[1] Univ Edinburgh, Wellcome Trust Ctr Cell Biol, Edinburgh EH9 3JR, Midlothian, Scotland
[2] Univ Edinburgh, Inst Cell Biol, Sch Biol Sci, Edinburgh EH9 3JR, Midlothian, Scotland
[3] Baylor Coll Med, Dept Mol & Human Genet, Houston, TX 77030 USA
[4] MRC Human Genet Unit, Edinburgh EH4 2XU, Midlothian, Scotland