GRHL2 and Encephalocele: However, the expression pattern of Grhl2 (25,49), and the association of Grhl2 loss of function with comparable malformations that we observe in Axd mutants (midline fusion defects, cleft palate and a brain abnormality that resembles encephalocele (26)), strongly suggest that the anomalies observed in Axd mutant embryos result from over-expression of Grhl2 as shown for spinal neurulation defects in this strain (25).