[1] Kyoto Univ, Fac Med, Dept Pharmacol, Kyoto 6068501, Japan
[2] Kyoto Univ, Fac Med, Horizontal Med Res Org, Kyoto 6068501, Japan
[3] RIKEN, Lab Cellular Morphogenesis, Ctr Dev Biol, Kobe, Hyogo 6500047, Japan
[4] Univ Minnesota, Dept Genet Cell Biol & Dev, Minneapolis, MN 55455 USA