1. Howard Hughes Medical Institute and Department of Cell and Developmental Biology, Vanderbilt University School of Medicine, Nashville, TN 37232, USA.
2. Department of Biological Sciences, Department of Cell and Developmental Biology, Department of Pediatrics, Vanderbilt University, Nashville, TN 37235, USA.
3. Department of Developmental Biology, Washington University School of Medicine, St Louis, MO 63110, USA.