Whitehead Institute for Biomedical Research, 9 Cambridge Center, Cambridge, MA 02142, Harvard-MIT Division of Health Sciences and Technology, MIT, Cambridge, MA 02139 and Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA 02142, USA Whitehead Institute for Biomedical Research, 9 Cambridge Center, Cambridge, MA 02142, Harvard-MIT Division of Health Sciences and Technology, MIT, Cambridge, MA 02139 and Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA 02142, USA Whitehead Institute for Biomedical Research, 9 Cambridge Center, Cambridge, MA 02142, Harvard-MIT Division of Health Sciences and Technology, MIT, Cambridge, MA 02139 and Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA 02142, USA.