SMAD4 and hepatocellular carcinoma: In addition to these previous findings, we revealed that ACVR2A impairment increased the expression levels of genes involved in the glycolysis pathway via the activin/SMAD signal transduction in non-viral HCC (Figures 2A–2D) and identified that SMAD4 repressed LDHA expression by directly binding to the promoter region (Figures 5I and 5J).