HSPG2 and hepatocellular carcinoma: Aggressive hepatoma cells (SK-Hep1 and HA 22T), which express high levels of the VEGF protein, showed enhanced recruitment of the SPZ1–TWIST1 complex to the VEGF promoter compared with that observed in cells with low or no VEGF (Hep 3B and PLC, respectively) expression (Fig. 3h).