[1] Univ Tokyo, Grad Sch Agr & Life Sci, Tokyo 1138657, Japan
[2] Japan Int Res Ctr Agr Sci, Biol Resources Div, Tsukuba, Ibaraki 3058686, Japan
[3] RIKEN, Plant Sci Ctr, Yokohama, Kanagawa 2300045, Japan
[4] Japan Sci & Technol Agcy, Core Res Evolut Sci & Technol, Kawaguchi, Saitama 3320012, Japan