HNF4A and cancer: Among the target genes activated by Wnt/β-catenin signaling and SOX9 is the oncogene MYC. In CRC cells, SOX9 is associated with many WREs, including several cancer risk-associated enhancers of MYC. Our characterization of the cis-regulatory grammar that allows SOX9 to activate Wnt target genes reveals the presence of SOX9-binding sites in the c-myc-335 WRE, which regulate enhancer activity along with TCF sites.