{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T12:28:08Z","timestamp":1786969688081,"version":"build-2736575974"},"reference-count":114,"publisher":"MDPI AG","issue":"16","license":[{"start":{"date-parts":[[2026,8,15]],"date-time":"2026-08-15T00:00:00Z","timestamp":1786752000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Int. J. Mol. Sci.","IJMS"],"abstract":"<jats:p>Gastrointestinal stromal tumors (GISTs) are the most common mesenchymal tumors of the gastrointestinal tract and are usually driven by activating mutations in KIT or PDGFRA. Although these alterations define the core molecular biology of GISTs and guide targeted therapy, they do not fully explain the variability observed in tumor behavior, recurrence risk, or response to tyrosine kinase inhibitors. Non-coding RNAs (ncRNAs), including microRNAs (miRNAs), long non-coding RNAs (lncRNAs), and circular RNAs (circRNAs), have been increasingly studied as regulators of gene expression in GISTs. Recent evidence suggests that these molecules may influence KIT-centered signaling, autophagy, apoptosis, invasion, angiogenesis, and drug resistance. However, the strength of evidence differs considerably across individual ncRNAs, ranging from bioinformatic associations to functional validation in cell lines and in vivo models. This review summarizes current knowledge on ncRNA-mediated regulation in GIST biology, with emphasis on tumor progression, therapeutic resistance, and possible clinical relevance. Rather than treating ncRNAs as isolated biomarkers, they function as part of broader regulatory networks that interact with oncogenic signaling and epigenetic mechanisms. Although several ncRNAs appear promising as prognostic or predictive candidates, further validation in independent clinical cohorts is required before their integration into routine risk stratification or treatment decision-making.<\/jats:p>","DOI":"10.3390\/ijms27167286","type":"journal-article","created":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T12:01:26Z","timestamp":1786968086000},"page":"7286","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Non-Coding RNAs in Gastrointestinal Stromal Tumors: Regulatory Networks, Drug Resistance, and Clinical Implications"],"prefix":"10.3390","volume":"27","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2974-4084","authenticated-orcid":false,"given":"Georgios","family":"Mandrakis","sequence":"first","affiliation":[{"name":"First Department of Pathology, School of Medicine, National and Kapodistrian University of Athens, 10679 Athens, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1583-1125","authenticated-orcid":false,"given":"Stavros P.","family":"Papadakos","sequence":"additional","affiliation":[{"name":"First Department of Pathology, School of Medicine, National and Kapodistrian University of Athens, 10679 Athens, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1398-178X","authenticated-orcid":false,"given":"Georgia","family":"Levidou","sequence":"additional","affiliation":[{"name":"Department of Pathology, Paracelsus Medical University, 90419 Nuremberg, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-6032-5251","authenticated-orcid":false,"given":"Panoraia","family":"Keratsa","sequence":"additional","affiliation":[{"name":"First Department of Pathology, School of Medicine, National and Kapodistrian University of Athens, 10679 Athens, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1659-4993","authenticated-orcid":false,"given":"Maria-Ioanna","family":"Christodoulou","sequence":"additional","affiliation":[{"name":"Tumor Immunology and Biomarkers Laboratory, Basic and Translational Cancer Research Center, Department of Life Sciences, European University Cyprus, Nicosia 2404, Cyprus"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stamatios","family":"Theocharis","sequence":"additional","affiliation":[{"name":"First Department of Pathology, School of Medicine, National and Kapodistrian University of Athens, 10679 Athens, Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,15]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"144","DOI":"10.21037\/jgo.2018.08.20","article-title":"Gastrointestinal stromal tumors: A comprehensive review","volume":"10","author":"Parab","year":"2019","journal-title":"J. Gastrointest. Oncol."},{"key":"ref_2","first-page":"31","article-title":"Gastrointestinal stromal tumours: Outcomes of surgical management and analysis of prognostic variables","volume":"52","author":"Unalp","year":"2009","journal-title":"Can. J. Surg."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1469","DOI":"10.4251\/wjgo.v14.i8.1469","article-title":"Angiogenesis in gastrointestinal stromal tumors: From bench to bedside","volume":"14","author":"Papadakos","year":"2022","journal-title":"World J. Gastrointest. Oncol."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Vallilas, C., Sarantis, P., Kyriazoglou, A., Koustas, E., Theocharis, S., Papavassiliou, A.G., and Karamouzis, M.V. (2021). Gastrointestinal Stromal Tumors (GISTs): Novel Therapeutic Strategies with Immunotherapy and Small Molecules. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22020493"},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Khan, J., Ullah, A., Waheed, A., Karki, N.R., Adhikari, N., Vemavarapu, L., Belakhlef, S., Bendjemil, S.M., Mehdizadeh Seraj, S., and Sidhwa, F. (2022). Gastrointestinal Stromal Tumors (GIST): A Population-Based Study Using the SEER Database, Including Management and Recent Advances in Targeted Therapy. Cancers, 14.","DOI":"10.3390\/cancers14153689"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"150","DOI":"10.1007\/s10238-025-01667-2","article-title":"Regulatory roles of non-coding RNAs in programmed cell death pathways and drug resistance in gastrointestinal stromal tumors","volume":"25","author":"Ma","year":"2025","journal-title":"Clin. Exp. Med."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1007\/978-3-319-23730-5_4","article-title":"MicroRNAs and Gastrointestinal Stromal Tumor","volume":"889","author":"Akcakaya","year":"2015","journal-title":"Adv. Exp. Med. Biol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1007\/s00428-010-0891-y","article-title":"Gastrointestinal stromal tumors","volume":"456","author":"Fletcher","year":"2010","journal-title":"Virchows Arch."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1097\/PAP.0000000000000158","article-title":"What is New in Gastrointestinal Stromal Tumor?","volume":"24","author":"Schaefer","year":"2017","journal-title":"Adv. Anat. Pathol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"849","DOI":"10.1038\/nature09409","article-title":"ETV1 is a lineage survival factor that cooperates with KIT in gastrointestinal stromal tumours","volume":"467","author":"Chi","year":"2010","journal-title":"Nature"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1055\/s-0044-1789204","article-title":"Gene Mutations in Gastrointestinal Stromal Tumors: Advances in Treatment and Mechanism Research","volume":"11","author":"Cao","year":"2024","journal-title":"Glob. Med. Genet."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"514","DOI":"10.1016\/j.biocel.2014.05.033","article-title":"Succinate dehydrogenase deficient gastrointestinal stromal tumors (GISTs)\u2014A review","volume":"53","author":"Miettinen","year":"2014","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"922","DOI":"10.1001\/jamaoncol.2016.0256","article-title":"Molecular Subtypes of KIT\/PDGFRA Wild-Type Gastrointestinal Stromal Tumors: A Report from the National Institutes of Health Gastrointestinal Stromal Tumor Clinic","volume":"2","author":"Boikos","year":"2016","journal-title":"JAMA Oncol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1158\/1541-7786.MCR-16-0376","article-title":"Genome-Wide Analysis Identifies MEN1 and MAX Mutations and a Neuroendocrine-Like Molecular Heterogeneity in Quadruple WT GIST","volume":"15","author":"Pantaleo","year":"2017","journal-title":"Mol. Cancer Res."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"853","DOI":"10.1002\/gcc.20589","article-title":"Novel V600E BRAF mutations in imatinib-naive and imatinib-resistant gastrointestinal stromal tumors","volume":"47","author":"Agaram","year":"2008","journal-title":"Genes Chromosom. Cancer"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1136\/jcp.2009.064550","article-title":"V600E BRAF mutations are alternative early molecular events in a subset of KIT\/PDGFRA wild-type gastrointestinal stromal tumours","volume":"62","author":"Agaimy","year":"2009","journal-title":"J. Clin. Pathol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1547","DOI":"10.1097\/PAS.0000000000000911","article-title":"Pan-Trk Immunohistochemistry Is an Efficient and Reliable Screen for the Detection of NTRK Fusions","volume":"41","author":"Hechtman","year":"2017","journal-title":"Am. J. Surg. Pathol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"15","DOI":"10.4291\/wjgp.v13.i1.15","article-title":"Current treatment strategies and future perspectives for gastrointestinal stromal tumors","volume":"13","author":"Sugiyama","year":"2022","journal-title":"World J. Gastrointest. Pathophysiol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"17588359231192388","DOI":"10.1177\/17588359231192388","article-title":"2023 GEIS Guidelines for gastrointestinal stromal tumors","volume":"15","author":"Serrano","year":"2023","journal-title":"Ther. Adv. Med. Oncol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"104314","DOI":"10.5306\/wjco.v16.i6.104314","article-title":"Immunotherapy in gastrointestinal stromal tumors: Current landscape and future horizons","volume":"16","author":"Papadakos","year":"2025","journal-title":"World J. Clin. Oncol."},{"key":"ref_21","first-page":"14","article-title":"The current state of chemotherapy for the treatment of gastrointestinal stromal tumors with different genotypes: A narrative review","volume":"05","author":"Xu","year":"2021","journal-title":"J. Bio-X Res."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"612","DOI":"10.1038\/s41416-019-0389-6","article-title":"Complementary activity of tyrosine kinase inhibitors against secondary kit mutations in imatinib-resistant gastrointestinal stromal tumours","volume":"120","author":"Serrano","year":"2019","journal-title":"Br. J. Cancer"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"4182","DOI":"10.1158\/1078-0432.CCR-04-2245","article-title":"Acquired resistance to imatinib in gastrointestinal stromal tumor occurs through secondary gene mutation","volume":"11","author":"Antonescu","year":"2005","journal-title":"Clin. Cancer Res."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"4764","DOI":"10.1200\/JCO.2006.06.2265","article-title":"Molecular correlates of imatinib resistance in gastrointestinal stromal tumors","volume":"24","author":"Heinrich","year":"2006","journal-title":"J. Clin. Oncol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"64","DOI":"10.1002\/path.2382","article-title":"Heterogeneity of kinase inhibitor resistance mechanisms in GIST","volume":"216","author":"Liegl","year":"2008","journal-title":"J. Pathol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"189","DOI":"10.3892\/ol.2022.13309","article-title":"Ki67 for evaluating the prognosis of gastrointestinal stromal tumors: A systematic review and meta-analysis","volume":"23","author":"Li","year":"2022","journal-title":"Oncol. Lett."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"e8399","DOI":"10.1590\/1414-431x20198399","article-title":"Downregulation of lncRNA CCDC26 contributes to imatinib resistance in human gastrointestinal stromal tumors through IGF-1R upregulation","volume":"52","author":"Yan","year":"2019","journal-title":"Braz. J. Med. Biol. Res."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"8387","DOI":"10.1073\/pnas.0803383105","article-title":"Insulin-like growth factor 1 receptor is a potential therapeutic target for gastrointestinal stromal tumors","volume":"105","author":"Tarn","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1186\/s12964-023-01411-x","article-title":"KIT mutations and expression: Current knowledge and new insights for overcoming IM resistance in GIST","volume":"22","author":"Zhou","year":"2024","journal-title":"Cell Commun. Signal"},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Boichuk, S., Dunaev, P., Galembikova, A., and Valeeva, E. (2024). Fibroblast Growth Factor 2 (FGF2) Activates Vascular Endothelial Growth Factor (VEGF) Signaling in Gastrointestinal Stromal Tumors (GIST): An Autocrine Mechanism Contributing to Imatinib Mesylate (IM) Resistance. Cancers, 16.","DOI":"10.3390\/cancers16173103"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"959","DOI":"10.1002\/ijc.26095","article-title":"Long-term exposure of gastrointestinal stromal tumor cells to sunitinib induces epigenetic silencing of the PTEN gene","volume":"130","author":"Yang","year":"2012","journal-title":"Int. J. Cancer"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"e79655","DOI":"10.7554\/eLife.79655","article-title":"Non-coding RNAs in drug and radiation resistance of bone and soft-tissue sarcoma: A systematic review","volume":"11","author":"Chen","year":"2022","journal-title":"elife"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"252","DOI":"10.2174\/1566524018666181004113436","article-title":"Coding and Non-coding: Molecular Portrait of GIST and its Clinical Implication","volume":"18","author":"Bure","year":"2018","journal-title":"Curr. Mol. Med."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"118502","DOI":"10.3748\/wjg.v32.i17.118502","article-title":"Epigenetic landscape of gastrointestinal stromal tumors: Mechanistic insights and therapeutic opportunities","volume":"32","author":"Papadakos","year":"2026","journal-title":"World J. Gastroenterol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"609","DOI":"10.1038\/s41422-024-00975-8","article-title":"Coding, or non-coding, that is the question","volume":"34","author":"Poliseno","year":"2024","journal-title":"Cell Res."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1151","DOI":"10.1002\/pmic.200500372","article-title":"Differentially expressed proteins in gastrointestinal stromal tumors with KIT and PDGFRA mutations","volume":"6","author":"Kang","year":"2006","journal-title":"Proteomics"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"829","DOI":"10.1177\/0300985820960132","article-title":"Validating Immunohistochemistry Assay Specificity in Investigative Studies: Considerations for a Weight of Evidence Approach","volume":"58","author":"Webster","year":"2021","journal-title":"Vet. Pathol."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Xi, X., Li, T., Huang, Y., Sun, J., Zhu, Y., Yang, Y., and Lu, Z.J. (2017). RNA Biomarkers: Frontier of Precision Medicine for Cancer. Non-Coding RNA, 3.","DOI":"10.3390\/ncrna3010009"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"627","DOI":"10.1038\/s41576-021-00370-8","article-title":"Integrating single-cell and spatial transcriptomics to elucidate intercellular tissue dynamics","volume":"22","author":"Longo","year":"2021","journal-title":"Nat. Rev. Genet."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"190","DOI":"10.1038\/s41419-024-06571-3","article-title":"Deciphering the tumor immune microenvironment of imatinib-resistance in advanced gastrointestinal stromal tumors at single-cell resolution","volume":"15","author":"Liu","year":"2024","journal-title":"Cell Death Dis."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"430","DOI":"10.1038\/s41580-022-00566-8","article-title":"Long non-coding RNAs: Definitions, functions, challenges and recommendations","volume":"24","author":"Mattick","year":"2023","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"D155","DOI":"10.1093\/nar\/gky1141","article-title":"miRBase: From microRNA sequences to function","volume":"47","author":"Kozomara","year":"2019","journal-title":"Nucleic Acids Res."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"869","DOI":"10.1016\/j.molcel.2013.05.007","article-title":"Alternative splicing regulates biogenesis of miRNAs located across exon-intron junctions","volume":"50","author":"Melamed","year":"2013","journal-title":"Mol. Cell"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1016\/j.cell.2009.01.002","article-title":"MicroRNAs: Target recognition and regulatory functions","volume":"136","author":"Bartel","year":"2009","journal-title":"Cell"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1157","DOI":"10.1038\/cr.2008.311","article-title":"miRNAs: Small changes, widespread effects","volume":"18","author":"Linsen","year":"2008","journal-title":"Cell Res."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1038\/nrm3838","article-title":"Regulation of microRNA biogenesis","volume":"15","author":"Ha","year":"2014","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1038\/s41580-018-0059-1","article-title":"Regulation of microRNA biogenesis and its crosstalk with other cellular pathways","volume":"20","author":"Treiber","year":"2019","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"189","DOI":"10.4161\/bioa.22592","article-title":"Sizing up long non-coding RNAs: Do lncRNAs have secondary and tertiary structure?","volume":"2","author":"Novikova","year":"2012","journal-title":"Bioarchitecture"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1016\/j.cell.2011.07.014","article-title":"A ceRNA hypothesis: The Rosetta Stone of a hidden RNA language?","volume":"146","author":"Salmena","year":"2011","journal-title":"Cell"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"111389","DOI":"10.1016\/j.biopha.2021.111389","article-title":"The role of a ceRNA regulatory network based on lncRNA MALAT1 site in cancer progression","volume":"137","author":"Su","year":"2021","journal-title":"Biomed. Pharmacother."},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"O\u2019Brien, J., Hayder, H., Zayed, Y., and Peng, C. (2018). Overview of MicroRNA Biogenesis, Mechanisms of Actions, and Circulation. Front. Endocrinol., 9.","DOI":"10.3389\/fendo.2018.00402"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"710","DOI":"10.1136\/jmedgenet-2015-103334","article-title":"ceRNA in cancer: Possible functions and clinical implications","volume":"52","author":"Qi","year":"2015","journal-title":"J. Med. Genet."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1261\/rna.035667.112","article-title":"Circular RNAs are abundant, conserved, and associated with ALU repeats","volume":"19","author":"Jeck","year":"2013","journal-title":"RNA"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"384","DOI":"10.1038\/nature11993","article-title":"Natural RNA circles function as efficient microRNA sponges","volume":"495","author":"Hansen","year":"2013","journal-title":"Nature"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"256","DOI":"10.1038\/nsmb.2959","article-title":"Exon-intron circular RNAs regulate transcription in the nucleus","volume":"22","author":"Li","year":"2015","journal-title":"Nat. Struct. Mol. Biol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"106781","DOI":"10.1016\/j.isci.2023.106781","article-title":"Microproteins: Overlooked regulators of physiology and disease","volume":"26","author":"Hassel","year":"2023","journal-title":"iScience"},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Gyvyte, U., Kupcinskas, J., Juzenas, S., Inciuraite, R., Poskiene, L., Salteniene, V., Link, A., Fassan, M., Franke, A., and Kupcinskas, L. (2018). Identification of long intergenic non-coding RNAs (lincRNAs) deregulated in gastrointestinal stromal tumors (GISTs). PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0209342"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"e70089","DOI":"10.1111\/cpr.70089","article-title":"LncRNA H19-Encoded Micropeptide altH19 Promotes DNA Replication and Mitosis in Myeloma Cells by Enhancing the Phosphorylation of CDK2 at Threonine 160","volume":"59","author":"Zhang","year":"2026","journal-title":"Cell Prolif."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1080\/26895293.2021.1889685","article-title":"LncRNA PVT1 promotes cells proliferation via PI3K\u2013AKT\u2013mTOR Pathway in gastrointestinal stromal tumor","volume":"14","author":"Bi","year":"2021","journal-title":"All Life"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1342","DOI":"10.1158\/2326-6066.CIR-20-0964","article-title":"CD8+ T-cell Immune Surveillance against a Tumor Antigen Encoded by the Oncogenic Long Noncoding RNA PVT1","volume":"9","author":"Kikuchi","year":"2021","journal-title":"Cancer Immunol. Res."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1186\/s12943-019-1010-6","article-title":"A novel protein encoded by a circular RNA circPPP1R12A promotes tumor pathogenesis and metastasis of colon cancer via Hippo-YAP signaling","volume":"18","author":"Zheng","year":"2019","journal-title":"Mol. Cancer"},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Isosaka, M., Niinuma, T., Nojima, M., Kai, M., Yamamoto, E., Maruyama, R., Nobuoka, T., Nishida, T., Kanda, T., and Taguchi, T. (2015). A Screen for Epigenetically Silenced microRNA Genes in Gastrointestinal Stromal Tumors. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0133754"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"1519","DOI":"10.1007\/s13258-022-01301-5","article-title":"Methyltransferase-like 3 (METTL3) mediated N6-methyladenosine (m6A) modifications facilitate mir-25-3p maturation to promote gastrointestinal stromal tumors (GISTs) progression","volume":"44","author":"Qian","year":"2022","journal-title":"Genes Genom."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"113131","DOI":"10.1016\/j.yexcr.2022.113131","article-title":"CTCF-activated SNHG16 facilitates gastrointestinal stromal tumor by targeting miR-128-3p\/CASC3 axis","volume":"417","author":"Yang","year":"2022","journal-title":"Exp. Cell Res."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"e70047","DOI":"10.1002\/mog2.70047","article-title":"Decreased M6A Modification Promotes Progression of Gastrointestinal Stromal Tumor and KIT-Derived Imatinib Resistance Through FTO-Regulated Axis","volume":"4","author":"Li","year":"2025","journal-title":"MedComm\u2013Oncology"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"2780","DOI":"10.1002\/elps.201600258","article-title":"High-throughput miRNA sequencing and identification of biomarkers for forensically relevant biological fluids","volume":"37","author":"Lewis","year":"2016","journal-title":"Electrophoresis"},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Gyvyte, U., Lukosevicius, R., Inciuraite, R., Streleckiene, G., Gudoityte, G., Bekampyte, J., Valentini, S., Salteniene, V., Ruzgys, P., and Satkauskas, S. (2020). The Role of miR-375-3p and miR-200b-3p in Gastrointestinal Stromal Tumors. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21145151"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1421","DOI":"10.1016\/j.molonc.2015.03.013","article-title":"miRNA-221 and miRNA-222 induce apoptosis via the KIT\/AKT signalling pathway in gastrointestinal stromal tumours","volume":"9","author":"Ihle","year":"2015","journal-title":"Mol. Oncol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"7584","DOI":"10.1158\/1078-0432.CCR-11-0166","article-title":"MicroRNA-494 downregulates KIT and inhibits gastrointestinal stromal tumor cell proliferation","volume":"17","author":"Kim","year":"2011","journal-title":"Clin. Cancer Res."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"2091","DOI":"10.1038\/bjc.2014.548","article-title":"microRNA expression signatures of gastrointestinal stromal tumours: Associations with imatinib resistance and patient outcome","volume":"111","author":"Akcakaya","year":"2014","journal-title":"Br. J. Cancer"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1186\/s12964-018-0228-z","article-title":"miR-148b-3p functions as a tumor suppressor in GISTs by directly targeting KIT","volume":"16","author":"Wang","year":"2018","journal-title":"Cell Commun. Signal"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"198","DOI":"10.1038\/s41419-020-2390-7","article-title":"MicroRNA-30a targets BECLIN-1 to inactivate autophagy and sensitizes gastrointestinal stromal tumor cells to imatinib","volume":"11","author":"Chen","year":"2020","journal-title":"Cell Death Dis."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1038\/s41419-021-03650-7","article-title":"LncRNA-HOTAIR activates autophagy and promotes the imatinib resistance of gastrointestinal stromal tumor cells through a mechanism involving the miR-130a\/ATG2B pathway","volume":"12","author":"Zhang","year":"2021","journal-title":"Cell Death Dis."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"652","DOI":"10.1038\/labinvest.2015.157","article-title":"PIK3C2A is a gene-specific target of microRNA-518a-5p in imatinib mesylate-resistant gastrointestinal stromal tumor","volume":"96","author":"Shi","year":"2016","journal-title":"Lab. Investig."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"5711","DOI":"10.1002\/jcp.29506","article-title":"miR-4510 acts as a tumor suppressor in gastrointestinal stromal tumor by targeting APOC2","volume":"235","author":"Chen","year":"2020","journal-title":"J. Cell. Physiol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"4209","DOI":"10.1007\/s13277-013-1551-z","article-title":"MicroRNA-218 inhibits gastrointestinal stromal tumor cell and invasion by targeting KIT","volume":"35","author":"Fan","year":"2014","journal-title":"Tumor Biol."},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Jia, N., Tong, H., Zhang, Y., Katayama, H., Wang, Y., Lu, W., Zhang, S., and Wang, J. (2019). CeRNA Expression Profiling Identifies KIT-Related circRNA-miRNA-mRNA Networks in Gastrointestinal Stromal Tumour. Front. Genet., 10.","DOI":"10.3389\/fgene.2019.00825"},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Fernandez-Serra, A., Moura, D.S., Sanchez-Izquierdo, M.D., Calabuig-Farinas, S., Lopez-Alvarez, M., Martinez-Martinez, A., Carrasco-Garcia, I., Ramirez-Calvo, M., Blanco-Alcaina, E., and Lopez-Reig, R. (2020). Prognostic Impact of let-7e MicroRNA and Its Target Genes in Localized High-Risk Intestinal GIST: A Spanish Group for Research on Sarcoma (GEIS) Study. Cancers, 12.","DOI":"10.3390\/cancers12102979"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"9131","DOI":"10.1007\/s13277-014-2177-5","article-title":"MiR-137 regulates epithelial-mesenchymal transition in gastrointestinal stromal tumor","volume":"35","author":"Liu","year":"2014","journal-title":"Tumor Biol."},{"key":"ref_80","first-page":"5703","article-title":"Downregulation of miR-186 is associated with metastatic recurrence of gastrointestinal stromal tumors","volume":"14","author":"Niinuma","year":"2017","journal-title":"Oncol. Lett."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"100423","DOI":"10.1016\/j.clinsp.2024.100423","article-title":"CircRNA amyloid precursor protein by competitive adsorption of microRNA-6838-5p mediates CDV3 expression to enhance malignant behavior and Warburg effect in Gastrointestinal Stromal Tumor","volume":"79","author":"Zhaorigetu","year":"2024","journal-title":"Clinics"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"935","DOI":"10.1007\/s11596-023-2715-8","article-title":"MicroRNA-409-5p Inhibits GIST Tumorigenesis and Improves Imatinib Resistance by Targeting KDM4D Expression","volume":"43","author":"Qiu","year":"2023","journal-title":"Curr. Med. Sci."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"683","DOI":"10.4149\/neo_2018_170906N575","article-title":"Identification of critical microRNAs in gastrointestinal stromal tumor patients treated with Imatinib","volume":"65","author":"Zhang","year":"2018","journal-title":"Neoplasma"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"563","DOI":"10.1038\/modpathol.2012.198","article-title":"Fascin-1 overexpression and miR-133b downregulation in the progression of gastrointestinal stromal tumor","volume":"26","author":"Yamamoto","year":"2013","journal-title":"Mod. Pathol."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"1126","DOI":"10.1158\/0008-5472.CAN-11-1803","article-title":"Upregulation of miR-196a and HOTAIR drive malignant character in gastrointestinal stromal tumors","volume":"72","author":"Niinuma","year":"2012","journal-title":"Cancer Res."},{"key":"ref_86","first-page":"3705","article-title":"miR-182 controls cell growth in gastrointestinal stromal tumors by negatively regulating CYLD expression","volume":"40","author":"Ling","year":"2018","journal-title":"Oncol. Rep."},{"key":"ref_87","first-page":"532","article-title":"MiR-374b Promotes Proliferation and Inhibits Apoptosis of Human GIST Cells by Inhibiting PTEN Through Activation of the PI3K\/Akt Pathway","volume":"41","author":"Long","year":"2018","journal-title":"Mol. Cells"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"1347","DOI":"10.2217\/epi-2016-0030","article-title":"Integrating miRNA and gene expression profiling analysis revealed regulatory networks in gastrointestinal stromal tumors","volume":"8","author":"Pantaleo","year":"2016","journal-title":"Epigenomics"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"669","DOI":"10.1038\/s41419-021-03942-y","article-title":"Integrated analysis of long non-coding RNAs and mRNAs associated with malignant transformation of gastrointestinal stromal tumors","volume":"12","author":"Yin","year":"2021","journal-title":"Cell Death Dis."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"584","DOI":"10.1002\/gcc.22672","article-title":"Long noncoding RNA HOTAIR is upregulated in an aggressive subgroup of gastrointestinal stromal tumors (GIST) and mediates the establishment of gene-specific DNA methylation patterns","volume":"57","author":"Bure","year":"2018","journal-title":"Genes Chromosom. Cancer"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"526","DOI":"10.4149\/neo_2023_230523N272","article-title":"LncRNA HIF1A-AS2 mediates imatinib resistance by regulating autophagy in gastrointestinal stromal tumor cells","volume":"70","author":"Yan","year":"2023","journal-title":"Neoplasma"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1016\/j.omtn.2021.05.017","article-title":"RP11-616M22.7 recapitulates imatinib resistance in gastrointestinal stromal tumor","volume":"25","author":"Shao","year":"2021","journal-title":"Mol. Ther. Nucleic Acids"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"2522543","DOI":"10.1080\/15384047.2025.2522543","article-title":"Highly enriched exosomal lncRNA OIP5-AS1 regulates gastrointestinal stromal tumor (GIST) resistance to sunitinib through miR-145 and SOX9","volume":"26","author":"Wang","year":"2025","journal-title":"Cancer Biol. Ther."},{"key":"ref_94","first-page":"1026","article-title":"Long noncoding RNA SPRY4-IT1 acts as a miR-101-5p sponge to promote gastrointestinal stromal tumor progression by inhibiting ZEB1","volume":"15","author":"Huang","year":"2023","journal-title":"Am. J. Transl. Res."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1007\/s13577-024-01167-7","article-title":"Long non-coding RNA MSC-AS1 confers imatinib resistance of gastrointestinal stromal tumor cells by activating FNDC1 and ANLN-mediated PI3K\/AKT pathway","volume":"38","author":"Chen","year":"2025","journal-title":"Hum. Cell"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"661","DOI":"10.1002\/mc.23199","article-title":"PCAT6 mediates cellular biological functions in gastrointestinal stromal tumor via upregulation of PRDX5 and activation of Wnt pathway","volume":"59","author":"Bai","year":"2020","journal-title":"Mol. Carcinog."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"e41934","DOI":"10.1016\/j.heliyon.2025.e41934","article-title":"LINC00870 promotes imatinib resistance in gastrointestinal stromal tumor via inhibiting PIGR glycosylation modifications","volume":"11","author":"Li","year":"2025","journal-title":"Heliyon"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"2817","DOI":"10.21873\/anticanres.15062","article-title":"An Integrated Epigenome and Transcriptome Analysis to Clarify the Effect of Epigenetic Inhibitors on GIST","volume":"41","author":"Niinuma","year":"2021","journal-title":"Anticancer Res."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1186\/s13000-019-0877-2","article-title":"LncRNA DNM3OS promotes proliferation and inhibits apoptosis through modulating IGF1 expression by sponging MiR-126 in CHON-001 cells","volume":"14","author":"Ai","year":"2019","journal-title":"Diagn. Pathol."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"75307","DOI":"10.18632\/oncotarget.12171","article-title":"Promoter methylation of PCDH10 by HOTAIR regulates the progression of gastrointestinal stromal tumors","volume":"7","author":"Lee","year":"2016","journal-title":"Oncotarget"},{"key":"ref_101","doi-asserted-by":"crossref","unstructured":"Zou, F.-W., Cao, D., Tang, Y.-F., Shu, L., Zuo, Z., and Zhang, L.-Y. (2020). Identification of CircRNA\u2013miRNA\u2013mRNA Regulatory Network in Gastrointestinal Stromal Tumor. Front. Genet., 11.","DOI":"10.3389\/fgene.2020.00403"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"1338","DOI":"10.4103\/jcrt.jcrt_625_22","article-title":"Circ-CCS enhances autophagy during imatinib resistance of gastrointestinal stromal tumor by regulating miR-197-3p\/ATG10 signaling","volume":"18","author":"Sui","year":"2022","journal-title":"J. Cancer Res. Ther."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"22004","DOI":"10.1038\/s41598-024-73393-w","article-title":"circ_SMA4 promotes gastrointestinal stromal tumors malignant progression by sponging miR-494-3p\/KIT axis and activating JAK\/STAT pathway","volume":"14","author":"Zou","year":"2024","journal-title":"Sci. Rep."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"601","DOI":"10.1186\/s12967-025-06598-w","article-title":"RNA profiling and immunohistochemistry analyses of circRNAs in imatinib-resistant gastrointestinal stromal tumors","volume":"23","author":"Tong","year":"2025","journal-title":"J. Transl. Med."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1093\/abbs\/gmt118","article-title":"MiR-320a downregulation is associated with imatinib resistance in gastrointestinal stromal tumors","volume":"46","author":"Gao","year":"2014","journal-title":"Acta Biochim. Biophys. Sin."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"1015","DOI":"10.1007\/s12038-018-9805-y","article-title":"Serum miR-518e-5p is a potential biomarker for secondary imatinib-resistant gastrointestinal stromal tumor","volume":"43","author":"Kou","year":"2018","journal-title":"J. Biosci."},{"key":"ref_107","doi-asserted-by":"crossref","unstructured":"Huang, W.K., Shi, H., Akcakaya, P., Zeljic, K., Gangaev, A., Caramuta, S., Yeh, C.N., Branstrom, R., Larsson, C., and Lui, W.O. (2021). Imatinib Regulates miR-483-3p and Mitochondrial Respiratory Complexes in Gastrointestinal Stromal Tumors. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms221910600"},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1038\/s41580-020-00324-8","article-title":"Ferroptosis: Mechanisms, biology and role in disease","volume":"22","author":"Jiang","year":"2021","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"839","DOI":"10.1038\/s41419-023-06300-2","article-title":"Imatinib induces ferroptosis in gastrointestinal stromal tumors by promoting STUB1-mediated GPX4 ubiquitination","volume":"14","author":"Sun","year":"2023","journal-title":"Cell Death Dis."},{"key":"ref_110","doi-asserted-by":"crossref","unstructured":"Delvaux, M., Hague, P., Craciun, L., Wozniak, A., Demetter, P., Schoffski, P., Erneux, C., and Vanderwinden, J.M. (2022). Ferroptosis Induction and YAP Inhibition as New Therapeutic Targets in Gastrointestinal Stromal Tumors (GISTs). Cancers, 14.","DOI":"10.3390\/cancers14205050"},{"key":"ref_111","doi-asserted-by":"crossref","unstructured":"Zheng, X., and Zhang, C. (2023). The Regulation of Ferroptosis by Noncoding RNAs. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms241713336"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1038\/s41573-021-00219-z","article-title":"Noncoding RNA therapeutics\u2014Challenges and potential solutions","volume":"20","author":"Winkle","year":"2021","journal-title":"Nat. Rev. Drug Discov."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"1630","DOI":"10.1038\/s41416-020-0802-1","article-title":"Phase 1 study of MRX34, a liposomal miR-34a mimic, in patients with advanced solid tumours","volume":"122","author":"Hong","year":"2020","journal-title":"Br. J. Cancer"},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"1386","DOI":"10.1016\/S1470-2045(17)30621-6","article-title":"Safety and activity of microRNA-loaded minicells in patients with recurrent malignant pleural mesothelioma: A first-in-man, phase 1, open-label, dose-escalation study","volume":"18","author":"Pavlakis","year":"2017","journal-title":"Lancet Oncol."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/16\/7286\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T12:04:11Z","timestamp":1786968251000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/16\/7286"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,15]]},"references-count":114,"journal-issue":{"issue":"16","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["ijms27167286"],"URL":"https:\/\/doi.org\/10.3390\/ijms27167286","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,15]]}}}