Department of Biology, College of Arts and Sciences, University of Pennsylvania, Philadelphia, PA 19143, USA, Department of Pharmacology and Department of Genetics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA 19143, USA.|Department of Biology, College of Arts and Sciences, University of Pennsylvania, Philadelphia, PA 19143, USA, Department of Pharmacology and Department of Genetics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA 19143, USA Department of Biology, College of Arts and Sciences, University of Pennsylvania, Philadelphia, PA 19143, USA, Department of Pharmacology and Department of Genetics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA 19143, USA.