COL1A1 and fatty liver disease: However, feeding with an HFD induced more severe hepatic steatosis and fibrogenesis in Foxo1/3/4 triple knockout mice compared to wild-type mice via upregulating profibrotic genes such as C-C motif chemokine ligand 2 (CCL2), alpha-1 type I collagen (Col1A1), and TGF-β [56].