About the Authors

Charley Xia

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom

Carmen Amador

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom

Jennifer Huffman

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom

Holly Trochet

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom

Archie Campbell

Affiliation Generation Scotland, Centre for Genomic and Experimental Medicine, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom

David Porteous

Affiliation Generation Scotland, Centre for Genomic and Experimental Medicine, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom

Generation Scotland

Membership of the Generation Scotland team is provided in the Acknowledgments.

Affiliation A collaboration between the University Medical School and NHS in Aberdeen, Dundee, Edinburgh and Glasgow, Scotland, United Kingdom

Nicholas D. Hastie

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom

Caroline Hayward

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom

Veronique Vitart

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom

Pau Navarro

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom

Chris S. Haley

chris.haley@igmm.ed.ac.uk; chris.haley@roslin.ed.ac.uk

Affiliations MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom, The Roslin Institute and R(D)SVS, University of Edinburgh, Midlothian, United Kingdom

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: CX CA PN CSH. Performed the experiments: CX CA. Analyzed the data: CX. Contributed reagents/materials/analysis tools: CX CA PN CSH NDH. Wrote the paper: CX CA PN CSH. Recruited participants: HT AC. Collected and prepared data: JH HT AC DP CH VV. Raised funding: CH DP NDH.