{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T13:49:15Z","timestamp":1790603355876,"version":"4.1.0"},"reference-count":60,"publisher":"MDPI AG","issue":"10","license":[{"start":{"date-parts":[[2025,5,21]],"date-time":"2025-05-21T00:00:00Z","timestamp":1747785600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100015999","name":"Oracle Cancer Trust","doi-asserted-by":"publisher","award":["Nu-000109"],"award-info":[{"award-number":["Nu-000109"]}],"id":[{"id":"10.13039\/100015999","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Papillary thyroid cancer (PTC) contains mesenchymal stem\/stromal cells (MSCs), but their contribution to PTC progression is not clear. In this study, we compared the transcriptional signatures of normal thyroid (NT) and PTC-derived MSCs with the aim of determining if these have distinct transcriptomes that might influence PTC progression. We used flow cytometry in combination with a panel of MSC clusters of differentiation (CD) markers and showed that both thyroid MSC populations expressed MSC markers and lacked expression of markers not normally expressed by MSCs. In addition, we determined that both MSC populations could differentiate to adipocytes and osteocytes. Analysis of single cell RNA sequencing data from both MSC populations revealed, regardless of tissue of origin, that both contained similar numbers of subpopulations. Cluster analysis revealed similarity in expression of both MSC populations for stromal markers, the vascular marker VEGFA and the smooth muscle marker CALD1, while smaller subpopulations expressed markers of more lineage-committed thyroid cells. PTC MSCs also showed upregulated expression of 28 genes, many of which are known to be involved in epithelial\u2013mesenchymal transition (EMT) and\/or disease progression in several types of cancers, including but not limited to breast cancer, gastric cancer, cervical carcinoma, bladder cancer and thyroid cancer. This included several members of the S100 and IGFBP gene families. Taken together, these data support a role for PTC MSCs in PTC progression.<\/jats:p>","DOI":"10.3390\/ijms26104957","type":"journal-article","created":{"date-parts":[[2025,5,21]],"date-time":"2025-05-21T13:59:43Z","timestamp":1747835983000},"page":"4957","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Single Cell RNA Sequencing of Papillary Cancer Mesenchymal Stem\/Stromal Cells Reveals a Transcriptional Profile That Supports a Role for These Cells in Cancer Progression"],"prefix":"10.3390","volume":"26","author":[{"given":"Danny","family":"Jandu","sequence":"first","affiliation":[{"name":"Bioscience Institute, Newcastle University, International Centre for Life, Newcastle upon Tyne NE1 3BZ, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5508-0659","authenticated-orcid":false,"given":"Nani","family":"Latar","sequence":"additional","affiliation":[{"name":"Bioscience Institute, Newcastle University, International Centre for Life, Newcastle upon Tyne NE1 3BZ, UK"},{"name":"Department of Surgery, Faculty of Medicine, Universiti Kebangsaan Malaysia Medical Centre, Cheras, Kuala Lumpur 56000, Malaysia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Artida","family":"Bajrami","sequence":"additional","affiliation":[{"name":"Bioscience Institute, Newcastle University, International Centre for Life, Newcastle upon Tyne NE1 3BZ, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rachel","family":"Queen","sequence":"additional","affiliation":[{"name":"Bioscience Institute, Newcastle University, International Centre for Life, Newcastle upon Tyne NE1 3BZ, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0325-4298","authenticated-orcid":false,"given":"Megan","family":"Hasoon","sequence":"additional","affiliation":[{"name":"Computational Biology Facility, Liverpool Shared Research Facilities, Faculty of Health and Life Sciences, University of Liverpool, Liverpool L69 3DR, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7376-9975","authenticated-orcid":false,"given":"Matthew","family":"Teasdale","sequence":"additional","affiliation":[{"name":"Bioinformatics Support Unit, The Faculty of Medical Sciences, Newcastle University, Newcastle upon Tyne NE2 4HH, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rafiqul","family":"Hussain","sequence":"additional","affiliation":[{"name":"Bioscience Institute, Newcastle University, International Centre for Life, Newcastle upon Tyne NE1 3BZ, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jonathan","family":"Coxhead","sequence":"additional","affiliation":[{"name":"Bioscience Institute, Newcastle University, International Centre for Life, Newcastle upon Tyne NE1 3BZ, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sebastian","family":"Aspinall","sequence":"additional","affiliation":[{"name":"Department of General Surgery, Aberdeen Royal Infirmary, Aberdeen AB25 2ZN, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5024-5511","authenticated-orcid":false,"given":"Annette","family":"Meeson","sequence":"additional","affiliation":[{"name":"Bioscience Institute, Newcastle University, International Centre for Life, Newcastle upon Tyne NE1 3BZ, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,5,21]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"2115","DOI":"10.1002\/sctm.17-0138","article-title":"Concise Review: Cancer Cells, Cancer Stem Cells, and Mesenchymal Stem Cells: Influence in Cancer Development","volume":"6","author":"Papaccio","year":"2017","journal-title":"Stem Cells Transl. Med."},{"key":"ref_2","first-page":"4","article-title":"TGF-\u03b2, to target or not to target; to prevent thyroid cancer progression?","volume":"1877","author":"Karnik","year":"2022","journal-title":"BBA-Rev. Cancer"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1007\/s10549-010-0734-1","article-title":"Potential role of mesenchymal stem cells (MSCs) in the breast tumour microenvironment: Stimulation of epithelial to mesenchymal transition (EMT)","volume":"124","author":"Martin","year":"2010","journal-title":"Breast Cancer Res. Treat."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"429","DOI":"10.1016\/j.neo.2017.02.010","article-title":"Mesenchymal Stem Cells Induce Epithelial to Mesenchymal Transition in Colon Cancer Cells through Direct Cell-to-Cell Contact","volume":"19","author":"Takigawa","year":"2017","journal-title":"Neoplasia"},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Parascandolo, A., Rappa, F., Cappello, F., Kim, J., Cantu, D.A., Chen, H., Mazzoccoli, G., Hematti, P., Castellone, M.D., and Salvatore, M. (2017). Extracellular Superoxide Dismutase Expression in Papillary Thyroid Cancer Mesenchymal Stem\/Stromal Cells Modulates Cancer Cell Growth and Migration. Sci. Rep., 7.","DOI":"10.1038\/srep41416"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1080\/14653240500319234","article-title":"Clarification of the nomenclature for MSC: The International Society for Cellular Therapy position statement","volume":"7","author":"Horwitz","year":"2005","journal-title":"Cytotherapy"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1080\/14653240600855905","article-title":"Minimal criteria for defining multipotent mesenchymal stromal cells. The International Society for Cellular Therapy position statement","volume":"8","author":"Dominici","year":"2006","journal-title":"Cytotherapy"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1019","DOI":"10.1016\/j.jcyt.2019.08.002","article-title":"Mesenchymal stem versus stromal cells: International Society for Cell & Gene Therapy (ISCT\u00ae) Mesenchymal Stromal Cell committee position statement on nomenclature","volume":"21","author":"Viswanathan","year":"2019","journal-title":"Cytotherapy"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1111\/jcmm.12168","article-title":"CD146 expression on mesenchymal stem cells is associated with their vascular smooth muscle commitment","volume":"18","author":"Espagnolle","year":"2014","journal-title":"J. Cell Mol. Med."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"118","DOI":"10.15283\/ijsc.2014.7.2.118","article-title":"Comparison of Mesenchymal Stem Cell Markers in Multiple Human Adult Stem Cells","volume":"7","author":"Maleki","year":"2014","journal-title":"Int. J. Stem Cells"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1408","DOI":"10.1002\/stem.1681","article-title":"The Surface Markers and Identity of Human Mesenchymal Stem Cells","volume":"32","author":"Lv","year":"2014","journal-title":"Stem Cells"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"6689","DOI":"10.1096\/fj.201802224R","article-title":"Thy-1: More than a marker for mesenchymal stromal cells","volume":"33","author":"Saalbach","year":"2019","journal-title":"FASEB J."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1038\/s41536-022-00258-z","article-title":"Enforced mesenchymal stem cell tissue colonization counteracts immunopathology","volume":"7","author":"Blanquer","year":"2022","journal-title":"npj Regen. Med."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"100031","DOI":"10.1016\/j.gene.2020.100031","article-title":"ALCAM (CD166) as a gene expression marker for human mesenchymal stromal cell characterisation","volume":"763","author":"Brinkhof","year":"2020","journal-title":"Gene"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1038\/s41419-017-0176-3","article-title":"Human colorectal cancer-derived mesenchymal stem cells promote colorectal cancer progression through IL-6\/JAK2\/STAT3 signaling","volume":"9","author":"Zhang","year":"2018","journal-title":"Cell Death Dis."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1637","DOI":"10.4251\/wjgo.v14.i9.1637","article-title":"Emerging role of caldesmon in cancer: A potential biomarker for colorectal cancer and other cancers","volume":"14","author":"Alnuaimi","year":"2022","journal-title":"World J. Gastrointest. Oncol."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Brun, J., Lutz, K.A., Neumayer, K.M.H., Klein, G., Seeger, T., Uynuk-Ool, T., W\u00f6rg\u00f6tter, K., Schmid, S., Kraushaar, U., and Guenther, E. (2015). Smooth Muscle Like Cells Generated from Human Mesenchymal Stromal Cells Display Marker Gene Expression and Electrophysiological Competence Comparable to Bladder Smooth Muscle Cells. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0145153"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"3682","DOI":"10.1038\/onc.2011.85","article-title":"SCUBE3 is an endogenous TGF-\u03b2 receptor ligand and regulates the epithelial-mesenchymal transition in lung cancer","volume":"30","author":"Wu","year":"2011","journal-title":"Oncogene"},{"key":"ref_19","first-page":"2272","article-title":"MARCKSL1 promotes the proliferation, migration and invasion of lung adenocarcinoma cells","volume":"19","author":"Liang","year":"2020","journal-title":"Oncol. Lett."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Tian, T., Leng, Y., Tang, B., Dong, X., Ren, Q., Liang, J., Liu, T., Liu, Y., Feng, W., and Liu, S. (2024). The oncogenic role and regulatory mechanism of PGK1 in human non-small cell lung cancer. Biol. Direct., 19.","DOI":"10.1186\/s13062-023-00448-9"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"4167","DOI":"10.1007\/s13277-015-3051-9","article-title":"SOX4 overexpression is a novel biomarker of malignant status and poor prognosis in breast cancer patients","volume":"36","author":"Song","year":"2015","journal-title":"Tumour Biol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"468","DOI":"10.1186\/s12935-020-01568-2","article-title":"SOX4 promotes the growth and metastasis of breast cancer","volume":"29","author":"Zhang","year":"2020","journal-title":"Cancer Cell Int."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"4695","DOI":"10.1038\/s41388-020-1319-z","article-title":"SOX4 activates CXCL12 in hepatocellular carcinoma cells to modulate endothelial cell migration and angiogenesis in vivo","volume":"39","author":"Tsai","year":"2020","journal-title":"Oncogene"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1207","DOI":"10.1007\/s12094-019-02042-w","article-title":"Elevated expression of GNAS promotes breast cancer cell proliferation and migration via the PI3K\/AKT\/Snail1\/E-cadherin axis","volume":"21","author":"Jin","year":"2019","journal-title":"Clin. Transl. Oncol."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Ding, H., Zhang, X., Su, Y., Jia, C., and Dai, C. (2020). GNAS promotes inflammation-related hepatocellular carcinoma progression by promoting STAT3 activation. Cell Mol. Biol. Lett., 25.","DOI":"10.1186\/s11658-020-00204-1"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"2019","DOI":"10.1042\/CS20190704","article-title":"A novel function for HEG1 in promoting metastasis in hepatocellular carcinoma","volume":"133","author":"Dewdney","year":"2019","journal-title":"Clin. Sci."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1186\/s12935-021-02117-1","article-title":"Plastin-3 is a diagnostic and prognostic marker for pancreatic adenocarcinoma and distinguishes from diffuse large B-cell lymphoma","volume":"21","author":"Xiong","year":"2021","journal-title":"Cancer Cell Int."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1456","DOI":"10.1007\/s12325-023-02428-x","article-title":"Bioinformatics Identification of Therapeutic Gene Targets for Gastric Cancer","volume":"40","author":"Li","year":"2023","journal-title":"Adv. Ther."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1038\/s41388-022-02537-x","article-title":"CST1 inhibits ferroptosis and promotes gastric cancer metastasis by regulating GPX4 protein stability via OTUB1","volume":"42","author":"Li","year":"2023","journal-title":"Oncogene"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"873","DOI":"10.1007\/s00109-017-1537-1","article-title":"Elevated expression of CST1 promotes breast cancer progression and predicts a poor prognosis","volume":"95","author":"Dai","year":"2017","journal-title":"J. Mol. Med."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1038\/s41420-020-00322-x","article-title":"EDIL3 promotes epithelial-mesenchymal transition and paclitaxel resistance through its interaction with integrin \u03b1V\u03b23 in cancer cells","volume":"16","author":"Gasca","year":"2020","journal-title":"Cell Death Discov."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"236","DOI":"10.1038\/nature11674","article-title":"COUP-TFII inhibits TGF-\u03b2-induced growth barrier to promote prostate tumorigenesis","volume":"493","author":"Qin","year":"2013","journal-title":"Nature"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1186\/s12967-020-02310-2","article-title":"Ltbp1 promotes esophageal squamous cell carcinoma progression through epithelial-mesenchymal transition and cancer-associated fibroblasts transformation","volume":"18","author":"Cai","year":"2020","journal-title":"J. Transl. Med."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"926","DOI":"10.1016\/j.ejca.2010.11.008","article-title":"TAZ\/WWTR1 is overexpressed in papillary thyroid carcinoma","volume":"47","author":"Ferraro","year":"2011","journal-title":"Eur. J. Cancer"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"667","DOI":"10.1038\/onc.2016.237","article-title":"Cadherin-6 promotes EMT and cancer metastasis by restraining autophagy","volume":"36","author":"Gugnoni","year":"2017","journal-title":"Oncogene"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"2342","DOI":"10.1016\/j.ajpath.2014.04.011","article-title":"The Roles of the Epithelial-Mesenchymal Transition Marker PRRX1 and miR-146b-5p in Papillary Thyroid Carcinoma Progression","volume":"184","author":"Hardin","year":"2014","journal-title":"Am. J. Pathol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1186\/s12935-019-0730-4","article-title":"SIRT6\/HIF-1\u03b1 axis promotes papillary thyroid cancer progression by inducing epithelial\u2013mesenchymal transition","volume":"19","author":"Yang","year":"2019","journal-title":"Cancer Cell Int."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1277","DOI":"10.1038\/sj.bjc.6602856","article-title":"S100A4 (Mts1) gene overexpression is associated with invasion and metastasis of papillary thyroid carcinoma","volume":"93","author":"Zou","year":"2005","journal-title":"Br. J. Cancer"},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Zheng, C., Wu, H., Zhang, S., Tian, R., Long, H., Zhang, H., Guo, X., Li, D., Tan, S., and Zhu, X. (2023). Myc-Associated Zinc Finger Protein Promotes Metastasis of Papillary Thyroid Cancer. Front. Biosci., 28.","DOI":"10.31083\/j.fbl2808162"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"MON-536","DOI":"10.1210\/jendso\/bvaa046.305","article-title":"MON-536 SPOCK1 Promotes the Progression of Papillary Thyroid Cancer ViaPI3K\/Akt Signaling Activation","volume":"4","author":"Li","year":"2020","journal-title":"J. Endocr. Soc."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"12079","DOI":"10.18632\/oncotarget.23951","article-title":"Integrated analysis of fine-needle-aspiration cystic fluid proteome, cancer cell secretome, and public transcriptome datasets for papillary thyroid cancer biomarker discovery","volume":"9","author":"Wu","year":"2018","journal-title":"Oncotarget"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1147","DOI":"10.2147\/IJGM.S346058","article-title":"Identification of a four-gene signature for determining the prognosis of papillary thyroid carcinoma by integrated bioinformatics analysis","volume":"15","author":"Luo","year":"2022","journal-title":"Int. J. Gen. Med."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1007\/s00109-011-0815-6","article-title":"S100-A10, thioredoxin, and S100-A6 as biomarkers of papillary thyroid carcinoma with lymph node metastasis identified by MALDI imaging","volume":"90","author":"Nipp","year":"2012","journal-title":"J. Mol. Med."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"515","DOI":"10.1007\/s10238-020-00642-3","article-title":"IGFBP3 as an indicator of lymph node metastasis and unfavorable prognosis for papillary thyroid carcinoma","volume":"20","author":"Huang","year":"2020","journal-title":"Clin. Exp. Med."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1016\/S0304-3835(02)00679-1","article-title":"Differential expression of IGFBP-5 and two human ESTs in thyroid glands with goiter, adenoma and papillary or follicular carcinomas","volume":"191","author":"Stolf","year":"2003","journal-title":"Cancer Lett."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1007\/s12079-015-0288-4","article-title":"Recent insights into the actions of IGFBP-6","volume":"9","author":"Bach","year":"2015","journal-title":"J. Cell Commun. Signal"},{"key":"ref_47","first-page":"1860","article-title":"Insulin-like growth factor binding protein-6 released from human mesenchymal stem cells confers neuronal protection through IGF-1R-mediated signaling","volume":"40","author":"Jeon","year":"2017","journal-title":"Int. J. Mol. Med."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"3835","DOI":"10.1038\/onc.2010.136","article-title":"IGFBP7: An oncosuppressor gene in thyroid carcinogenesis","volume":"29","author":"Vizioli","year":"2010","journal-title":"Oncogene"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1038\/s41420-023-01336-x","article-title":"Cancer-associated fibroblast-secreted IGFBP7 promotes gastric cancer by enhancing tumor associated macrophage infiltration via FGF2\/FGFR1\/PI3K\/AKT axis","volume":"9","author":"Li","year":"2023","journal-title":"Cell Death Discov."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Infante, A., and Rodr\u00edguez, C.I. (2018). Secretome analysis of in vitro aged human mesenchymal stem cells reveals IGFBP7 as a putative factor for promoting osteogenesis. Sci. Rep., 8.","DOI":"10.1038\/s41598-018-22855-z"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"2218","DOI":"10.1007\/s11427-021-2096-0","article-title":"Insulin-like growth factor binding proteins 7 prevents dental pulp-derived mesenchymal stem cell senescence via metabolic downregulation of p21","volume":"65","author":"Li","year":"2022","journal-title":"Sci. China Life Sci."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"36","DOI":"10.1038\/s41368-021-00146-0","article-title":"From bulk, single-cell to spatial RNA sequencing","volume":"13","author":"Li","year":"2021","journal-title":"Int. J. Oral. Sci."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1186\/s13287-018-0876-3","article-title":"Changes in phenotype and differentiation potential of human mesenchymal stem cells aging in vitro","volume":"9","author":"Yang","year":"2018","journal-title":"Stem Cell Res. Ther."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"273","DOI":"10.2217\/rme-2016-0009","article-title":"Human limbal mesenchymal stem cells express ABCB5 and can grow on amniotic membrane","volume":"11","author":"Shaharuddin","year":"2016","journal-title":"Regen. Med."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"748","DOI":"10.1038\/modpathol.2008.51","article-title":"S100A4 expression is associated with lymph node metastasis in papillary microcarcinoma of the thyroid","volume":"21","author":"Min","year":"2008","journal-title":"Mod. Pathol."},{"key":"ref_56","unstructured":"R Core Team (2022). R: A Language and Environment for Statistical Computing, R Foundation for Statistical Computing. Available online: https:\/\/www.R-project.org\/."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1158\/2159-8290.CD-12-0095","article-title":"The cBio cancer genomics portal: An open platform for exploring multidimensional cancer genomics data","volume":"2","author":"Cerami","year":"2012","journal-title":"Cancer Discov."},{"key":"ref_58","first-page":"pl1","article-title":"Integrative analysis of complex cancer genomics and clinical profiles using the cBioPortal","volume":"2","author":"Gao","year":"2013","journal-title":"Sci. Signal"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"3861","DOI":"10.1158\/0008-5472.CAN-23-0816","article-title":"Analysis and Visualization of Longitudinal Genomic and Clinical Data from the AACR Project GENIE Biopharma Collaborative in cBioPortal","volume":"83","author":"Kundra","year":"2023","journal-title":"Cancer Res."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"464","DOI":"10.1016\/j.ydbio.2004.08.048","article-title":"An integrated regulatory network controlling survival and migration in thyroid organogenesis","volume":"276","author":"Parlato","year":"2004","journal-title":"Dev. Biol."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/26\/10\/4957\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T17:36:45Z","timestamp":1760031405000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/26\/10\/4957"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,5,21]]},"references-count":60,"journal-issue":{"issue":"10","published-online":{"date-parts":[[2025,5]]}},"alternative-id":["ijms26104957"],"URL":"https:\/\/doi.org\/10.3390\/ijms26104957","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,5,21]]}}}