[1] Hokkaido Univ, Inst Low Temp Sci, Sapporo, Hokkaido, Japan
[2] Hokkaido Univ, Grad Sch Environm Sci, Sapporo, Hokkaido, Japan
[3] Hokkaido Univ, Arctic Res Ctr, Sapporo, Hokkaido, Japan
[4] Japan Agcy Marine Earth Sci & Technol, Yokosuka, Kanagawa, Japan
[5] Tokyo Univ Marine Sci & Technol, Tokyo, Japan
[6] Japan Fisheries Res & Educ Agcy, Yokohama, Kanagawa, Japan