[1] Kobe Univ, Sch Med, Dept Biomed Regulat, Chuo Ku, Kobe, Hyogo 6500017, Japan
[2] Kyoto Univ, Grad Sch Sci, Ctr Mol & Dev Biol, Sakyo Ku, Kyoto 6068502, Japan
[3] Kobe Univ, Sch Med, Dept Anat, Chuo Ku, Kobe, Hyogo 6500017, Japan
[4] Japan Sci & Technol Corp, JST, ERATO, Kondoh Differentiat Signalling Project,Sakyo Ku, Kyoto 6068305, Japan