Donnelly Centre for Cellular & Biomolecular Research, University of Toronto, M5S 3E1, Toronto, ON, Canada, Department of Molecular Genetics, University of Toronto, M5S 1A8, Toronto, ON, Canada, Genetics and Genome Biology, Hospital for Sick Children, M5G 0A4, Toronto, ON, Canada and Banting and Best Department of Medical Research, University of Toronto, M5G 1L6, Toronto, ON, Canada.