[1] Queens Univ, Dept Biol, Kingston, ON K7L 3N6, Canada
[2] Cornell Univ, Dept Mol Biol & Genet, Ithaca, NY 14853 USA
[3] Univ Penn, Dept Biol, Philadelphia, PA 19104 USA
[4] Univ Penn, Dept Cell & Dev Biol, Philadelphia, PA 19104 USA
[5] Univ Toronto, Banting & Best Dept Med Res, Toronto, ON M5G 1L6, Canada
[6] Univ Toronto, Dept Mol & Med Genet, Toronto, ON M5G 1L6, Canada