[1] Univ Utah, Hlth Sci Ctr, Dept Neurobiol & Anat, John A Moran Eye Ctr, Salt Lake City, UT 84132 USA
[2] Univ Utah, John A Moran Eye Ctr, Dept Ophthalmol & Visual Sci, Salt Lake City, UT 84132 USA
[3] Netherlands Inst Dev Biol, Ctr Biomed Res, Hubrecht Inst, Utrecht, Netherlands
[4] Kyoto Univ, Grad Sch Med, Dept Pharmacol, Kyoto, Japan