1. Nakagawa Initiative Research Unit, RIKEN, 2-1 Hirosawa, Wako 351-0198, Japan
2. RIKEN Center for Developmental Biology, 2-2-3 Minatojima Minamimachi, Chuo-ku, Kobe 650-0047, Japan
3. Department of Cell and Developmental Biology, Graduate School of Biostudies, Kyoto University, Kitashirakawa, Sakyo-ku, Kyoto 606-8502, Japan