[1] Kyoto Univ, Fac Med, JST, CREST Res Project,Sakyo Ku, Kyoto 6068501, Japan
[2] Kyoto Univ, Grad Sch Biostudies, Dept Gene Mech, Sakyo Ku, Kyoto 6068502, Japan
[3] Univ Edinburgh, Inst Cell & Mol Biol, Wellcome Trust Ctr Cell Biol, Edinburgh EH9 3JR, Midlothian, Scotland
[4] Niigata Univ, Fac Med, Dept Anat & Histol, Niigata 9518510, Japan
[5] Grad Univ Adv Studies, JST, PRESTO, Shizuoka 4118540, Japan
[6] Res Inst Mol Pathol, A-1030 Vienna, Austria