Required for cilia formation during embryonal development. Acts as a negative regulator of Shh signaling. Component of the IFT complex A (IFT-A) complex. Reduced number of cilia in embryonal pronephric duct and shortened primary cilia in Kupffer's vesicle. Embryos show shortened body axis and curvature, cardiac edema, small eyes, pronephric cysts and a distended cranium; shh signaling is reduced.