Using a murine model with constitutional Smoothened mutant protein (SmoM2) expression (K14-CreER/Rosa-SmoM2 mice) that resulted in BCC development, they discovered increased Hedgehog signaling activity with YAP and GLI2 (a downstream target of Hedgehog signaling in the skin) nuclear expression accompanied by the increased expression of certain YAP target genes including Ctgf [36]. Here, GLI2 is linked to skin basal cell carcinoma.