[1] Tohoku Univ, Grad Sch Agr Sci, Dept Life Sci, Sendai, Miyagi 9818555, Japan
[2] Natl Agr Res Ctr Hokkaido Reg, Dept Upland Agr, Memuro, Hokkaido 0820071, Japan
[3] Tohoku Univ, Grad Sch Agr Sci, Dept Biol Resource Sci, Sendai, Miyagi 9818555, Japan
[4] Nagoya Univ, Grad Sch Bioagr Sci, Lab Def Plant Pathogen Interact, Nagoya, Aichi 4648601, Japan
[5] John Innes Ctr Plant Sci Res, Sainsbury Lab, Norwich NR4 7UH, Norfolk, England