LGR4 and gastric cancer: Analysis of proteins associated with the AKT pathway in GC cells overexpressing or downregulating ONECUT2 found that ONECUT2 can inhibit PP2A phosphatase activity and increase AKT(S473)/β-catenin(S552) phosphorylation, resulting in detectable changes in genes whose transcription is upregulated downstream of β-catenin, including CCND1 and the stemness-related factor LGR4 (Figs. 5C, 4C).