We tested four of the hits from this screen (S1 ExtendedData Table, S2 Extended Data Table): Pax2, which is known to be a target for mutations that cause similar ocular and kidney phenotypes as the loss of Capn15 [38,39], Pax5, where haploinsufficiencies cause neurodevelopmental disorders [40], Runx2, a transcription factor implicated in bone development also may play a role in developmental patterning [41]and Smarca4, a component of the mammalian SWI/SNF complex linked to neurodevelopmental disorders such as Coffin Sirius syndrome [42] and eye development [43]. Here, RUNX2 is linked to neurodevelopmental disorder.