About the Authors

Scott Kilpatrick

Affiliation Royal (Dick) School of Veterinary Studies and The Roslin Institute, The University of Edinburgh, Hospital for Small Animals, Easter Bush Veterinary Centre, Roslin, Midlothian, United Kingdom

Margaret Dreistadt

Affiliation Royal (Dick) School of Veterinary Studies and The Roslin Institute, The University of Edinburgh, Hospital for Small Animals, Easter Bush Veterinary Centre, Roslin, Midlothian, United Kingdom

Polly Frowde

Affiliation Davies Veterinary Specialists Limited, Manor Farm Business Park, Higham Gobion, Herts, United Kingdom

Roger Powell

Affiliation PTDS, Unit 2a, Manor Farm Business Park, Higham Gobion, Herts, United Kingdom

Elspeth Milne

Affiliation Royal (Dick) School of Veterinary Studies and The Roslin Institute, The University of Edinburgh, Hospital for Small Animals, Easter Bush Veterinary Centre, Roslin, Midlothian, United Kingdom

Sionagh Smith

Affiliation Royal (Dick) School of Veterinary Studies and The Roslin Institute, The University of Edinburgh, Hospital for Small Animals, Easter Bush Veterinary Centre, Roslin, Midlothian, United Kingdom

Linda Morrison

Affiliation Royal (Dick) School of Veterinary Studies and The Roslin Institute, The University of Edinburgh, Hospital for Small Animals, Easter Bush Veterinary Centre, Roslin, Midlothian, United Kingdom

Adam G. Gow

Affiliation Royal (Dick) School of Veterinary Studies and The Roslin Institute, The University of Edinburgh, Hospital for Small Animals, Easter Bush Veterinary Centre, Roslin, Midlothian, United Kingdom

Ian Handel

Affiliation Royal (Dick) School of Veterinary Studies and The Roslin Institute, The University of Edinburgh, Hospital for Small Animals, Easter Bush Veterinary Centre, Roslin, Midlothian, United Kingdom

Richard J. Mellanby

Richard.Mellanby@ed.ac.uk

Affiliation Royal (Dick) School of Veterinary Studies and The Roslin Institute, The University of Edinburgh, Hospital for Small Animals, Easter Bush Veterinary Centre, Roslin, Midlothian, United Kingdom

Competing Interests

The authors’ affiliation with the funders does not alter the authors’ adherence to PLOS ONE policies on sharing data and materials.

Author Contributions

Conceived and designed the experiments: RJM SK. Performed the experiments: SK RJM. Analyzed the data: SK RJM IH. Contributed reagents/materials/analysis tools: SK MD PF RP EM SS AGG IH RJM LM. Wrote the paper: SK MD PF RP EM SS AGG IH LM RJM.