Aldh1a3-/- (Raldh3) knockout mice display coloboma, and although Aldh1a1-/- (Raldh1) knockout mice did not have a recognizable ophthalmic phenotype, double knockout Aldh1a1-/-; Alldh1a3-/- mice displayed an even more severe eye phenotype [26]. Here, ALDH1A3 is linked to coloboma.