[1] Univ Edinburgh, Roslin Inst, Edinburgh, Midlothian, Scotland
[2] Univ Edinburgh, R D SVS, Edinburgh, Midlothian, Scotland
[3] Univ Edinburgh, Edinburgh Genom, Edinburgh, Midlothian, Scotland
[4] Ctr Environm Fisheries & Aquaculture Sci, Cefas Weymouth Lab, Weymouth, Scotland