Department of Cellular and Molecular Physiology, Yale University School of Medicine, 333 Cedar Street, New Haven, CT 06510, USA; Department of Neuroscience, Yale University School of Medicine, 333 Cedar Street, New Haven, CT 06510, USA; Program in Cellular Neuroscience, Neurodegeneration and Repair, Yale University School of Medicine, 333 Cedar Street, New Haven, CT 06510, USA.