About the Authors

Hazel Wilkie

Affiliation The Roslin Institute & R(D)SVS, University of Edinburgh, Easter Bush, Midlothian, EH25 9RG, United Kingdom

Anton Gossner

Affiliation The Roslin Institute & R(D)SVS, University of Edinburgh, Easter Bush, Midlothian, EH25 9RG, United Kingdom

Stephen Bishop

† Deceased.

Affiliation The Roslin Institute & R(D)SVS, University of Edinburgh, Easter Bush, Midlothian, EH25 9RG, United Kingdom

John Hopkins

john.hopkins@ed.ac.uk

Affiliation The Roslin Institute & R(D)SVS, University of Edinburgh, Easter Bush, Midlothian, EH25 9RG, United Kingdom

Competing Interests

This project was funded by the Biotechnology and Biological Sciences Research Council (BBSRC) Institute Strategic Programme Grant to The Roslin Institute and by a BBSRC Industrial CASE scholarship to JH and SB, for HW, supported by Sheep Improved Genetics Ltd (no funding was obtained from SIG Ltd) and AHDB Beef and Lamb (Grant BB/J012653/1). This does not alter the authors' adherence to PLOS ONE policies on sharing data. The initial work was funded by the Natural Environment Research Council (NERC) through its ‘Post-Genomics and Proteomics’ programme (grant NE/D000645/1).

Author Contributions

Conceived and designed the experiments: JH SB AG. Performed the experiments: HW JH. Analyzed the data: HW AG JH. Contributed reagents/materials/analysis tools: JH AG. Wrote the paper: JH AG HW. Played a major role in obtaining funding: SB.