[1] Harvard Univ, Sch Med, Dept Cell Biol, Boston, MA 02115 USA
[2] Harvard Univ, Sch Med, Dept Ophthalmol, Boston, MA 02114 USA
[3] Russian Acad Sci, Sechenov Inst, St Petersburg 196140, Russia
[4] Brandeis Univ, Dept Biol, Waltham, MA 02454 USA
[5] Brandeis Univ, Rosenstiel Basic Med Sci Res Ctr, Waltham, MA 02454 USA