{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,22]],"date-time":"2026-07-22T19:48:33Z","timestamp":1784749713328,"version":"3.55.0"},"reference-count":95,"publisher":"The Company of Biologists","issue":"2","license":[{"start":{"date-parts":[[2026,3,5]],"date-time":"2026-03-05T00:00:00Z","timestamp":1772668800000},"content-version":"vor","delay-in-days":32,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"National Cancer Institute","award":["R01CA222570-NIH-NCI"],"award-info":[{"award-number":["R01CA222570-NIH-NCI"]}]},{"name":"National Cancer Institute","award":["P30CA042014"],"award-info":[{"award-number":["P30CA042014"]}]},{"DOI":"10.13039\/100000048","name":"American Cancer Society","doi-asserted-by":"crossref","award":["RSG-22-014-01-CCB"],"award-info":[{"award-number":["RSG-22-014-01-CCB"]}],"id":[{"id":"10.13039\/100000048","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000048","name":"American Cancer Society","doi-asserted-by":"crossref","award":["DBG-23-1037804-01-TBE"],"award-info":[{"award-number":["DBG-23-1037804-01-TBE"]}],"id":[{"id":"10.13039\/100000048","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100001021","name":"Damon Runyon Cancer Research Foundation","doi-asserted-by":"crossref","award":["DR 61-20"],"award-info":[{"award-number":["DR 61-20"]}],"id":[{"id":"10.13039\/100001021","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100007747","name":"University of Utah","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100007747","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100010637","name":"Huntsman Cancer Foundation","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100010637","id-type":"DOI","asserted-by":"crossref"}]},{"name":"Huntsman Cancer Institute, University of Utah"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,2,1]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>The prevalence of hepatocellular carcinoma (HCC) is rising in parallel with increasing obesity and metabolic dysfunction-associated steatohepatitis (MASH). MicroRNAs (miRNAs) are key post-transcriptional regulators of gene expression and attractive targets for HCC therapy. Here, we sought to identify and characterize dysregulated miRNAs in MASH-driven HCC (MASH-HCC). We profiled miRNA expression in liver tissue from patients with MASH or MASH-HCC and in zebrafish HCC driven by activated \u03b2-catenin (CTNNB1), one of the most commonly mutated oncogenes in MASH-HCC. We found overlap between dysregulated human and zebrafish miRNAs, including microRNA-21 (miR-21), which was increasingly upregulated from normal liver to MASH to MASH-HCC. We generated transgenic zebrafish that overexpress or sponge miR-21 in hepatocytes. We found that miR-21 overexpression caused larval liver overgrowth and increased HCC, while miR-21 sponge suppressed \u03b2-catenin-driven larval liver overgrowth. By performing histological and lipidomics analysis, we found that overexpression of miR-21, like activated \u03b2-catenin (ABC), suppressed lipid accumulation in response to a high cholesterol diet and increased accumulation of acylcarnitines. Thus, miR-21, which is similarly upregulated in human and zebrafish HCC, promotes lipid metabolic changes that may help drive hepatocarcinogenesis.<\/jats:p>","DOI":"10.1242\/dmm.052583","type":"journal-article","created":{"date-parts":[[2026,1,26]],"date-time":"2026-01-26T08:32:04Z","timestamp":1769416324000},"source":"Crossref","is-referenced-by-count":2,"title":["MicroRNA-21 promotes dysregulated lipid metabolism and hepatocellular carcinoma"],"prefix":"10.1242","volume":"19","author":[{"given":"Chad","family":"VanSant-Webb","sequence":"first","affiliation":[{"name":"University of Utah 1 Department of Pathology , , 15 North Medical Drive East, Salt Lake City, UT 84101 ,","place":["USA"]},{"name":"2 Huntsman Cancer Institute, 2000 Circle of Hope Way, Salt Lake City, UT 84101, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jessye C.","family":"Castro","sequence":"additional","affiliation":[{"name":"University of Utah 1 Department of Pathology , , 15 North Medical Drive East, Salt Lake City, UT 84101 ,","place":["USA"]},{"name":"2 Huntsman Cancer Institute, 2000 Circle of Hope Way, Salt Lake City, UT 84101, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Audrey Y.","family":"Su","sequence":"additional","affiliation":[{"name":"University of Utah 1 Department of Pathology , , 15 North Medical Drive East, Salt Lake City, UT 84101 ,","place":["USA"]},{"name":"2 Huntsman Cancer Institute, 2000 Circle of Hope Way, Salt Lake City, UT 84101, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kiandra","family":"Hawkins","sequence":"additional","affiliation":[{"name":"University of Utah 1 Department of Pathology , , 15 North Medical Drive East, Salt Lake City, UT 84101 ,","place":["USA"]},{"name":"2 Huntsman Cancer Institute, 2000 Circle of Hope Way, Salt Lake City, UT 84101, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aavrati","family":"Saxena","sequence":"additional","affiliation":[{"name":"University of Utah 1 Department of Pathology , , 15 North Medical Drive East, Salt Lake City, UT 84101 ,","place":["USA"]},{"name":"2 Huntsman Cancer Institute, 2000 Circle of Hope Way, Salt Lake City, UT 84101, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jillian","family":"Wright","sequence":"additional","affiliation":[{"name":"2 Huntsman Cancer Institute, 2000 Circle of Hope Way, Salt Lake City, UT 84101, USA"},{"name":"University of Utah 3 Department of Biochemistry , , 15 N Medical Drive E, Salt Lake City, UT 84112 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Richard","family":"Smith","sequence":"additional","affiliation":[{"name":"University of Utah 1 Department of Pathology , , 15 North Medical Drive East, Salt Lake City, UT 84101 ,","place":["USA"]},{"name":"2 Huntsman Cancer Institute, 2000 Circle of Hope Way, Salt Lake City, UT 84101, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marco","family":"Fragoso-Garc\u00eda","sequence":"additional","affiliation":[{"name":"Albert Einstein College of Medicine 4 Department of Pathology , , 1301 Morris Park Avenue, Bronx, NY 10461 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yian Ann","family":"Chen","sequence":"additional","affiliation":[{"name":"University of Utah 5 Department of Population Health Sciences , , 295 Chipeta Way, Salt Lake City, UT 84108 ,","place":["USA"]},{"name":"University of Utah 6 Department of Internal Medicine , , 30 N Mario Capecchi Dr., Salt Lake City, UT 84112 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carrie","family":"Barton","sequence":"additional","affiliation":[{"name":"Office of Comparative Medicine 7 , 50 S 2030 E, Salt Lake City, UT 84132 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chris","family":"Stubben","sequence":"additional","affiliation":[{"name":"2 Huntsman Cancer Institute, 2000 Circle of Hope Way, Salt Lake City, UT 84101, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryan M.","family":"O'Connell","sequence":"additional","affiliation":[{"name":"University of Utah 1 Department of Pathology , , 15 North Medical Drive East, Salt Lake City, UT 84101 ,","place":["USA"]},{"name":"2 Huntsman Cancer Institute, 2000 Circle of Hope Way, Salt Lake City, UT 84101, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8651-6846","authenticated-orcid":false,"given":"Gregory S.","family":"Ducker","sequence":"additional","affiliation":[{"name":"2 Huntsman Cancer Institute, 2000 Circle of Hope Way, Salt Lake City, UT 84101, USA"},{"name":"University of Utah 3 Department of Biochemistry , , 15 N Medical Drive E, Salt Lake City, UT 84112 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0471-2473","authenticated-orcid":false,"given":"Kimberley J.","family":"Evason","sequence":"additional","affiliation":[{"name":"University of Utah 1 Department of Pathology , , 15 North Medical Drive East, Salt Lake City, UT 84101 ,","place":["USA"]},{"name":"2 Huntsman Cancer Institute, 2000 Circle of Hope Way, Salt Lake City, UT 84101, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2026,3,5]]},"reference":[{"key":"2026030505115514100_DMM052583C2","first-page":"161","article-title":"The role of microRNAs in human diseases","volume":"2","author":"Ardekani","year":"2010","journal-title":"Avicenna J. Med. Biotechnol."},{"key":"2026030505115514100_DMM052583C3","doi-asserted-by":"publisher","first-page":"1972","DOI":"10.1002\/hep.23527","article-title":"The incidence and risk factors of hepatocellular carcinoma in patients with nonalcoholic steatohepatitis","volume":"51","author":"Ascha","year":"2010","journal-title":"Hepatology"},{"key":"2026030505115514100_DMM052583C4","doi-asserted-by":"publisher","first-page":"16878","DOI":"10.1038\/s41598-017-17204-5","article-title":"QuPath: open source software for digital pathology image analysis","volume":"7","author":"Bankhead","year":"2017","journal-title":"Sci. Rep."},{"key":"2026030505115514100_DMM052583C5","doi-asserted-by":"publisher","first-page":"886","DOI":"10.1002\/hep.20868","article-title":"Cholesterol accumulation and liver cell death in mice with Niemann-Pick type C disease","volume":"42","author":"Beltroy","year":"2005","journal-title":"Hepatology"},{"key":"2026030505115514100_DMM052583C6","doi-asserted-by":"publisher","first-page":"407","DOI":"10.1073\/pnas.0406252102","article-title":"tp53 mutant zebrafish develop malignant peripheral nerve sheath tumors","volume":"102","author":"Berghmans","year":"2005","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2026030505115514100_DMM052583C7","doi-asserted-by":"publisher","first-page":"1326","DOI":"10.1038\/s41467-019-09152-7","article-title":"Sterol regulatory element binding protein 1 couples mechanical cues and lipid metabolism","volume":"10","author":"Bertolio","year":"2019","journal-title":"Nat. Commun."},{"key":"2026030505115514100_DMM052583C8","doi-asserted-by":"publisher","first-page":"505","DOI":"10.1016\/S0022-2275(20)42221-7","article-title":"Multivalent feedback regulation of HMG CoA reductase, a control mechanism coordinating isoprenoid synthesis and cell growth","volume":"21","author":"Brown","year":"1980","journal-title":"J. Lipid Res."},{"key":"2026030505115514100_DMM052583C9","doi-asserted-by":"publisher","first-page":"8293","DOI":"10.1038\/sj.onc.1206118","article-title":"New targets of \u03b2-catenin signaling in the liver are involved in the glutamine metabolism","volume":"21","author":"Cadoret","year":"2002","journal-title":"Oncogene"},{"key":"2026030505115514100_DMM052583C10","doi-asserted-by":"publisher","first-page":"46","DOI":"10.1038\/cdd.2014.136","article-title":"MicroRNAs in liver cancer: a model for investigating pathogenesis and novel therapeutic approaches","volume":"22","author":"Callegari","year":"2015","journal-title":"Cell Death Differ."},{"key":"2026030505115514100_DMM052583C11","doi-asserted-by":"publisher","first-page":"1327","DOI":"10.1016\/j.cell.2017.05.046","article-title":"Comprehensive and integrative genomic characterization of hepatocellular carcinoma","volume":"169","author":"Cancer Genome Atlas Research Network","year":"2017","journal-title":"Cell"},{"key":"2026030505115514100_DMM052583C12","doi-asserted-by":"publisher","first-page":"918","DOI":"10.1186\/s12967-023-04801-4","article-title":"Integrated plasma metabolomics and lipidomics profiling highlights distinctive signature of hepatocellular carcinoma in HCV patients","volume":"21","author":"Caponigro","year":"2023","journal-title":"J. Transl. Med."},{"key":"2026030505115514100_DMM052583C13","doi-asserted-by":"publisher","first-page":"1712","DOI":"10.3390\/ijms17101712","article-title":"MicroRNA in control of gene expression: an overview of nuclear functions","volume":"17","author":"Catalanotto","year":"2016","journal-title":"Int. J. Mol. Sci."},{"key":"2026030505115514100_DMM052583C14","doi-asserted-by":"publisher","first-page":"386","DOI":"10.1126\/science.aav3722","article-title":"Targeting a ceramide double bond improves insulin resistance and hepatic steatosis","volume":"365","author":"Chaurasia","year":"2019","journal-title":"Science"},{"key":"2026030505115514100_DMM052583C15","doi-asserted-by":"publisher","first-page":"Unit14.11","DOI":"10.1002\/0471250953.bi1411s37","article-title":"LC-MS data processing with MAVEN: a metabolomic analysis and visualization engine","volume":"Chapter 14","author":"Clasquin","year":"2012","journal-title":"Curr. Protoc. Bioinformatics"},{"key":"2026030505115514100_DMM052583C16","doi-asserted-by":"publisher","first-page":"e71","DOI":"10.1093\/nar\/gkv1507","article-title":"TCGAbiolinks: an R\/Bioconductor package for integrative analysis of TCGA data","volume":"44","author":"Colaprico","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"2026030505115514100_DMM052583C17","doi-asserted-by":"publisher","first-page":"4983","DOI":"10.3390\/cancers13194983","article-title":"Mir-21 suppression promotes mouse hepatocarcinogenesis","volume":"13","author":"Correia De Sousa","year":"2021","journal-title":"Cancers"},{"key":"2026030505115514100_DMM052583C18","doi-asserted-by":"publisher","first-page":"42","DOI":"10.1186\/s12986-015-0036-z","article-title":"High fat plus high cholesterol diet lead to hepatic steatosis in zebrafish larvae: a novel model for screening anti-hepatic steatosis drugs","volume":"12","author":"Dai","year":"2015","journal-title":"Nutr. Metab."},{"key":"2026030505115514100_DMM052583C19","doi-asserted-by":"publisher","first-page":"1846","DOI":"10.1093\/bioinformatics\/btm254","article-title":"GEOquery: a bridge between the Gene Expression Omnibus (GEO) and BioConductor","volume":"23","author":"Davis","year":"2007","journal-title":"Bioinformatics"},{"key":"2026030505115514100_DMM052583C20","doi-asserted-by":"publisher","first-page":"710","DOI":"10.1016\/j.jhep.2018.11.034","article-title":"Metformin modulates innate immune-mediated inflammation and early progression of NAFLD-associated hepatocellular carcinoma in zebrafish","volume":"70","author":"De Oliveira","year":"2019","journal-title":"J. Hepatol."},{"key":"2026030505115514100_DMM052583C21","doi-asserted-by":"publisher","first-page":"227","DOI":"10.1016\/S0167-4781(00)00102-0","article-title":"cDNA sequence and tissue-specific expression of a basic liver-type fatty acid binding protein in adult zebrafish (Danio rerio)","volume":"1492","author":"Denovan-Wright","year":"2000","journal-title":"Biochim. Biophys. Acta"},{"key":"2026030505115514100_DMM052583C22","doi-asserted-by":"publisher","first-page":"3439","DOI":"10.1093\/bioinformatics\/bti525","article-title":"BioMart and Bioconductor: a powerful link between biological databases and microarray data analysis","volume":"21","author":"Durinck","year":"2005","journal-title":"Bioinformatics"},{"key":"2026030505115514100_DMM052583C96","doi-asserted-by":"publisher","first-page":"1184","DOI":"10.1038\/nprot.2009.97","article-title":"Mapping identifiers for the integration of genomic datasets with the R\/Bioconductor package biomaRt","volume":"4","author":"Durinck","year":"2009","journal-title":"Nat. Protoc."},{"key":"2026030505115514100_DMM052583C23","doi-asserted-by":"publisher","first-page":"10663","DOI":"10.1038\/s41598-019-47216-2","article-title":"Altered serum acylcarnitine profile is associated with the status of nonalcoholic fatty liver disease (NAFLD) and NAFLD-related hepatocellular carcinoma","volume":"9","author":"Enooku","year":"2019","journal-title":"Sci. Rep."},{"key":"2026030505115514100_DMM052583C24","doi-asserted-by":"publisher","first-page":"e1005305","DOI":"10.1371\/journal.pgen.1005305","article-title":"Identification of chemical inhibitors of \u03b2-catenin-driven liver tumorigenesis in zebrafish","volume":"11","author":"Evason","year":"2015","journal-title":"PLoS Genet."},{"key":"2026030505115514100_DMM052583C25","doi-asserted-by":"publisher","first-page":"e168476","DOI":"10.1172\/jci.insight.168476","article-title":"miR-33 deletion in hepatocytes attenuates MASLD-MASH-HCC progression","volume":"9","author":"Fern\u00e1ndez-Tussy","year":"2024","journal-title":"JCI Insight"},{"key":"2026030505115514100_DMM052583C26","doi-asserted-by":"publisher","first-page":"272","DOI":"10.1210\/en.2017-00850","article-title":"\u03b2-catenin directs long-chain fatty acid catabolism in the osteoblasts of male mice","volume":"159","author":"Frey","year":"2018","journal-title":"Endocrinology"},{"key":"2026030505115514100_DMM052583C27","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1101\/gr.082701.108","article-title":"Most mammalian mRNAs are conserved targets of microRNAs","volume":"19","author":"Friedman","year":"2009","journal-title":"Genome Res."},{"key":"2026030505115514100_DMM052583C28","doi-asserted-by":"publisher","first-page":"707","DOI":"10.1046\/j.1523-1747.2003.12123.x","article-title":"Identification of the \u0394-6 desaturase of human sebaceous glands: expression and enzyme activity","volume":"120","author":"Ge","year":"2003","journal-title":"J. Investig. Dermatol."},{"key":"2026030505115514100_DMM052583C29","doi-asserted-by":"publisher","first-page":"2628","DOI":"10.1093\/bioinformatics\/btz931","article-title":"ShinyGO: a graphical gene-set enrichment tool for animals and plants","volume":"36","author":"Ge","year":"2020","journal-title":"Bioinformatics"},{"key":"2026030505115514100_DMM052583C30","doi-asserted-by":"publisher","first-page":"710","DOI":"10.1210\/js.2018-00122","article-title":"Transcriptomic profiling of obesity-related nonalcoholic steatohepatitis reveals a core set of fibrosis-specific genes","volume":"2","author":"Gerhard","year":"2018","journal-title":"J. Endocr. Soc."},{"key":"2026030505115514100_DMM052583C31","doi-asserted-by":"publisher","first-page":"eaba4448","DOI":"10.1126\/scitranslmed.aba4448","article-title":"Transcriptomic profiling across the nonalcoholic fatty liver disease spectrum reveals gene signatures for steatohepatitis and fibrosis","volume":"12","author":"Govaere","year":"2020","journal-title":"Sci. Transl. Med."},{"key":"2026030505115514100_DMM052583C32","volume-title":"Histopathology of Preclinical Toxicity Studies: Interpretation and Relevance in Drug Safety Evaluation","author":"Greaves","year":"2007"},{"key":"2026030505115514100_DMM052583C33","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1016\/j.molcel.2007.06.017","article-title":"MicroRNA targeting specificity in mammals: determinants beyond seed pairing","volume":"27","author":"Grimson","year":"2007","journal-title":"Mol. Cell"},{"key":"2026030505115514100_DMM052583C34","doi-asserted-by":"publisher","first-page":"356","DOI":"10.7555\/JBR.29.20150067","article-title":"Hypercholesterolemia, low density lipoprotein receptor and proprotein convertase subtilisin\/kexin-type 9","volume":"29","author":"Gu","year":"2015","journal-title":"J. Biomed. Res."},{"key":"2026030505115514100_DMM052583C35","doi-asserted-by":"publisher","first-page":"971","DOI":"10.1177\/0192623312448935","article-title":"Liver hypertrophy: a review of adaptive (adverse and non-adverse) changes--conclusions from the 3rd International ESTP expert workshop","volume":"40","author":"Hall","year":"2012","journal-title":"Toxicol. Pathol."},{"key":"2026030505115514100_DMM052583C36","doi-asserted-by":"publisher","first-page":"573","DOI":"10.1111\/j.1755-5949.2012.00344.x","article-title":"MicroRNA-21 expression is regulated by \u03b2-catenin\/STAT3 pathway and promotes glioma cell invasion by direct targeting RECK","volume":"18","author":"Han","year":"2012","journal-title":"CNS Neurosci. Ther."},{"key":"2026030505115514100_DMM052583C37","doi-asserted-by":"publisher","first-page":"23056","DOI":"10.1038\/s41598-023-50328-5","article-title":"Semaglutide reduces tumor burden in the GAN diet-induced obese and biopsy-confirmed mouse model of NASH-HCC with advanced fibrosis","volume":"13","author":"Hansen","year":"2023","journal-title":"Sci. Rep."},{"key":"2026030505115514100_DMM052583C38","doi-asserted-by":"publisher","first-page":"1920","DOI":"10.1002\/1873-3468.14134","article-title":"Ablation of fatty acid desaturase 2 (FADS2) exacerbates hepatic triacylglycerol and cholesterol accumulation in polyunsaturated fatty acid-depleted mice","volume":"595","author":"Hayashi","year":"2021","journal-title":"FEBS Lett."},{"key":"2026030505115514100_DMM052583C39","doi-asserted-by":"publisher","first-page":"1372","DOI":"10.1038\/s41598-020-80621-6","article-title":"Stimulation of hepatocarcinogenesis by activated cholangiocytes via Il17a\/f1 pathway in kras transgenic zebrafish model","volume":"11","author":"Helal","year":"2021","journal-title":"Sci. Rep."},{"key":"2026030505115514100_DMM052583C40","doi-asserted-by":"publisher","first-page":"14896","DOI":"10.1073\/pnas.0906348106","article-title":"Distinct populations of quiescent and proliferative pancreatic \u03b2-cells identified by HOTcre mediated labeling","volume":"106","author":"Hesselson","year":"2009","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2026030505115514100_DMM052583C41","doi-asserted-by":"publisher","first-page":"1125","DOI":"10.1172\/JCI0215593","article-title":"SREBPs: activators of the complete program of cholesterol and fatty acid synthesis in the liver","volume":"109","author":"Horton","year":"2002","journal-title":"J. Clin. Invest."},{"key":"2026030505115514100_DMM052583C42","doi-asserted-by":"publisher","first-page":"18561","DOI":"10.1073\/pnas.1905221116","article-title":"Phenotypical microRNA screen reveals a noncanonical role of CDK2 in regulating neutrophil migration","volume":"116","author":"Hsu","year":"2019","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2026030505115514100_DMM052583C44","doi-asserted-by":"publisher","first-page":"102752","DOI":"10.1016\/j.omtn.2025.102752","article-title":"MicroRNA-21 is a potential therapeutic agent targeting Tgfbi and mitigating high-fat-diet-induced liver disease and cancer","volume":"36","author":"Jagtap","year":"2025","journal-title":"Mol. Ther. Nucleic Acids"},{"key":"2026030505115514100_DMM052583C45","doi-asserted-by":"publisher","first-page":"100637","DOI":"10.1016\/j.jlr.2024.100637","article-title":"A high-cholesterol zebrafish diet promotes hypercholesterolemia and fasting-associated liver steatosis","volume":"65","author":"Jin","year":"2024","journal-title":"J. Lipid Res."},{"key":"2026030505115514100_DMM052583C46","doi-asserted-by":"publisher","first-page":"bio047829","DOI":"10.1242\/bio.047829","article-title":"Heterogeneous beta-catenin activation is sufficient to cause hepatocellular carcinoma in zebrafish","volume":"8","author":"Kalasekar","year":"2019","journal-title":"Biol. Open"},{"key":"2026030505115514100_DMM052583C47","doi-asserted-by":"publisher","first-page":"5524","DOI":"10.3390\/cancers13215524","article-title":"Intratumor heterogeneity in hepatocellular carcinoma: challenges and opportunities","volume":"13","author":"Kalasekar","year":"2021","journal-title":"Cancers"},{"key":"2026030505115514100_DMM052583C48","doi-asserted-by":"publisher","first-page":"1212","DOI":"10.1097\/HEP.0000000000000670","article-title":"Metabolic dysfunction-associated steatotic liver disease: Update and impact of new nomenclature on the American Association for the Study of Liver Diseases practice guidance on nonalcoholic fatty liver disease","volume":"79","author":"Kanwal","year":"2024","journal-title":"Hepatology"},{"key":"2026030505115514100_DMM052583C49","doi-asserted-by":"publisher","first-page":"132960","DOI":"10.1155\/2008\/132960","article-title":"Peroxisome proliferator activated receptors and lipoprotein metabolism","volume":"2008","author":"Kersten","year":"2008","journal-title":"PPAR Res."},{"key":"2026030505115514100_DMM052583C50","doi-asserted-by":"publisher","first-page":"dmm050786","DOI":"10.1242\/dmm.050786","article-title":"Optimized methods to image hepatic lipid droplets in zebrafish larvae","volume":"17","author":"Khan","year":"2024","journal-title":"Dis. Model. Mech."},{"key":"2026030505115514100_DMM052583C51","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1002\/aja.1002030302","article-title":"Stages of embryonic development of the zebrafish","volume":"203","author":"Kimmel","year":"1995","journal-title":"Dev. Dyn."},{"key":"2026030505115514100_DMM052583C52","doi-asserted-by":"publisher","first-page":"155664","DOI":"10.1016\/j.metabol.2023.155664","article-title":"From non-alcoholic fatty liver disease (NAFLD) to steatotic liver disease (SLD): an ongoing journey towards refining the terminology for this prevalent metabolic condition and unmet clinical need","volume":"147","author":"Kokkorakis","year":"2023","journal-title":"Metabolism"},{"key":"2026030505115514100_DMM052583C53","doi-asserted-by":"publisher","first-page":"306","DOI":"10.3390\/cells12020306","article-title":"Network regulation of microRNA biogenesis and target interaction","volume":"12","author":"Komatsu","year":"2023","journal-title":"Cells"},{"key":"2026030505115514100_DMM052583C97","doi-asserted-by":"publisher","DOI":"10.1101\/060012","article-title":"Fast gene set enrichment analysis","author":"Korotkevich","year":"2021","journal-title":"bioRxiv"},{"key":"2026030505115514100_DMM052583C54","doi-asserted-by":"publisher","first-page":"60744","DOI":"10.3791\/60744","article-title":"Quantifying liver size in larval zebrafish using brightfield microscopy","volume":"156","author":"Kotiyal","year":"2020","journal-title":"J. Vis. Exp."},{"key":"2026030505115514100_DMM052583C55","doi-asserted-by":"publisher","first-page":"7168","DOI":"10.3390\/ijms24087168","article-title":"An insight into the arising role of MicroRNAs in hepatocellular carcinoma: future diagnostic and therapeutic approaches","volume":"24","author":"Koustas","year":"2023","journal-title":"Int. J. Mol. Sci."},{"key":"2026030505115514100_DMM052583C56","doi-asserted-by":"publisher","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":"2026030505115514100_DMM052583C57","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1016\/S0012-1606(02)00079-9","article-title":"Suppression of lens growth by alphaA-crystallin promoter-driven expression of diphtheria toxin results in disruption of retinal cell organization in zebrafish","volume":"255","author":"Kurita","year":"2003","journal-title":"Dev. Biol."},{"key":"2026030505115514100_DMM052583C58","doi-asserted-by":"publisher","first-page":"519","DOI":"10.3892\/ijo.2011.1215","article-title":"Genome-wide identification of TCF7L2\/TCF4 target miRNAs reveals a role for miR-21 in Wnt-driven epithelial cancer","volume":"40","author":"Lan","year":"2012","journal-title":"Int. J. Oncol."},{"key":"2026030505115514100_DMM052583C59","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1016\/j.cell.2004.12.035","article-title":"Conserved seed pairing, often flanked by adenosines, indicates that thousands of human genes are microRNA targets","volume":"120","author":"Lewis","year":"2005","journal-title":"Cell"},{"key":"2026030505115514100_DMM052583C60","doi-asserted-by":"publisher","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol."},{"key":"2026030505115514100_DMM052583C61","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1016\/j.canlet.2018.08.006","article-title":"Fatty acid oxidation: an emerging facet of metabolic transformation in cancer","volume":"435","author":"Ma","year":"2018","journal-title":"Cancer Lett."},{"key":"2026030505115514100_DMM052583C62","doi-asserted-by":"publisher","first-page":"100642","DOI":"10.1016\/j.jlr.2024.100642","article-title":"Fads2 knockout mice reveal that ALA prevention of hepatic steatosis is dependent on delta-6 desaturase activity","volume":"65","author":"MacLeod","year":"2024","journal-title":"J. Lipid Res."},{"key":"2026030505115514100_DMM052583C63","doi-asserted-by":"publisher","first-page":"776","DOI":"10.1177\/0192623310373778","article-title":"Hepatic enzyme induction: histopathology","volume":"38","author":"Maronpot","year":"2010","journal-title":"Toxicol. Pathol."},{"key":"2026030505115514100_DMM052583C64","doi-asserted-by":"publisher","first-page":"86","DOI":"10.1038\/nature09284","article-title":"OncomiR addiction in an in vivo model of microRNA-21-induced pre-B-cell lymphoma","volume":"467","author":"Medina","year":"2010","journal-title":"Nature"},{"key":"2026030505115514100_DMM052583C65","doi-asserted-by":"publisher","first-page":"228","DOI":"10.1038\/s41419-018-0313-7","article-title":"The carnitine system and cancer metabolic plasticity","volume":"9","author":"Melone","year":"2018","journal-title":"Cell Death Dis."},{"key":"2026030505115514100_DMM052583C66","doi-asserted-by":"publisher","first-page":"1021","DOI":"10.1016\/j.biocel.2009.09.001","article-title":"Role of Wnt\/\u03b2-catenin signaling in liver metabolism and cancer","volume":"43","author":"Monga","year":"2011","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"2026030505115514100_DMM052583C67","doi-asserted-by":"publisher","first-page":"447","DOI":"10.3390\/cancers10110447","article-title":"Lipid metabolic reprogramming in hepatocellular carcinoma","volume":"10","author":"Nakagawa","year":"2018","journal-title":"Cancers"},{"key":"2026030505115514100_DMM052583C68","doi-asserted-by":"publisher","first-page":"164","DOI":"10.1002\/hep.30811","article-title":"Clinical impact of genomic diversity from early to advanced hepatocellular carcinoma","volume":"71","author":"Nault","year":"2020","journal-title":"Hepatology"},{"key":"2026030505115514100_DMM052583C69","doi-asserted-by":"publisher","first-page":"402","DOI":"10.3389\/fendo.2018.00402","article-title":"Overview of MicroRNA biogenesis, mechanisms of actions, and circulation","volume":"9","author":"O'Brien","year":"2018","journal-title":"Front. Endocrinol."},{"key":"2026030505115514100_DMM052583C70","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1002\/hep.1840110114","article-title":"The natural history of nonalcoholic steatohepatitis: a follow-up study of forty-two patients for up to 21 years","volume":"11","author":"Powell","year":"1990","journal-title":"Hepatology"},{"key":"2026030505115514100_DMM052583C71","doi-asserted-by":"publisher","first-page":"5864","DOI":"10.1038\/ncomms6864","article-title":"Dietary cholesterol directly induces acute inflammasome-dependent intestinal inflammation","volume":"5","author":"Progatzky","year":"2014","journal-title":"Nat. Commun."},{"key":"2026030505115514100_DMM052583C72","doi-asserted-by":"publisher","first-page":"1466","DOI":"10.12669\/pjms.35.5.685","article-title":"MicroRNA-21 as a diagnostic marker for hepatocellular carcinoma: a systematic review and meta-analysis","volume":"35","author":"Qu","year":"2019","journal-title":"Pak. J. Med. Sci."},{"key":"2026030505115514100_DMM052583C73","doi-asserted-by":"publisher","first-page":"493","DOI":"10.1007\/BF00316069","article-title":"Quantitation of adipose conversion and triglycerides by staining intracytoplasmic lipids with Oil red O","volume":"97","author":"Ram\u00edrez-Zacar\u00edas","year":"1992","journal-title":"Histochemistry"},{"key":"2026030505115514100_DMM052583C74","doi-asserted-by":"publisher","first-page":"2791","DOI":"10.3390\/cells11182791","article-title":"From molecular mechanisms to therapeutics: understanding MicroRNA-21 in cancer","volume":"11","author":"Rhim","year":"2022","journal-title":"Cells"},{"key":"2026030505115514100_DMM052583C75","doi-asserted-by":"publisher","first-page":"e2825","DOI":"10.1038\/cddis.2017.246","article-title":"miR-21 ablation and obeticholic acid ameliorate nonalcoholic steatohepatitis in mice","volume":"8","author":"Rodrigues","year":"2017","journal-title":"Cell Death Dis."},{"key":"2026030505115514100_DMM052583C76","doi-asserted-by":"publisher","first-page":"2256","DOI":"10.1111\/liv.15682","article-title":"miR-21-5p promotes NASH-related hepatocarcinogenesis","volume":"43","author":"Rodrigues","year":"2023","journal-title":"Liver Int."},{"key":"2026030505115514100_DMM052583C77","doi-asserted-by":"publisher","first-page":"1409","DOI":"10.1194\/jlr.M700537-JLR200","article-title":"Intracellular cholesterol transporter StarD4 binds free cholesterol and increases cholesteryl ester formation","volume":"49","author":"Rodriguez-Agudo","year":"2008","journal-title":"J. Lipid Res."},{"key":"2026030505115514100_DMM052583C78","doi-asserted-by":"publisher","first-page":"1598","DOI":"10.1016\/j.jhep.2022.08.021","article-title":"Global burden of primary liver cancer in 2020 and predictions to 2040","volume":"77","author":"Rumgay","year":"2022","journal-title":"J. Hepatol."},{"key":"2026030505115514100_DMM052583C79","doi-asserted-by":"publisher","first-page":"322","DOI":"10.1136\/gutjnl-2017-315448","article-title":"\u03b2-catenin-activated hepatocellular carcinomas are addicted to fatty acids","volume":"68","author":"Senni","year":"2019","journal-title":"Gut"},{"key":"2026030505115514100_DMM052583C80","doi-asserted-by":"publisher","first-page":"710","DOI":"10.1038\/nrendo.2017.91","article-title":"SREBP-regulated lipid metabolism: convergent physiology \u2013 divergent pathophysiology","volume":"13","author":"Shimano","year":"2017","journal-title":"Nat. Rev. Endocrinol."},{"key":"2026030505115514100_DMM052583C81","doi-asserted-by":"publisher","first-page":"G462","DOI":"10.1152\/ajpgi.00358.2018","article-title":"Hepatic transcriptome signatures in patients with varying degrees of nonalcoholic fatty liver disease compared with healthy normal-weight individuals","volume":"316","author":"Suppli","year":"2019","journal-title":"Am. J. Physiol. Gastroint. Liver Physiol."},{"key":"2026030505115514100_DMM052583C82","volume-title":"Target Organ Pathology: A Basic Text","author":"Target Organ Pathology: A Basic Text","year":"1998"},{"key":"2026030505115514100_DMM052583C83","doi-asserted-by":"publisher","first-page":"142","DOI":"10.2174\/1566523214666141224095610","article-title":"miR-122 is a unique molecule with great potential in diagnosis, prognosis of liver disease, and therapy both as miRNA mimic and antimir","volume":"15","author":"Thakral","year":"2015","journal-title":"Curr. Gene Ther."},{"key":"2026030505115514100_DMM052583C84","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1016\/S0925-4773(02)00218-6","article-title":"I-SceI meganuclease mediates highly efficient transgenesis in fish","volume":"118","author":"Thermes","year":"2002","journal-title":"Mech. Dev."},{"key":"2026030505115514100_DMM052583C85","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1038\/s43856-024-00465-3","article-title":"Transcriptomics-driven metabolic pathway analysis reveals similar alterations in lipid metabolism in mouse MASH model and human","volume":"4","author":"Tsouka","year":"2024","journal-title":"Commun. Med."},{"key":"2026030505115514100_DMM052583C86","doi-asserted-by":"publisher","first-page":"2054","DOI":"10.1101\/gr.139733.112","article-title":"Extensive alternative polyadenylation during zebrafish development","volume":"22","author":"Ulitsky","year":"2012","journal-title":"Genome Res."},{"key":"2026030505115514100_DMM052583C87","doi-asserted-by":"publisher","first-page":"159514","DOI":"10.1016\/j.bbalip.2024.159514","article-title":"Phospholipid isotope tracing suggests \u03b2-catenin-driven suppression of phosphatidylcholine metabolism in hepatocellular carcinoma","volume":"1869","author":"VanSant-Webb","year":"2024","journal-title":"Biochim. Biophys. Acta Mol. Cell Biol. Lipids"},{"key":"2026030505115514100_DMM052583C88","doi-asserted-by":"publisher","first-page":"2501","DOI":"10.1161\/ATVBAHA.113.301948","article-title":"Apolipoprotein A-IV expression in mouse liver enhances triglyceride secretion and reduces hepatic lipid content by promoting very low density lipoprotein particle expansion","volume":"33","author":"VerHague","year":"2013","journal-title":"Arterioscler. Thromb. Vasc. Biol."},{"key":"2026030505115514100_DMM052583C89","doi-asserted-by":"publisher","first-page":"2257","DOI":"10.1073\/pnas.0510565103","article-title":"A microRNA expression signature of human solid tumors defines cancer gene targets","volume":"103","author":"Volinia","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2026030505115514100_DMM052583C90","doi-asserted-by":"publisher","first-page":"16923","DOI":"10.1038\/srep16923","article-title":"Efficient test and visualization of multi-set intersections","volume":"5","author":"Wang","year":"2015","journal-title":"Sci. Rep."},{"key":"2026030505115514100_DMM052583C91","doi-asserted-by":"publisher","first-page":"1303","DOI":"10.1097\/HEP.0000000000000786","article-title":"Molecular mechanisms in MASLD\/MASH-related HCC","volume":"82","author":"Wang","year":"2025","journal-title":"Hepatology"},{"key":"2026030505115514100_DMM052583C92","doi-asserted-by":"publisher","first-page":"e1008941","DOI":"10.1371\/journal.pgen.1008941","article-title":"A point mutation decouples the lipid transfer activities of microsomal triglyceride transfer protein","volume":"16","author":"Wilson","year":"2020","journal-title":"PLoS Genet."},{"key":"2026030505115514100_DMM052583C93","doi-asserted-by":"publisher","first-page":"e154515","DOI":"10.1172\/JCI154515","article-title":"\u03b2-Catenin signaling in hepatocellular carcinoma","volume":"132","author":"Xu","year":"2022","journal-title":"J. Clin. Invest."},{"key":"2026030505115514100_DMM052583C94","doi-asserted-by":"publisher","first-page":"884066","DOI":"10.3389\/fonc.2022.884066","article-title":"ELOVLs predict distinct prognosis value and immunotherapy efficacy in patients with hepatocellular carcinoma","volume":"12","author":"Zhang","year":"2022","journal-title":"Front. Oncol."},{"key":"2026030505115514100_DMM052583C95","doi-asserted-by":"publisher","first-page":"1810","DOI":"10.1016\/j.celrep.2018.01.058","article-title":"MicroRNA-223 suppresses the canonical NF-\u03baB pathway in basal keratinocytes to dampen neutrophilic inflammation","volume":"22","author":"Zhou","year":"2018","journal-title":"Cell Rep."}],"container-title":["Disease Models &amp; Mechanisms"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/dmm\/article-pdf\/doi\/10.1242\/dmm.052583\/3682294\/dmm052583.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/dmm\/article-pdf\/doi\/10.1242\/dmm.052583\/3688794\/dmm052583.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/dmm\/article-pdf\/doi\/10.1242\/dmm.052583\/3688794\/dmm052583.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,3,5]],"date-time":"2026-03-05T10:12:05Z","timestamp":1772705525000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/dmm\/article\/19\/2\/dmm052583\/370942\/MicroRNA-21-promotes-dysregulated-lipid-metabolism"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,1]]},"references-count":95,"journal-issue":{"issue":"2","published-print":{"date-parts":[[2026,2,1]]}},"URL":"https:\/\/doi.org\/10.1242\/dmm.052583","relation":{},"ISSN":["1754-8403","1754-8411"],"issn-type":[{"value":"1754-8403","type":"print"},{"value":"1754-8411","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,1]]},"article-number":"dmm052583"}}