[1] Carnegie Inst Washington, Dept Embryol, Baltimore, MD 21218 USA
[2] Howard Hughes Med Inst, Baltimore, MD 21218 USA
[3] Johns Hopkins Univ, Dept Biol, Baltimore, MD 21218 USA
[4] Northwestern Univ, Feinberg Sch Med, Dept Cell & Mol Biol, Chicago, IL 60611 USA