{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,27]],"date-time":"2026-04-27T20:18:08Z","timestamp":1777321088923,"version":"3.51.4"},"reference-count":70,"publisher":"MDPI AG","issue":"14","license":[{"start":{"date-parts":[[2019,7,12]],"date-time":"2019-07-12T00:00:00Z","timestamp":1562889600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000289","name":"Cancer Research UK","doi-asserted-by":"publisher","award":["A13044"],"award-info":[{"award-number":["A13044"]}],"id":[{"id":"10.13039\/501100000289","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000781","name":"European Research Council","doi-asserted-by":"publisher","award":["250170"],"award-info":[{"award-number":["250170"]}],"id":[{"id":"10.13039\/501100000781","id-type":"DOI","asserted-by":"publisher"}]},{"name":"CRUK Beatson Institute","award":["C596\/A17196"],"award-info":[{"award-number":["C596\/A17196"]}]},{"DOI":"10.13039\/501100000268","name":"Biotechnology and Biological Sciences Research Council","doi-asserted-by":"publisher","award":["1906000"],"award-info":[{"award-number":["1906000"]}],"id":[{"id":"10.13039\/501100000268","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Background: Cutaneous squamous cell carcinoma (cSCC) incidence continues to rise with increasing morbidity and mortality, with limited treatment options for advanced disease. Future improvements in targeted therapy will rely on advances in genomic\/transcriptomic understanding and the use of model systems for basic research. We describe here the panel of 16 primary and metastatic cSCC cell lines developed and characterised over the past three decades in our laboratory in order to provide such a resource for future preclinical research and drug screening. Methods: Primary keratinocytes were isolated from cSCC tumours and metastases, and cell lines were established. These were characterised using short tandem repeat (STR) profiling and genotyped by whole exome sequencing. Multiple in vitro assays were performed to document their morphology, growth characteristics, migration and invasion characteristics, and in vivo xenograft growth. Results: STR profiles of the cSCC lines allow the confirmation of their unique identity. Phylogenetic trees derived from exome sequence analysis of the matched primary and metastatic lines provide insight into the genetic basis of disease progression. The results of in vivo and in vitro analyses allow researchers to select suitable cell lines for specific experimentation. Conclusions: There are few well-characterised cSCC lines available for widespread preclinical experimentation and drug screening. The described cSCC cell line panel provides a critical tool for in vitro and in vivo experimentation.<\/jats:p>","DOI":"10.3390\/ijms20143428","type":"journal-article","created":{"date-parts":[[2019,7,12]],"date-time":"2019-07-12T11:49:38Z","timestamp":1562932178000},"page":"3428","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":21,"title":["A Unique Panel of Patient-Derived Cutaneous Squamous Cell Carcinoma Cell Lines Provides a Preclinical Pathway for Therapeutic Testing"],"prefix":"10.3390","volume":"20","author":[{"given":"Sakinah","family":"Hassan","sequence":"first","affiliation":[{"name":"Blizard Institute, Barts and the London School of Medicine and Dentistry, QMUL, London E1 2AT, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Karin J.","family":"Purdie","sequence":"additional","affiliation":[{"name":"Blizard Institute, Barts and the London School of Medicine and Dentistry, QMUL, London E1 2AT, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2509-9599","authenticated-orcid":false,"given":"Jun","family":"Wang","sequence":"additional","affiliation":[{"name":"Barts Cancer Institute, QMUL, London EC1M 6BQ, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Catherine A.","family":"Harwood","sequence":"additional","affiliation":[{"name":"Blizard Institute, Barts and the London School of Medicine and Dentistry, QMUL, London E1 2AT, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Charlotte M.","family":"Proby","sequence":"additional","affiliation":[{"name":"Division of Cancer, Ninewells Hospital and Medical School, University of Dundee, Dundee DD1 9SY, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0312-9734","authenticated-orcid":false,"given":"Celine","family":"Pourreyron","sequence":"additional","affiliation":[{"name":"Division of Cancer, Ninewells Hospital and Medical School, University of Dundee, Dundee DD1 9SY, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Nikol","family":"Mladkova","sequence":"additional","affiliation":[{"name":"Blizard Institute, Barts and the London School of Medicine and Dentistry, QMUL, London E1 2AT, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2354-660X","authenticated-orcid":false,"given":"Ai","family":"Nagano","sequence":"additional","affiliation":[{"name":"Barts Cancer Institute, QMUL, London EC1M 6BQ, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Sandeep","family":"Dhayade","sequence":"additional","affiliation":[{"name":"Cancer Research UK Beatson Institute, Garscube Estate, Switchback Rd, Glasgow G61 1BD, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5298-7941","authenticated-orcid":false,"given":"Dimitris","family":"Athineos","sequence":"additional","affiliation":[{"name":"Cancer Research UK Beatson Institute, Garscube Estate, Switchback Rd, Glasgow G61 1BD, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0504-9922","authenticated-orcid":false,"given":"Matthew","family":"Caley","sequence":"additional","affiliation":[{"name":"Blizard Institute, Barts and the London School of Medicine and Dentistry, QMUL, London E1 2AT, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Viviana","family":"Mannella","sequence":"additional","affiliation":[{"name":"Blizard Institute, Barts and the London School of Medicine and Dentistry, QMUL, London E1 2AT, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Karen","family":"Blyth","sequence":"additional","affiliation":[{"name":"Cancer Research UK Beatson Institute, Garscube Estate, Switchback Rd, Glasgow G61 1BD, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6264-4253","authenticated-orcid":false,"given":"Gareth J.","family":"Inman","sequence":"additional","affiliation":[{"name":"Cancer Research UK Beatson Institute, Garscube Estate, Switchback Rd, Glasgow G61 1BD, UK"},{"name":"Institute of Cancer Sciences, University of Glasgow, Glasgow G61 1GH, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Irene M.","family":"Leigh","sequence":"additional","affiliation":[{"name":"Blizard Institute, Barts and the London School of Medicine and Dentistry, QMUL, London E1 2AT, UK"},{"name":"Division of Cancer, Ninewells Hospital and Medical School, University of Dundee, Dundee DD1 9SY, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2019,7,12]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"298","DOI":"10.1001\/jamadermatol.2018.4219","article-title":"Nationwide Incidence of Metastatic Cutaneous Squamous Cell Carcinoma in England","volume":"155","author":"Venables","year":"2019","journal-title":"JAMA Dermatol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1056\/NEJMoa1805131","article-title":"PD-1 Blockade with Cemiplimab in Advanced Cutaneous Squamous-Cell Carcinoma","volume":"379","author":"Migden","year":"2018","journal-title":"N. Eng. J. Med."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"3667","DOI":"10.1038\/s41467-018-06027-1","article-title":"The genomic landscape of cutaneous SCC reveals drivers and a novel azathioprine associated mutational signature","volume":"9","author":"Inman","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"2630","DOI":"10.1038\/jid.2014.154","article-title":"NOTCH1 Mutations Occur Early during Cutaneous Squamous Cell Carcinogenesis","volume":"134","author":"South","year":"2014","journal-title":"J. Investig. Dermatol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"3","DOI":"10.2340\/00015555-2181","article-title":"The Promise of Genomics and the Development of Targeted Therapies for Cutaneous Squamous Cell Carcinoma","volume":"96","author":"Harwood","year":"2016","journal-title":"Acta Derm. Venereol."},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Zhao, M., Sano, D., Pickering, C.R., Jasser, S.A., Henderson, Y.C., Clayman, G.L., Sturgis, E.M., Ow, T.J., Lotan, R., and Carey, T.E. (2011). Assembly and Initial Characterization of a Panel of 85 Genomically Validated Cell Lines from Diverse Head and Neck Tumor Sites. Clin. Cancer Res., 17.","DOI":"10.1158\/1078-0432.CCR-11-0690"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1034\/j.1600-0625.2000.009002104.x","article-title":"Spontaneous keratinocyte cell lines representing early and advanced stages of malignant transformation of the epidermis","volume":"9","author":"Proby","year":"2000","journal-title":"Exp. Dermatol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1417","DOI":"10.1093\/jnci\/51.5.1417","article-title":"In Vitro Cultivation of Human Tumors: Establishment of Cell Lines Derived From a Series of Solid Tumors2","volume":"51","author":"Giard","year":"1973","journal-title":"JNCI J. Natl.Cancer Inst."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"741","DOI":"10.1016\/0092-8674(81)90100-8","article-title":"Epidermal growth factor induces rapid tyrosine phosphorylation of proteins in A431 human tumor cells","volume":"24","author":"Hunter","year":"1981","journal-title":"Cell"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1007\/BF02621233","article-title":"Incomplete epidermal differentiation of A431 epidermoid carcinoma cells","volume":"22","author":"Rosdy","year":"1986","journal-title":"In Vitro Cell. Dev. Biol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1016\/0092-8674(80)90373-6","article-title":"Defective terminal differentiation in culture as a consistent and selectable character of malignant human keratinocytes","volume":"22","author":"Rheinwald","year":"1980","journal-title":"Cell"},{"key":"ref_12","first-page":"3972","article-title":"Abnormal Expression of Retinoic Acid Receptors and Keratin 19 by Human Oral and Epidermal Squamous Cell Carcinoma Cell Lines","volume":"51","author":"Hu","year":"1991","journal-title":"Cancer Res."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"5267","DOI":"10.1038\/sj.onc.1210332","article-title":"p38\u03b1 and p38\u03b4 mitogen-activated protein kinase isoforms regulate invasion and growth of head and neck squamous carcinoma cells","volume":"26","author":"Junttila","year":"2007","journal-title":"Oncogene"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1110","DOI":"10.1016\/j.ajpath.2011.05.012","article-title":"Serpin peptidase inhibitor clade A member 1 (SerpinA1) is a novel biomarker for progression of cutaneous squamous cell carcinoma","volume":"179","author":"Farshchian","year":"2011","journal-title":"Am. J. Pathol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"45057","DOI":"10.1038\/srep45057","article-title":"Significant Role of Collagen XVII And Integrin \u03b24 in Migration and Invasion of The Less Aggressive Squamous Cell Carcinoma Cells","volume":"7","author":"Moilanen","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"45614","DOI":"10.1074\/jbc.M106559200","article-title":"Mitochondrial Respiration Is Uniquely Associated with the Prooxidant and Apoptotic Effects ofN-(4-Hydroxyphenyl)retinamide","volume":"276","author":"Hail","year":"2001","journal-title":"J. Biol. Chem."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1002\/ijc.2910610119","article-title":"Induction of apoptotic cell death in non-melanoma skin cancer by interferon-\u03b1","volume":"61","author":"McDonnell","year":"1995","journal-title":"Int. J. Cancer"},{"key":"ref_18","first-page":"2684","article-title":"A Human Squamous Cell Carcinoma Cell Line","volume":"35","author":"Moore","year":"1975","journal-title":"Cancer Res."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"2239","DOI":"10.3892\/ol.2017.6458","article-title":"Inactivated Wnt signaling in resveratrol-treated epidermal squamous cancer cells and its biological implication","volume":"14","author":"Liu","year":"2017","journal-title":"Oncol. Lett."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1111\/j.1600-0625.2011.01392.x","article-title":"CRABP-II- and FABP5-independent all-trans retinoic acid resistance in COLO 16 human cutaneous squamous cancer cells","volume":"21","author":"Chen","year":"2012","journal-title":"Exp. Dermatol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1963","DOI":"10.1158\/1078-0432.CCR-04-1665","article-title":"AEE788, a Dual Tyrosine Kinase Receptor Inhibitor, Induces Endothelial Cell Apoptosis in Human Cutaneous Squamous Cell Carcinoma Xenografts in Nude Mice","volume":"11","author":"Park","year":"2005","journal-title":"Clin. Cancer Res."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1002\/(SICI)1097-4652(199712)173:3<433::AID-JCP16>3.0.CO;2-C","article-title":"PTHrP and cell division: Expression and localization of PTHrP in a keratinocyte cell line (HaCaT) during the cell cycle","volume":"173","author":"Lam","year":"1997","journal-title":"J. Cell. Physiol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"147863","DOI":"10.1155\/2012\/147863","article-title":"Topical curcumin-based cream is equivalent to dietary curcumin in a skin cancer model","volume":"2012","author":"Sonavane","year":"2012","journal-title":"J. Skin Cancer"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1177\/0194599811400711","article-title":"Curcumin Inhibits Skin Squamous Cell Carcinoma Tumor Growth In Vivo","volume":"145","author":"Phillips","year":"2011","journal-title":"Otolaryngol. Head Neck Surg."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Kleiner-Hancock, H.E., Shi, R., Remeika, A., Robbins, D., Prince, M., Gill, J.N., Syed, Z., Adegboyega, P., Mathis, J.M., and Clifford, J.L. (2010). Effects of ATRA combined with citrus and ginger-derived compounds in human SCC xenografts. BMC Cancer, 10.","DOI":"10.1186\/1471-2407-10-394"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"364","DOI":"10.1200\/JCO.2002.20.2.364","article-title":"Phase II and Biologic Study of Interferon Alfa, Retinoic Acid, and Cisplatin in Advanced Squamous Skin Cancer","volume":"20","author":"Shin","year":"2002","journal-title":"J. Clin. Oncol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1920","DOI":"10.1016\/j.jid.2016.05.110","article-title":"MEK Is a Therapeutic and Chemopreventative Target in Squamous Cell Carcinoma","volume":"136","author":"Adelmann","year":"2016","journal-title":"J. Investig. Dermatol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1450","DOI":"10.1038\/nm.2265","article-title":"Invasive three-dimensional organotypic neoplasia from multiple normal human epithelia","volume":"16","author":"Ridky","year":"2010","journal-title":"Nat. Med."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Vaughan, M.B., Ramirez, R.D., Andrews, C.M., Wright, W.E., and Shay, J.W. (2009). H-Ras Expression in Immortalized Keratinocytes Produces an Invasive Epithelium in Cultured Skin Equivalents. PLoS ONE, 4.","DOI":"10.1371\/journal.pone.0007908"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1002\/(SICI)1098-2744(199811)23:3<144::AID-MC3>3.0.CO;2-U","article-title":"Multiple stages and genetic alterations in immortalization, malignant transformation, and tumor progression of human skin keratinocytes","volume":"23","author":"Fusenig","year":"1998","journal-title":"Mol. Carcinog."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1567","DOI":"10.1016\/S0002-9440(10)62541-2","article-title":"Tumor progression of skin carcinoma cells in vivo promoted by clonal selection, mutagenesis, and autocrine growth regulation by granulocyte colony-stimulating factor and granulocyte-macrophage colony-stimulating factor","volume":"159","author":"Mueller","year":"2001","journal-title":"Am. J. Pathol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"3522","DOI":"10.1158\/0008-5472.CAN-11-2996","article-title":"Fibroblast-Derived Dermal Matrix Drives Development of Aggressive Cutaneous Squamous Cell Carcinoma in Patients with Recessive Dystrophic Epidermolysis Bullosa","volume":"72","author":"Ng","year":"2012","journal-title":"Cancer Res."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"985","DOI":"10.1002\/jcb.24453","article-title":"New insights into the role of Runx1 in epithelial stem cell biology and pathology","volume":"114","author":"Scheitz","year":"2013","journal-title":"J. Cell. Biochem."},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Pourreyron, C., Reilly, L., Proby, C., Panteleyev, A., Fleming, C., McLean, K., South, A.P., and Foerster, J. (2012). Wnt5a Is Strongly Expressed at the Leading Edge in Non-Melanoma Skin Cancer, Forming Active Gradients, while Canonical Wnt Signalling Is Repressed. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0031827"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"570","DOI":"10.1038\/nature11005","article-title":"Systematic identification of genomic markers of drug sensitivity in cancer cells","volume":"483","author":"Garnett","year":"2012","journal-title":"Nature"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"603","DOI":"10.1038\/nature11003","article-title":"The Cancer Cell Line Encyclopedia enables predictive modelling of anticancer drug sensitivity","volume":"483","author":"Barretina","year":"2012","journal-title":"Nature"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"20265","DOI":"10.18632\/oncotarget.24750","article-title":"Preclinical comparison of proteasome and ubiquitin E1 enzyme inhibitors in cutaneous squamous cell carcinoma: The identification of mechanisms of differential sensitivity","volume":"9","author":"McHugh","year":"2018","journal-title":"Oncotarget"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"23029","DOI":"10.18632\/oncotarget.25196","article-title":"Targeting the spliceosome for cutaneous squamous cell carcinoma therapy: A role for c-MYC and wild-type p53 in determining the degree of tumour selectivity","volume":"9","author":"Hepburn","year":"2018","journal-title":"Oncotarget"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"946","DOI":"10.1038\/nm.3878","article-title":"Genomic landscape of carcinogen-induced and genetically induced mouse skin squamous cell carcinoma","volume":"21","author":"Nassar","year":"2015","journal-title":"Nat. Med."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1158\/1940-6207.CAPR-16-0218","article-title":"Whole-Exome Sequencing Validates a Preclinical Mouse Model for the Prevention and Treatment of Cutaneous Squamous Cell Carcinoma","volume":"10","author":"Knatko","year":"2017","journal-title":"Cancer Prev. Res."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"12493","DOI":"10.1038\/ncomms12493","article-title":"Inactivation of TGF\u03b2 receptors in stem cells drives cutaneous squamous cell carcinoma","volume":"7","author":"Cammareri","year":"2016","journal-title":"Nat. Commun."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1111\/j.1600-6143.2012.04292.x","article-title":"A Surveillance Model for Skin Cancer in Organ Transplant Recipients: A 22-Year Prospective Study in an Ethnically Diverse Population","volume":"13","author":"Harwood","year":"2013","journal-title":"Am. J. Transplant."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1095","DOI":"10.1046\/j.1523-1747.2000.00173.x","article-title":"Genetic Characterization of a Human Skin Carcinoma Progression Model: From Primary Tumor to Metastasis","volume":"115","author":"Popp","year":"2000","journal-title":"J. Investig. Dermatol."},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Cree, I.A. (2011). Isolation and Culture of Squamous Cell Carcinoma Lines. Cancer Cell Culture: Methods and Protocols, Humana Press.","DOI":"10.1007\/978-1-61779-080-5"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"16910","DOI":"10.1073\/pnas.1009843107","article-title":"Allele-specific copy number analysis of tumors","volume":"107","author":"Nordgard","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"557","DOI":"10.1093\/biostatistics\/kxh008","article-title":"Circular binary segmentation for the analysis of array-based DNA copy number data","volume":"5","author":"Olshen","year":"2004","journal-title":"Biostatistics"},{"key":"ref_47","unstructured":"Bengtsson, H., Simpson, K., Bullard, J., and Hansen, K. (2008). aroma. affymetrix: A Generic Framework in R for Analyzing Small to Very Large Affymetrix Data Sets in Bounded Memory, Department of Epidemiology and Biostatistics, University of California San Francisco. Technical Report."},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Bengtsson, H., Neuvial, P., and Speed, T.P. (2010). TumorBoost: Normalization of allele-specific tumor copy numbers from a single pair of tumor-normal genotyping microarrays. BMC Bioinform., 11.","DOI":"10.1186\/1471-2105-11-245"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"2038","DOI":"10.1093\/bioinformatics\/btr329","article-title":"Parent-specific copy number in paired tumor-normal studies using circular binary segmentation","volume":"27","author":"Olshen","year":"2011","journal-title":"Bioinformatics"},{"key":"ref_50","first-page":"406","article-title":"The neighbor-joining method: a new method for reconstructing phylogenetic trees","volume":"4","author":"Nei","year":"1987","journal-title":"Mol. Biol. Evolut."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1446","DOI":"10.1038\/sj.bjc.6603135","article-title":"Overexpression of the Axl tyrosine kinase receptor in cutaneous SCC-derived cell lines and tumours","volume":"94","author":"Green","year":"2006","journal-title":"Br. J. Cancer"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1788","DOI":"10.1242\/jcs.042895","article-title":"Increased invasive behaviour in cutaneous squamous cell carcinoma with loss of basement-membrane type VII collagen","volume":"122","author":"Martins","year":"2009","journal-title":"J. Cell Sci."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"2277","DOI":"10.1038\/jid.2010.124","article-title":"The Flavonoid Luteolin Increases the Resistance of Normal, but Not Malignant Keratinocytes, Against UVB-Induced Apoptosis","volume":"130","author":"Verschooten","year":"2010","journal-title":"J. Investig. Dermatol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"2790","DOI":"10.1002\/ijc.25300","article-title":"Concomitant inhibition of AKT and autophagy is required for efficient cisplatin-induced apoptosis of metastatic skin carcinoma","volume":"127","author":"Claerhout","year":"2010","journal-title":"Int. J. Cancer"},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Ismail, F., Ikram, M., Purdie, K., Harwood, C., Leigh, I., and Storey, A. (2011). Cutaneous Squamous Cell Carcinoma (SCC) and the DNA Damage Response: pATM Expression Patterns in Pre-Malignant and Malignant Keratinocyte Skin Lesions. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0021271"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1038\/jid.2010.326","article-title":"Axl Promotes Cutaneous Squamous Cell Carcinoma Survival through Negative Regulation of Pro-Apoptotic Bcl-2 Family Members","volume":"131","author":"Papadakis","year":"2011","journal-title":"J. Investig. Dermatol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"17761","DOI":"10.1073\/pnas.1114669108","article-title":"Loss-of-function mutations in Notch receptors in cutaneous and lung squamous cell carcinoma","volume":"108","author":"Wang","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"4666","DOI":"10.1038\/onc.2011.180","article-title":"Integrative mRNA profiling comparing cultured primary cells with clinical samples reveals PLK1 and C20orf20 as therapeutic targets in cutaneous squamous cell carcinoma","volume":"30","author":"Watt","year":"2011","journal-title":"Oncogene"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1002\/path.2986","article-title":"The retinoid signalling molecule, TRIM16, is repressed during squamous cell carcinoma skin carcinogenesis in vivo and reduces skin cancer cell migration in vitro","volume":"226","author":"Cheung","year":"2012","journal-title":"J. Pathol."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Verschooten, L., Barrette, K., Van Kelst, S., Rubio Romero, N., Proby, C., De Vos, R., Agostinis, P., and Garmyn, M. (2012). Autophagy inhibitor chloroquine enhanced the cell death inducing effect of the flavonoid luteolin in metastatic squamous cell carcinoma cells. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0048264"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1039\/c2pp25064k","article-title":"Pattern of sensitivity of progressive cutaneous squamous cell carcinoma cells to UVB and oxidative stress-induced cell death","volume":"12","author":"Barrette","year":"2013","journal-title":"Photochem. Photobiol. Sci."},{"key":"ref_62","first-page":"4185","article-title":"The receptor tyrosine kinase Axl regulates cell\u2013cell adhesion and stemness in cutaneous squamous cell carcinoma","volume":"33","author":"Szentpetery","year":"2013","journal-title":"Oncogene"},{"key":"ref_63","first-page":"5303","article-title":"Eps8 controls Src- and FAK-dependent phenotypes in squamous carcinoma cells","volume":"127","author":"Schoenherr","year":"2014","journal-title":"J. Cell Sci."},{"key":"ref_64","first-page":"740","article-title":"Type VII collagen regulates expression of OATP1B3, promotes front-to-rear polarity and increases structural organisation in 3D spheroid cultures of RDEB tumour keratinocytes","volume":"127","author":"Dayal","year":"2014","journal-title":"J. Cell Sci."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"36426","DOI":"10.18632\/oncotarget.5536","article-title":"Cytoskeletal protein Flightless I inhibits apoptosis, enhances tumor cell invasion and promotes cutaneous squamous cell carcinoma progression","volume":"6","author":"Kopecki","year":"2015","journal-title":"Oncotarget"},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Martins, V.L., Caley, M.P., Moore, K., Szentpetery, Z., Marsh, S.T., Murrell, D.F., Kim, M.H., Avari, M., McGrath, J.A., and Cerio, R. (2015). Suppression of TGF\u03b2 and Angiogenesis by Type VII Collagen in Cutaneous SCC. JNCI J. Natl. Cancer Inst., 108.","DOI":"10.1093\/jnci\/djv293"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"14552","DOI":"10.18632\/oncotarget.24545","article-title":"Reduced SMAD2\/3 activation independently predicts increased depth of human cutaneous squamous cell carcinoma","volume":"9","author":"Rose","year":"2018","journal-title":"Oncotarget"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"e120750","DOI":"10.1172\/jci.insight.120750","article-title":"Ruxolitinib inhibits cyclosporine-induced proliferation of cutaneous squamous cell carcinoma","volume":"3","author":"Roudiani","year":"2018","journal-title":"JCI Insight"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1038\/s41467-017-02242-4","article-title":"Targeting the CoREST complex with dual histone deacetylase and demethylase inhibitors","volume":"9","author":"Kalin","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Kalin, J.H., Eroglu, A., Liu, H., Holtzclaw, W.D., Leigh, I., Proby, C.M., Fahey, J.W., Cole, P.A., and Dinkova-Kostova, A.T. (2019). Investigation into the use of histone deacetylase inhibitor MS-275 as a topical agent for the prevention and treatment of cutaneous squamous cell carcinoma in an SKH-1 hairless mouse model. PLoS ONE, 14.","DOI":"10.1371\/journal.pone.0213095"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/20\/14\/3428\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T13:04:58Z","timestamp":1760187898000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/20\/14\/3428"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,7,12]]},"references-count":70,"journal-issue":{"issue":"14","published-online":{"date-parts":[[2019,7]]}},"alternative-id":["ijms20143428"],"URL":"https:\/\/doi.org\/10.3390\/ijms20143428","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,7,12]]}}}