[1] Univ Edinburgh, Inst Cell & Mol Biol, Edinburgh EH9 3JH, Midlothian, Scotland
[2] RIKEN, Tsukuba Inst, Plant Mol Biol Lab, Tsukuba, Ibaraki 3050074, Japan
[3] RIKEN Genom Sci Ctr, Plant Mutat Explorat Team, Plant Funct Genom Res Grp, Tsurumi Ku, Kanagawa 2300045, Japan
[4] JST, CREST, Osaka, Japan