[1] Univ Edinburgh, Roslin Inst, Easter Bush EH25 9RG, Midlothian, Scotland
[2] Univ Edinburgh, Royal Dick Sch Vet Studies, Easter Bush EH25 9RG, Midlothian, Scotland
[3] Pirbright Inst, Genet & Genom Grp, Woking GU24 0NF, Surrey, England
[4] Ctr Cellular & Mol Physiol, Nuffield Dept Clin Med, Jenner Inst, Roosevelt Dr, Oxford OX3 7BN, England
[5] Univ Nottingham, Sch Vet Med & Sci, Sutton Bonington Campus, Loughborough LE12 5RD, Leics, England
[6] Scotlands Rural Coll, Easter Bush EH25 9RG, Midlothian, Scotland