Moreover, Sox4- and Sox11-deficient animal models display coloboma, cardiac malformations and brain defects like those seen in CHARGE syndrome (Wurm et al., 2008; Paul et al., 2014; Pillai-Kastoori et al., 2014; Schulz et al., 2014; Gnedeva and Hudspeth, 2015; Wen et al., 2015). Here, SOX11 is linked to coloboma.