[1] Kitasato Univ, Sch Fisheries Sci, Sanriku, Iwate 0220101, Japan
[2] Univ Tokyo, Grad Sch Agr & Life Sci, Dept Aquat Biosci, Tokyo 1138657, Japan
[3] Natl Res Inst Aquaculture, Nikko Branch, Nikko, Tochigi 3211661, Japan
[4] Natl Res Inst Aquaculture, Tamaki, Mie 5190423, Japan