[1] Hokkaido Univ, Grad Sch Agr, Lab Pathogen Plant Interact, Kita Ku, Sapporo, Hokkaido 0608589, Japan
[2] Sapporo Breweries Ltd, Hop Res Dept, Plant Bioengn Res Labs, Sapporo, Hokkaido, Japan
[3] Hirosaki Univ, Fac Agr & Life Sci, Dept Environm Biol, Hirosaki, Aomori, Japan