[1] Kyoto Univ, Grad Sch Sci, Dept Zool, Sakyo Ku, Kyoto 6068502, Japan
[2] Japan Sci & Technol Agcy JST, CREST, Tokyo, Japan
[3] RIKEN, Ctr Dev Biol, Chuo Ku, Kobe, Hyogo 6500047, Japan
[4] Kobe Univ, Grad Sch Med, Dept Physiol & Cell Biol, Chuo Ku, Kobe, Hyogo 6500017, Japan