[1] Newcastle Univ, Comparat Biol Ctr, Framlington Pl, Newcastle Upon Tyne NE2 4HH, Tyne & Wear, England
[2] Univ Liege, Fac Vet Med, Blvd Colonster, B-4000 Liege, Belgium
[3] Univ Nottingham, Sch Vet Med & Sci, Sutton Bonington Campus, Loughborough LE12 5RD, Leics, England
[4] Newcastle Univ, Inst Neurosci, Newcastle Upon Tyne NE1 7RU, Tyne & Wear, England