[1] Univ Edinburgh, Roslin Inst, Div Genet & Genom, Edinburgh EH25 9RG, Midlothian, Scotland
[2] Univ Edinburgh, Royal Dick Sch Vet Studies, Edinburgh EH25 9RG, Midlothian, Scotland
[3] Swedish Univ Agr Sci, Dept Anim Breeding & Genet, POB 7023, S-75007 Uppsala, Sweden
[4] Swedish Armed Forces Dog Training Ctr, POB 194, S-19524 Marsta, Sweden
[5] Scotlands Rural Coll, Anim & Vet Sci Grp, Edinburgh EH25 9RG, Midlothian, Scotland