SHH and omphalocele: This finding suggests that hedgehog signaling could contribute to the gut rotation phenotype as loss of SHH signaling has previously been identified as causative of omphalocele (Matsumaru et al., 2011; Negretti et al., 2022) and is a regulator of Pitx2; however, a direct SHH role in Hand1Hand2/Hand2 needs to be confirmed at earlier stages to make an informed conclusion.