About the Authors

Richard J. Leach

richard.j.leach@ars.usda.gov

Current address: United States Department of Agriculture, Agricultural Research Service, United States Meat Animal Research Center, Clay Center, Nebraska, United States of America

Affiliation Department of Genetics and Genomics, The Roslin Institute and Royal (Dick) School of Veterinary Studies, The University of Edinburgh, Easter Bush, Midlothian, United Kingdom

Ronan G. O'Neill

Affiliation Department of Agriculture, Fisheries and Food, Celbridge, Co. Kildare, Ireland

Julie L. Fitzpatrick

Affiliation Moredun Research Institute, Edinburgh, United Kingdom,

John L. Williams

Current address: Parco Tecnologico Padano, Polo Universitario, Lodi, Italy

Affiliation Department of Genetics and Genomics, The Roslin Institute and Royal (Dick) School of Veterinary Studies, The University of Edinburgh, Easter Bush, Midlothian, United Kingdom

Elizabeth J. Glass

Affiliation Department of Genetics and Genomics, The Roslin Institute and Royal (Dick) School of Veterinary Studies, The University of Edinburgh, Easter Bush, Midlothian, United Kingdom

Competing Interests

The authors have the following interest: Pfizer Animal Health provided a proportion of the funding for this investigation. This does not alter the authors' adherence to all the PLoS ONE policies on sharing data and materials.

Author Contributions

Conceived and designed the experiments: JLW JLF EJG. Performed the experiments: RGO. Analyzed the data: RJL. Wrote the paper: RJL. Helped to edit the manuscript: EJG JLW RGO JLF.