Zhenyu Gu, Masatoshi Nomura, Brenda B. Simpson, Hong Lei, Alie Feijen, Janny van den Eijnden-van Raaij, Patricia K. Donahoe, and En Li Cardiovascular Research Center, Massachusetts General Hospital East, Department of Medicine, Harvard Medical School, Charlestown, Massachusetts 02129, USA; Pediatric Surgical Research Laboratory, Massachusetts General Hospital, Department of Surgery, Harvard Medical School, Boston, Massachusetts 02114, USA; Hubrecht Laboratory, Netherlands Institute for Developmental Biology, Utrecht, The Netherlands