[1] Univ Edinburgh, Sch Phys & Astron, Peter Guthrie Tait Rd, Edinburgh EH9 3FD, Midlothian, Scotland
[2] Univ Edinburgh, Western Gen Hosp, MRC Human Genet Unit, Inst Genet & Mol Med, Crewe Rd, Edinburgh EH4 2XU, Midlothian, Scotland
[3] Univ Edinburgh, EPCC, Peter Guthrie Tait Rd, Edinburgh EH9 3FD, Midlothian, Scotland