MEIS2 and major depressive disorder: Conversely, accessible regions significantly associated with MDD were enriched for the MEIS2, OTX2 and GATA2-binding motifs, indicating a midbrain GABAergic identity (Fig. 5c and Extended Data Fig. 9b), with 29 of the 114 significant MDD regions also containing CNN-predicted OTX2-binding sites and 46 containing predicted GATA2-binding sites.