[1] Univ Exeter, Inst Clin Sci, Div Mol Genet, Exeter EX4 4QJ, Devon, England
[2] Univ Chicago, Howard Hughes Med Inst, Chicago, IL 60637 USA
[3] Univ Chicago, Dept Biochem & Mol Biol, Chicago, IL 60637 USA
[4] Univ Chicago, Dept Med, Chicago, IL 60637 USA