[1] Scottish Agr Coll, Sustainable Livestock Syst Grp, Edinburgh EH9 3JG, Midlothian, Scotland
[2] Scottish Agr Coll, Anim Hlth Grp, Edinburgh EH9 3JG, Midlothian, Scotland
[3] Univ Edinburgh, Roslin Inst, Easter Bush EH25 9RG, Midlothian, Scotland
[4] Minist Ciencia & Innovac, Dept Mejora Genet Anim, Inst Nacl Invest & Tecnol Agr & Alimentaria, Madrid 28040, Spain