 15, 733\u2013735, https:\/\/doi.org\/10.1586\/14737140.2015.1058711 (2015).","journal-title":"Expert Rev Anticancer Ther"},{"key":"28637_CR10","doi-asserted-by":"publisher","first-page":"1142","DOI":"10.3892\/mmr.2014.1965","volume":"9","author":"N Tokui","year":"2014","unstructured":"Tokui, N. et al. Extravasation during bladder cancer metastasis requires cortactinmediated invadopodia formation. Mol Med Rep 9, 1142\u20131146, https:\/\/doi.org\/10.3892\/mmr.2014.1965 (2014).","journal-title":"Mol Med Rep"},{"key":"28637_CR11","doi-asserted-by":"publisher","first-page":"724","DOI":"10.1242\/jcs.092726","volume":"125","author":"B Gligorijevic","year":"2012","unstructured":"Gligorijevic, B. et al. N-WASP-mediated invadopodium formation is involved in intravasation and lung metastasis of mammary tumors. Journal of cell science 125, 724\u2013734, https:\/\/doi.org\/10.1242\/jcs.092726 (2012).","journal-title":"Journal of cell science"},{"key":"28637_CR12","doi-asserted-by":"publisher","first-page":"e1001995","DOI":"10.1371\/journal.pbio.1001995","volume":"12","author":"B Gligorijevic","year":"2014","unstructured":"Gligorijevic, B., Bergman, A. & Condeelis, J. Multiparametric classification links tumor microenvironments with tumor cell phenotype. Plos Biol 12, e1001995, https:\/\/doi.org\/10.1371\/journal.pbio.1001995 (2014).","journal-title":"Plos Biol"},{"key":"28637_CR13","doi-asserted-by":"publisher","first-page":"e0121003","DOI":"10.1371\/journal.pone.0121003","volume":"10","author":"B Blouw","year":"2015","unstructured":"Blouw, B. et al. The invadopodia scaffold protein Tks5 is required for the growth of human breast cancer cells in vitro and in vivo. Plos One 10, e0121003, https:\/\/doi.org\/10.1371\/journal.pone.0121003 (2015).","journal-title":"Plos One"},{"key":"28637_CR14","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1016\/S0092-8674(03)00513-0","volume":"114","author":"KB Hotary","year":"2003","unstructured":"Hotary, K. B. et al. Membrane type I matrix metalloproteinase usurps tumor growth control imposed by the three-dimensional extracellular matrix. Cell 114, 33\u201345 (2003).","journal-title":"Cell"},{"key":"28637_CR15","doi-asserted-by":"publisher","first-page":"3037","DOI":"10.1242\/jcs.052704","volume":"122","author":"C Albiges-Rizo","year":"2009","unstructured":"Albiges-Rizo, C., Destaing, O., Fourcade, B., Planus, E. & Block, M. R. Actin machinery and mechanosensitivity in invadopodia, podosomes and focal adhesions. Journal of cell science 122, 3037\u20133049, https:\/\/doi.org\/10.1242\/jcs.052704 (2009).","journal-title":"Journal of cell science"},{"key":"28637_CR16","doi-asserted-by":"publisher","first-page":"541","DOI":"10.1083\/jcb.200909113","volume":"189","author":"M Schoumacher","year":"2010","unstructured":"Schoumacher, M., Goldman, R. D., Louvard, D. & Vignjevic, D. M. Actin, microtubules, and vimentin intermediate filaments cooperate for elongation of invadopodia. The Journal of cell biology 189, 541\u2013556, https:\/\/doi.org\/10.1083\/jcb.200909113 (2010).","journal-title":"The Journal of cell biology"},{"key":"28637_CR17","doi-asserted-by":"publisher","first-page":"642","DOI":"10.1016\/j.bbamcr.2006.07.001","volume":"1773","author":"H Yamaguchi","year":"2007","unstructured":"Yamaguchi, H. & Condeelis, J. Regulation of the actin cytoskeleton in cancer cell migration and invasion. Biochim Biophys Acta 1773, 642\u2013652, https:\/\/doi.org\/10.1016\/j.bbamcr.2006.07.001 (2007).","journal-title":"Biochim Biophys Acta"},{"key":"28637_CR18","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1146\/annurev-cellbio-092910-154216","volume":"27","author":"S Linder","year":"2011","unstructured":"Linder, S., Wiesner, C. & Himmel, M. Degrading devices: invadosomes in proteolytic cell invasion. Annual review of cell and developmental biology 27, 185\u2013211, https:\/\/doi.org\/10.1146\/annurev-cellbio-092910-154216 (2011).","journal-title":"Annual review of cell and developmental biology"},{"key":"28637_CR19","doi-asserted-by":"publisher","first-page":"13688","DOI":"10.18632\/oncotarget.7327","volume":"7","author":"C Hoffmann","year":"2016","unstructured":"Hoffmann, C. et al. CRP2, a new invadopodia actin bundling factor critically promotes breast cancer cell invasion and metastasis. Oncotarget 7, 13688\u201313705, https:\/\/doi.org\/10.18632\/oncotarget.7327 (2016).","journal-title":"Oncotarget"},{"key":"28637_CR20","doi-asserted-by":"publisher","first-page":"3053","DOI":"10.1128\/MCB.00651-14","volume":"34","author":"C Hoffmann","year":"2014","unstructured":"Hoffmann, C. et al. Human Muscle LIM Protein Dimerizes along the Actin Cytoskeleton and Cross-Links Actin Filaments. Molecular and cellular biology 34, 3053\u20133065, https:\/\/doi.org\/10.1128\/MCB.00651-14 (2014).","journal-title":"Molecular and cellular biology"},{"key":"28637_CR21","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2121-6-45","volume":"6","author":"TC Tran","year":"2005","unstructured":"Tran, T. C., Singleton, C., Fraley, T. S. & Greenwood, J. A. Cysteine-rich protein 1 (CRP1) regulates actin filament bundling. BMC Cell Biol 6, 45 (2005).","journal-title":"BMC Cell Biol"},{"key":"28637_CR22","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2164-7-96","volume":"7","author":"Z Hu","year":"2006","unstructured":"Hu, Z. et al. The molecular portraits of breast tumors are conserved across microarray platforms. BMC Genomics 7, 96, https:\/\/doi.org\/10.1186\/1471-2164-7-96 (2006).","journal-title":"BMC Genomics"},{"key":"28637_CR23","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1631\/jzus.B1400221","volume":"16","author":"ZJ Liu","year":"2015","unstructured":"Liu, Z. J., Semenza, G. L. & Zhang, H. F. Hypoxia-inducible factor 1 and breast cancer metastasis. J Zhejiang Univ Sci B 16, 32\u201343, https:\/\/doi.org\/10.1631\/jzus.B1400221 (2015).","journal-title":"J Zhejiang Univ Sci B"},{"key":"28637_CR24","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1016\/j.tips.2012.01.005","volume":"33","author":"GL Semenza","year":"2012","unstructured":"Semenza, G. L. Hypoxia-inducible factors: mediators of cancer progression and targets for cancer therapy. Trends Pharmacol Sci 33, 207\u2013214, https:\/\/doi.org\/10.1016\/j.tips.2012.01.005 (2012).","journal-title":"Trends Pharmacol Sci"},{"key":"28637_CR25","doi-asserted-by":"publisher","first-page":"32529","DOI":"10.1074\/jbc.271.51.32529","volume":"271","author":"GL Semenza","year":"1996","unstructured":"Semenza, G. L. et al. Hypoxia response elements in the aldolase A, enolase 1, and lactate dehydrogenase A gene promoters contain essential binding sites for hypoxia-inducible factor 1. J Biol Chem 271, 32529\u201332537 (1996).","journal-title":"J Biol Chem"},{"key":"28637_CR26","doi-asserted-by":"publisher","first-page":"re12","DOI":"10.1126\/stke.3062005re12","volume":"2005","author":"RH Wenger","year":"2005","unstructured":"Wenger, R. H., Stiehl, D. P. & Camenisch, G. Integration of oxygen signaling at the consensus HRE. Sci Stke 2005, re12, https:\/\/doi.org\/10.1126\/stke.3062005re12 (2005).","journal-title":"Sci Stke"},{"key":"28637_CR27","doi-asserted-by":"publisher","first-page":"333","DOI":"10.1093\/bioinformatics\/18.2.333","volume":"18","author":"X Messeguer","year":"2002","unstructured":"Messeguer, X. et al. PROMO: detection of known transcription regulatory elements using species-tailored searches. Bioinformatics 18, 333\u2013334 (2002).","journal-title":"Bioinformatics"},{"key":"28637_CR28","doi-asserted-by":"publisher","first-page":"2379","DOI":"10.1038\/sj.onc.1209273","volume":"25","author":"UM Munoz-Najar","year":"2006","unstructured":"Munoz-Najar, U. M., Neurath, K. M., Vumbaca, F. & Claffey, K. P. Hypoxia stimulates breast carcinoma cell invasion through MT1-MMP and MMP-2 activation. Oncogene 25, 2379\u20132392, https:\/\/doi.org\/10.1038\/sj.onc.1209273 (2006).","journal-title":"Oncogene"},{"key":"28637_CR29","doi-asserted-by":"publisher","first-page":"3034","DOI":"10.1158\/0008-5472.CAN-05-2177","volume":"66","author":"VV Artym","year":"2006","unstructured":"Artym, V. V., Zhang, Y., Seillier-Moiseiwitsch, F., Yamada, K. M. & Mueller, S. C. Dynamic interactions of cortactin and membrane type 1 matrix metalloproteinase at invadopodia: defining the stages of invadopodia formation and function. Cancer research 66, 3034\u20133043, https:\/\/doi.org\/10.1158\/0008-5472.CAN-05-2177 (2006).","journal-title":"Cancer research"},{"key":"28637_CR30","doi-asserted-by":"publisher","first-page":"2079","DOI":"10.1016\/j.cub.2013.08.044","volume":"23","author":"VP Sharma","year":"2013","unstructured":"Sharma, V. P. et al. Tks5 and SHIP2 regulate invadopodium maturation, but not initiation, in breast carcinoma cells. Curr Biol 23, 2079\u20132089, https:\/\/doi.org\/10.1016\/j.cub.2013.08.044 (2013).","journal-title":"Curr Biol"},{"key":"28637_CR31","doi-asserted-by":"publisher","first-page":"1623","DOI":"10.2217\/fon.13.92","volume":"9","author":"DM Gilkes","year":"2013","unstructured":"Gilkes, D. M. & Semenza, G. L. Role of hypoxia-inducible factors in breast cancer metastasis. Future Oncol 9, 1623\u20131636, https:\/\/doi.org\/10.2217\/fon.13.92 (2013).","journal-title":"Future Oncol"},{"key":"28637_CR32","doi-asserted-by":"publisher","first-page":"725","DOI":"10.1007\/s10549-009-0674-9","volume":"123","author":"B Gyorffy","year":"2010","unstructured":"Gyorffy, B. et al. An online survival analysis tool to rapidly assess the effect of 22,277 genes on breast cancer prognosis using microarray data of 1,809 patients. Breast cancer research and treatment 123, 725\u2013731, https:\/\/doi.org\/10.1007\/s10549-009-0674-9 (2010).","journal-title":"Breast cancer research and treatment"},{"key":"28637_CR33","doi-asserted-by":"publisher","first-page":"4","DOI":"10.3389\/fcell.2015.00004","volume":"3","author":"A Jacob","year":"2015","unstructured":"Jacob, A. & Prekeris, R. The regulation of MMP targeting to invadopodia during cancer metastasis. Front Cell Dev Biol 3, 4, https:\/\/doi.org\/10.3389\/fcell.2015.00004 (2015).","journal-title":"Front Cell Dev Biol"},{"key":"28637_CR34","doi-asserted-by":"publisher","first-page":"9096","DOI":"10.1158\/0008-5472.CAN-08-2522","volume":"68","author":"Y Nishida","year":"2008","unstructured":"Nishida, Y. et al. Activation of matrix metalloproteinase-2 (MMP-2) by membrane type 1 matrix metalloproteinase through an artificial receptor for proMMP-2 generates active MMP-2. Cancer research 68, 9096\u20139104, https:\/\/doi.org\/10.1158\/0008-5472.CAN-08-2522 (2008).","journal-title":"Cancer research"},{"key":"28637_CR35","doi-asserted-by":"publisher","first-page":"386","DOI":"10.1016\/S0006-291X(03)01405-0","volume":"308","author":"M Toth","year":"2003","unstructured":"Toth, M., Chvyrkova, I., Bernardo, M. M., Hernandez-Barrantes, S. & Fridman, R. Pro-MMP-9 activation by the MT1-MMP\/MMP-2 axis and MMP-3: role of TIMP-2 and plasma membranes. Biochem Biophys Res Commun 308, 386\u2013395 (2003).","journal-title":"Biochem Biophys Res Commun"},{"key":"28637_CR36","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1111\/cas.13134","volume":"108","author":"Z Li","year":"2017","unstructured":"Li, Z., Takino, T., Endo, Y. & Sato, H. Activation of MMP-9 by membrane type-1 MMP\/MMP-2 axis stimulates tumor metastasis. Cancer Sci 108, 347\u2013353, https:\/\/doi.org\/10.1111\/cas.13134 (2017).","journal-title":"Cancer Sci"},{"key":"28637_CR37","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1038\/ncb3295","volume":"18","author":"A Lin","year":"2016","unstructured":"Lin, A. et al. The LINK-A lncRNA activates normoxic HIF1alpha signalling in triple-negative breast cancer. Nat Cell Biol 18, 213\u2013224, https:\/\/doi.org\/10.1038\/ncb3295 (2016).","journal-title":"Nat Cell Biol"},{"key":"28637_CR38","doi-asserted-by":"publisher","first-page":"2078","DOI":"10.1172\/JCI66715","volume":"123","author":"SC Hanna","year":"2013","unstructured":"Hanna, S. C. et al. HIF1alpha and HIF2alpha independently activate SRC to promote melanoma metastases. J Clin Invest 123, 2078\u20132093, https:\/\/doi.org\/10.1172\/JCI66715 (2013).","journal-title":"J Clin Invest"},{"key":"28637_CR39","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1083\/jcb.201209151","volume":"201","author":"B Diaz","year":"2013","unstructured":"Diaz, B., Yuen, A., Iizuka, S., Higashiyama, S. & Courtneidge, S. A. Notch increases the shedding of HB-EGF by ADAM12 to potentiate invadopodia formation in hypoxia. The Journal of cell biology 201, 279\u2013292, https:\/\/doi.org\/10.1083\/jcb.201209151 (2013).","journal-title":"The Journal of cell biology"},{"key":"28637_CR40","doi-asserted-by":"publisher","first-page":"e55529","DOI":"10.1371\/journal.pone.0055529","volume":"8","author":"D Arsenault","year":"2013","unstructured":"Arsenault, D., Brochu-Gaudreau, K., Charbonneau, M. & Dubois, C. M. HDAC6 deacetylase activity is required for hypoxia-induced invadopodia formation and cell invasion. Plos One 8, e55529, https:\/\/doi.org\/10.1371\/journal.pone.0055529 (2013).","journal-title":"Plos One"},{"key":"28637_CR41","doi-asserted-by":"publisher","first-page":"e28851","DOI":"10.1371\/journal.pone.0028851","volume":"6","author":"F Lucien","year":"2011","unstructured":"Lucien, F., Brochu-Gaudreau, K., Arsenault, D., Harper, K. & Dubois, C. M. Hypoxia-induced invadopodia formation involves activation of NHE-1 by the p90 ribosomal S6 kinase (p90RSK). Plos One 6, e28851, https:\/\/doi.org\/10.1371\/journal.pone.0028851 (2011).","journal-title":"Plos One"},{"key":"28637_CR42","doi-asserted-by":"publisher","first-page":"2455","DOI":"10.1158\/0008-5472.CAN-13-3009","volume":"74","author":"X Zhao","year":"2014","unstructured":"Zhao, X. et al. Hypoxia-inducible factor-1 promotes pancreatic ductal adenocarcinoma invasion and metastasis by activating transcription of the actin-bundling protein fascin. Cancer research 74, 2455\u20132464, https:\/\/doi.org\/10.1158\/0008-5472.CAN-13-3009 (2014).","journal-title":"Cancer research"},{"key":"28637_CR43","doi-asserted-by":"publisher","first-page":"339","DOI":"10.1016\/j.cub.2009.12.035","volume":"20","author":"A Li","year":"2010","unstructured":"Li, A. et al. The actin-bundling protein fascin stabilizes actin in invadopodia and potentiates protrusive invasion. Curr Biol 20, 339\u2013345, https:\/\/doi.org\/10.1016\/j.cub.2009.12.035 (2010).","journal-title":"Curr Biol"},{"key":"28637_CR44","doi-asserted-by":"publisher","first-page":"1805","DOI":"10.1096\/fj.13-242537","volume":"28","author":"I Van Audenhove","year":"2014","unstructured":"Van Audenhove, I. et al. Stratifying fascin and cortactin function in invadopodium formation using inhibitory nanobodies and targeted subcellular delocalization. FASEB journal: official publication of the Federation of American Societies for Experimental Biology 28, 1805\u20131818, https:\/\/doi.org\/10.1096\/fj.13-242537 (2014).","journal-title":"FASEB journal: official publication of the Federation of American Societies for Experimental Biology"},{"key":"28637_CR45","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1016\/j.ebiom.2016.10.026","volume":"13","author":"M Ohtsuka","year":"2016","unstructured":"Ohtsuka, M. et al. H19 Noncoding RNA, an Independent Prognostic Factor, Regulates Essential Rb-E2F and CDK8-beta-Catenin Signaling in Colorectal Cancer. EBioMedicine 13, 113\u2013124, https:\/\/doi.org\/10.1016\/j.ebiom.2016.10.026 (2016).","journal-title":"EBioMedicine"},{"key":"28637_CR46","doi-asserted-by":"publisher","unstructured":"Wang, S. J. et al. Cysteine and glycine-rich protein 2 (CSRP2) transcript levels correlate with leukemia relapse and leukemia-free survival in adults with B-cell acute lymphoblastic leukemia and normal cytogenetics. Oncotarget, https:\/\/doi.org\/10.18632\/oncotarget.16416 (2017).","DOI":"10.18632\/oncotarget.16416"},{"key":"28637_CR47","doi-asserted-by":"publisher","unstructured":"Schito, L., Rey, S. & Konopleva, M. Integration of hypoxic HIF-alpha signaling in blood cancers. Oncogene, https:\/\/doi.org\/10.1038\/onc.2017.119 (2017).","DOI":"10.1038\/onc.2017.119"},{"key":"28637_CR48","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1111\/j.1365-2818.2006.01706.x","volume":"224","author":"S Bolte","year":"2006","unstructured":"Bolte, S. & Cordelieres, F. P. A guided tour into subcellular colocalization analysis in light microscopy. J Microsc 224, 213\u2013232, https:\/\/doi.org\/10.1111\/j.1365-2818.2006.01706.x (2006).","journal-title":"J Microsc"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-018-28637-x.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-018-28637-x","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-018-28637-x.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,21]],"date-time":"2022-12-21T07:12:00Z","timestamp":1671606720000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-018-28637-x"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2018,7,5]]},"references-count":48,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2018,12]]}},"alternative-id":["28637"],"URL":"https:\/\/doi.org\/10.1038\/s41598-018-28637-x","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2018,7,5]]},"assertion":[{"value":"8 February 2018","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"13 June 2018","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"5 July 2018","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing Interests"}}],"article-number":"10191"}}