{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T05:02:23Z","timestamp":1789621343120,"version":"build-2803163510"},"reference-count":66,"publisher":"MDPI AG","issue":"18","license":[{"start":{"date-parts":[[2026,9,13]],"date-time":"2026-09-13T00:00:00Z","timestamp":1789257600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Ministry of Science and Higher Education of the Russian Federation","award":["075-15-2024-536"],"award-info":[{"award-number":["075-15-2024-536"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Int. J. Mol. Sci.","IJMS"],"abstract":"<jats:p>MicroRNAs are frequently dysregulated in cancer, yet how their tissue specificity is remodeled during malignant transformation remains poorly characterized. Here, we systematically quantified the tissue specificity of miRNAs across normal (GTEx) and cancer (TCGA) tissues using the Tau index as the primary metric and the Gini index for independent validation. Our analysis revealed a negative association between tissue specificity in healthy tissues and changes in specificity during malignant transformation, indicating that miRNAs with higher tissue specificity in normal tissues tend to undergo greater loss of specificity in cancer. To robustly define dysregulation, we combined two independent analyses: a binomial test over per-project differential expression across 17 matched normal tissues within the TCGA cohort, and a TCGA\u2013GTEx pan-tissue comparison of mean expression. The change in specificity (\u0394Tau) separated up- from downregulated miRNAs, showing moderate agreement with the binomial signal and a strong correlation with the expression-based contrast. Finally, we identified six miRNAs that lose tissue specificity upon transformation while remaining consistently upregulated (miR-519a-5p, miR-512-3p, miR-522-3p, miR-105-5p, miR-935, miR-1269a). Functional analysis of their experimentally validated and predicted targets showed significant enrichment for converging on core oncogenic programs. Collectively, integrating specificity dynamics with dysregulation evidence pinpoints candidate miRNAs with coordinated, cancer-relevant regulatory roles and highlights those with favorable tissue-specificity profiles that may warrant further investigation as potential therapeutic candidates.<\/jats:p>","DOI":"10.3390\/ijms27188153","type":"journal-article","created":{"date-parts":[[2026,9,14]],"date-time":"2026-09-14T12:09:51Z","timestamp":1789387791000},"page":"8153","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["A Pan-Cancer Analysis of microRNA Tissue Specificity and Its Association with Dysregulation"],"prefix":"10.3390","volume":"27","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4506-0256","authenticated-orcid":false,"given":"Aly","family":"Ismailov","sequence":"first","affiliation":[{"name":"Centre for Biomedical Research and Technology, HSE University, 109028 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"conceptualization","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2033-9621","authenticated-orcid":false,"suffix":"Jr.","given":"Alexey","family":"Belogurov","sequence":"additional","affiliation":[{"name":"Laboratory of Hormonal Regulation Proteins, Shemyakin and Ovchinnikov Institute of Bioorganic Chemistry Russian Academy of Sciences (RAS), 117997 Moscow, Russia"},{"name":"Department of Biological Chemistry, Russian University of Medicine, Ministry of Health of Russian Federation, 127473 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alena","family":"Evpak","sequence":"additional","affiliation":[{"name":"Laboratory of Hormonal Regulation Proteins, Shemyakin and Ovchinnikov Institute of Bioorganic Chemistry Russian Academy of Sciences (RAS), 117997 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7198-8234","authenticated-orcid":false,"given":"Maria","family":"Poptsova","sequence":"additional","affiliation":[{"name":"Centre for Biomedical Research and Technology, HSE University, 109028 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"conceptualization","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,13]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"2628","DOI":"10.7150\/ijbs.47203","article-title":"miRNA-based biomarkers, therapies, and resistance in Cancer","volume":"16","author":"He","year":"2020","journal-title":"Int. J. Biol. Sci."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"3687","DOI":"10.1158\/0008-5472.CAN-22-0240","article-title":"Pan-Cancer Analysis of Canonical and Modified miRNAs Enhances the Resolution of the Functional miRNAome in Cancer","volume":"82","author":"Distefano","year":"2022","journal-title":"Cancer Res."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1016\/j.compbiomed.2018.10.021","article-title":"Identification of key differentially expressed MicroRNAs in cancer patients through pan-cancer analysis","volume":"103","author":"Hu","year":"2018","journal-title":"Comput Biol. Med."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Navon, R., Wang, H., Steinfeld, I., Tsalenko, A., Ben-Dor, A., and Yakhini, Z. (2009). Novel rank-based statistical methods reveal microRNAs with differential expression in multiple cancer types. PLoS ONE, 4.","DOI":"10.1371\/journal.pone.0008003"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"5150","DOI":"10.1038\/srep05150","article-title":"Genome-wide survey of tissue-specific microRNA and transcription factor regulatory networks in 12 tissues","volume":"4","author":"Guo","year":"2014","journal-title":"Sci. Rep."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1038\/nrm3854","article-title":"Regulation of microRNA function in somatic stem cell proliferation and differentiation","volume":"15","author":"Shenoy","year":"2014","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"3582","DOI":"10.1016\/j.jmb.2013.03.007","article-title":"The role of miRNAs in regulating gene expression networks","volume":"425","author":"Gurtan","year":"2013","journal-title":"J. Mol. Biol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"650","DOI":"10.1093\/bioinformatics\/bti042","article-title":"Genome-wide midrange transcription profiles reveal expression level relationships in human tissue specification","volume":"21","author":"Yanai","year":"2005","journal-title":"Bioinformatics"},{"key":"ref_9","first-page":"205","article-title":"A benchmark of gene expression tissue-specificity metrics","volume":"18","year":"2017","journal-title":"Brief. Bioinform."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Han, Y., and Zhou, Y. (2022). Comprehensive Identification of Human Cell Type Chromatin Activity-Specific and Cell Type Expression-Specific MicroRNAs. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23137324"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"D129","DOI":"10.1093\/nar\/gkae1036","article-title":"miRNATissueAtlas 2025: An update to the uniformly processed and annotated human and mouse non-coding RNA tissue atlas","volume":"53","author":"Rishik","year":"2025","journal-title":"Nucleic Acids Res."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Moradi, S., Kamal, A., Aboulkheyr Es, H., Farhadi, F., Ebrahimi, M., Chitsaz, H., Sharifi-Zarchi, A., and Baharvand, H. (2022). Pan-cancer analysis of microRNA expression profiles highlights microRNAs enriched in normal body cells as effective suppressors of multiple tumor types: A study based on TCGA database. PLoS ONE, 17.","DOI":"10.1371\/journal.pone.0267291"},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Sarver, A.L., Sarver, A.E., Yuan, C., and Subramanian, S. (2018). OMCD: OncomiR Cancer Database. BMC Cancer, 18.","DOI":"10.1186\/s12885-018-5085-z"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"5228","DOI":"10.1038\/s41467-018-07657-1","article-title":"Pan-cancer characterisation of microRNA across cancer hallmarks reveals microRNA-mediated downregulation of tumour suppressors","volume":"9","author":"Dhawan","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"e22490","DOI":"10.1002\/jbt.22490","article-title":"MiR-671-5p plays a promising role in restraining osteosarcoma cell characteristics through targeting TUFT1","volume":"34","author":"Ma","year":"2020","journal-title":"J. Biochem. Mol. Toxicol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1892","DOI":"10.7150\/ijbs.32429","article-title":"miR-671-5p Blocks The Progression Of Human Esophageal Squamous Cell Carcinoma By Suppressing FGFR2","volume":"15","author":"Li","year":"2019","journal-title":"Int. J. Biol. Sci."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"996","DOI":"10.1089\/dna.2019.4870","article-title":"miR-671-5p Inhibits Tumor Proliferation by Blocking Cell Cycle in Osteosarcoma","volume":"38","author":"Xin","year":"2019","journal-title":"DNA Cell Biol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"983","DOI":"10.1002\/kjm2.12424","article-title":"miR-671-5p repressed progression of papillary thyroid carcinoma via TRIM14","volume":"37","author":"Wang","year":"2021","journal-title":"Kaohsiung J. Med. Sci."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"2924","DOI":"10.1111\/1759-7714.15078","article-title":"MiR-671-5p sponging activity of circMMP1 promotes esophageal squamous cancer progression","volume":"14","author":"Li","year":"2023","journal-title":"Thorac. Cancer"},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Bernstein, D.L., Jiang, X., and Rom, S. (2021). let-7 microRNAs: Their Role in Cerebral and Cardiovascular Diseases, Inflammation, Cancer, and Their Regulation. Biomedicines, 9.","DOI":"10.3390\/biomedicines9060606"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1186\/s13046-019-1059-5","article-title":"MicroRNA-34 family: A potential tumor suppressor and therapeutic candidate in cancer","volume":"38","author":"Zhang","year":"2019","journal-title":"J. Exp. Clin. Cancer Res."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1016\/j.omtn.2019.06.015","article-title":"Expression and Role of MicroRNAs from the miR-200 Family in the Tumor Formation and Metastatic Propensity of Pancreatic Cancer","volume":"17","author":"Ginesta","year":"2019","journal-title":"Mol. Ther. Nucleic Acids"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"27","DOI":"10.14712\/fb2026.0003","article-title":"MiR-200a-5p Is a Potential Prognostic Marker for Lung Adenocarcinoma: Based on Transcriptomics, Bioinformatics, and In Vitro and In Vivo Experiments","volume":"72","author":"Liu","year":"2026","journal-title":"Folia Biol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"922","DOI":"10.1007\/s12672-026-05074-1","article-title":"The role of the miR-200a-5p\/NFIA axis in the progression and clinical prognosis of non-small cell lung cancer","volume":"17","author":"Wang","year":"2026","journal-title":"Discov. Oncol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1913","DOI":"10.1111\/jcmm.12358","article-title":"MicroRNA-145: A potent tumour suppressor that regulates multiple cellular pathways","volume":"18","author":"Cui","year":"2014","journal-title":"J. Cell Mol. Med."},{"key":"ref_26","first-page":"6739","article-title":"MiR-145: A potential biomarker of cancer migration and invasion","volume":"11","author":"Xu","year":"2019","journal-title":"Am. J. Transl. Res."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"769","DOI":"10.1186\/s12967-023-04649-8","article-title":"The important role of miR-1-3p in cancers","volume":"21","author":"Dai","year":"2023","journal-title":"J. Transl. Med."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Hong, Y., Ye, M., Wang, F., Fang, J., Wang, C., Luo, J., Liu, J., Liu, J., Liu, L., and Zhao, Q. (2021). MiR-21-3p Promotes Hepatocellular Carcinoma Progression via SMAD7\/YAP1 Regulation. Front. Oncol., 11.","DOI":"10.3389\/fonc.2021.642030"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"260","DOI":"10.3892\/ol.2022.13380","article-title":"Nicotine-induced miR-21-3p promotes chemoresistance in lung cancer by negatively regulating FOXO3a","volume":"24","author":"Zhang","year":"2022","journal-title":"Oncol. Lett."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1038\/s41420-022-01005-5","article-title":"Regulatory mechanism of miR-20a-5p expression in Cancer","volume":"8","author":"Huang","year":"2022","journal-title":"Cell Death Discov."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1186\/s40463-021-00497-y","article-title":"Comparison of miRNA expressions among benign, premalignant and malignant lesions of the larynx: Could they be transformation biomarkers?","volume":"50","author":"Tuncturk","year":"2021","journal-title":"J. Otolaryngol. Head Neck Surg."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Dos Santos, G.A., Chatsirisupachai, K., Avelar, R.A., and de Magalhaes, J.P. (2023). Transcriptomic analysis reveals a tissue-specific loss of identity during ageing and cancer. BMC Genom., 24.","DOI":"10.1186\/s12864-023-09756-w"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"9053","DOI":"10.2147\/OTT.S217324","article-title":"miR-522-3p Promotes Osteosarcoma Cell Growth by Regulating Glucose Uptake and GLUT1 Expression","volume":"12","author":"Chen","year":"2019","journal-title":"OncoTargets Ther."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1111\/jgh.16345","article-title":"Exosomal-miR-522-3p derived from cancer-associated fibroblasts accelerates tumor metastasis and angiogenesis via repression bone morphogenetic protein 5 in colorectal cancer","volume":"39","author":"Zhang","year":"2024","journal-title":"J. Gastroenterol. Hepatol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"1945","DOI":"10.1002\/cbin.11634","article-title":"BVES-AS1 inhibits the malignant behaviors of colon adenocarcinoma cells via regulating BVES","volume":"45","author":"Wen","year":"2021","journal-title":"Cell Biol. Int."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"334","DOI":"10.1111\/hepr.13870","article-title":"circTMEM181 upregulates ARHGAP29 to inhibit hepatocellular carcinoma migration and invasion by sponging miR-519a-5p","volume":"53","author":"Chen","year":"2023","journal-title":"Hepatol. Res."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1455","DOI":"10.1111\/1440-1681.13321","article-title":"Ginsenoside 20(S)-Rg3 upregulates HIF-1alpha-targeting miR-519a-5p to inhibit the Warburg effect in ovarian cancer cells","volume":"47","author":"Lu","year":"2020","journal-title":"Clin. Exp. Pharmacol. Physiol."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Pan, G., Jiang, B., Yi, Z., Yin, J., and Liu, Y. (2023). Exosomal miR-105-5p derived from bladder cancer stem cells targets for GPR12 to promote the malignancy of bladder cancer. BMC Urol., 23.","DOI":"10.1186\/s12894-023-01326-2"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"3022","DOI":"10.1097\/CM9.0000000000002689","article-title":"Hsa-miR-105-5p acts as an oncogene in triple-negative breast cancer","volume":"136","author":"Qian","year":"2023","journal-title":"Chin. Med. J."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1016\/j.canlet.2021.05.037","article-title":"miR-105-5p regulates PD-L1 expression and tumor immunogenicity in gastric cancer","volume":"518","author":"Miliotis","year":"2021","journal-title":"Cancer Lett."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1186\/s11658-024-00656-9","article-title":"The miR-1269a\/PCDHGA9\/CXCR4\/beta-catenin pathway promotes colorectal cancer invasion and metastasis","volume":"29","author":"Mei","year":"2024","journal-title":"Cell Mol. Biol. Lett."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1186\/s12957-025-04026-3","article-title":"LINC01082 inhibits malignant phenotypes of bladder cancer through sponging miR-1269a","volume":"23","author":"Su","year":"2025","journal-title":"World J. Surg. Oncol."},{"key":"ref_43","first-page":"111832","article-title":"Hsa-miR-1269a up-regulation fosters the malignant progression of esophageal squamous cell carcinoma via targeting FAM46C","volume":"827","author":"Ma","year":"2023","journal-title":"Mutat. Res."},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Zhang, Y., Wang, Q., Luo, N., Liu, J., Ren, H., Shao, X., Zhang, L., and Yu, Y. (2020). MicroRNA-1269a Promotes Proliferation and Arrest of Apoptosis of Glioma Cells by Directly Targeting ATRX. Front. Oncol., 10.","DOI":"10.3389\/fonc.2020.563901"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1007\/s12311-025-01812-3","article-title":"miR-512-3p as a Potential Biomarker of Poor Outcome in Pediatric Medulloblastoma","volume":"24","author":"Correa","year":"2025","journal-title":"Cerebellum"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"194921","DOI":"10.1016\/j.bbagrm.2023.194921","article-title":"Novel splice variants of LINC00963 suppress colorectal cancer cell proliferation via miR-10a\/miR-143\/miR-217\/miR-512-mediated regulation of PI3K\/AKT and Wnt\/beta-catenin signaling pathways","volume":"1866","author":"Ghaemi","year":"2023","journal-title":"Biochim. Biophys. Acta Gene Regul. Mech."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.trsl.2026.05.007","article-title":"Circ-PHC3 promotes malignant progression of ovarian cancer through regulating miR-512-3p\/SLC2A1\/HK2 mediated glucose metabolism","volume":"294","author":"Tian","year":"2026","journal-title":"Transl. Res."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"575","DOI":"10.1016\/j.ajur.2024.01.007","article-title":"LncRNA HCG18 promotes prostate cancer progression by regulating the miR-512-3p\/HK-2 axis","volume":"11","author":"Zhu","year":"2024","journal-title":"Asian J. Urol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"155281","DOI":"10.1016\/j.prp.2024.155281","article-title":"Hsa_circ_0087862 contributes to the progression of colorectal cancer through regulating miR-512-3p\/HK2 axis","volume":"257","author":"Chen","year":"2024","journal-title":"Pathol. Res. Pract."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"427","DOI":"10.3727\/096504016X14747300207374","article-title":"miR-935 Promotes Liver Cancer Cell Proliferation and Migration by Targeting SOX7","volume":"25","author":"Liu","year":"2017","journal-title":"Oncol. Res."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1016\/j.biopha.2016.01.011","article-title":"miR-935 promotes gastric cancer cell proliferation by targeting SOX7","volume":"79","author":"Yang","year":"2016","journal-title":"BioMed Pharmacother."},{"key":"ref_52","first-page":"3397","article-title":"Knockdown of miR-935 increases paclitaxel sensitivity via regulation of SOX7 in non-small-cell lung cancer","volume":"18","author":"Peng","year":"2018","journal-title":"Mol. Med. Rep."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"e13791","DOI":"10.1111\/aji.13791","article-title":"METTL3 promotes choriocarcinoma progression by activating the miR-935\/GJA1 pathway in an m6A-dependent manner","volume":"90","author":"Wang","year":"2023","journal-title":"Am. J. Reprod. Immunol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"10817","DOI":"10.2147\/OTT.S244409","article-title":"MiR-935\/HIF1alpha Feedback Loop Inhibits the Proliferation and Invasiveness of Glioma","volume":"13","author":"Huang","year":"2020","journal-title":"OncoTargets Ther."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"6101","DOI":"10.21873\/anticanres.14631","article-title":"miR-935 Inhibits Oral Squamous Cell Carcinoma and Targets Inositol Polyphosphate-4-phosphatase Type IA (INPP4A)","volume":"40","author":"Maruyama","year":"2020","journal-title":"Anticancer Res."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1186\/s13045-023-01468-8","article-title":"Safety and activity of the first-in-class locked nucleic acid (LNA) miR-221 selective inhibitor in refractory advanced cancer patients: A first-in-human, phase 1, open-label, dose-escalation study","volume":"16","author":"Tassone","year":"2023","journal-title":"J. Hematol. Oncol."},{"key":"ref_57","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_58","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1007\/s10637-016-0407-y","article-title":"Phase I study of MRX34, a liposomal miR-34a mimic, administered twice weekly in patients with advanced solid tumors","volume":"35","author":"Beg","year":"2017","journal-title":"Investig. New Drugs"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"580","DOI":"10.1038\/ng.2653","article-title":"The Genotype-Tissue Expression (GTEx) project","volume":"45","author":"Consortium","year":"2013","journal-title":"Nat. Genet."},{"key":"ref_60","first-page":"A68","article-title":"The Cancer Genome Atlas (TCGA): An immeasurable source of knowledge","volume":"19","author":"Tomczak","year":"2015","journal-title":"Contemp. Oncol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"btad547","DOI":"10.1093\/bioinformatics\/btad547","article-title":"PyDESeq2: A python package for bulk RNA-seq differential expression analysis","volume":"39","author":"Muzellec","year":"2023","journal-title":"Bioinformatics"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"W409","DOI":"10.1093\/nar\/gkab297","article-title":"miRTargetLink 2.0-interactive miRNA target gene and target pathway networks","volume":"49","author":"Kern","year":"2021","journal-title":"Nucleic Acids Res."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"D147","DOI":"10.1093\/nar\/gkae1072","article-title":"miRTarBase 2025: Updates to the collection of experimentally validated microRNA-target interactions","volume":"53","author":"Cui","year":"2025","journal-title":"Nucleic Acids Res."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"D217","DOI":"10.1093\/nar\/gkac1070","article-title":"MirDIP 5.2: Tissue context annotation and novel microRNA curation","volume":"51","author":"Hauschild","year":"2023","journal-title":"Nucleic Acids Res."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"D127","DOI":"10.1093\/nar\/gkz757","article-title":"miRDB: An online database for prediction of functional microRNA targets","volume":"48","author":"Chen","year":"2020","journal-title":"Nucleic Acids Res."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"D730","DOI":"10.1093\/nar\/gkae1113","article-title":"The STRING database in 2025: Protein networks with directionality of regulation","volume":"53","author":"Szklarczyk","year":"2025","journal-title":"Nucleic Acids Res."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/18\/8153\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T04:13:59Z","timestamp":1789618439000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/18\/8153"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,13]]},"references-count":66,"journal-issue":{"issue":"18","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["ijms27188153"],"URL":"https:\/\/doi.org\/10.3390\/ijms27188153","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,13]]}}}