Conversely, hyperactivation of the Hh pathway through overexpression of a constitutively activated form of ci or smo was sufficient to synergize with ykiACT, resulting in tumor overgrowth, enhanced proliferation, VNC invasion, and upregulation of Mmp1 (Figs. 3D–F and EV3J–O), phenocopying that of ykiACT/Toll-6ACT tumors. Here, SMO is linked to neoplasm.