1. Centre for Mathematical Biology, Mathematical Institute, University of Oxford, 24-29 St Giles', Oxford OX1 3PN, UK
2. Oxford Centre for Integrative Systems Biology, Department of Biochemistry, University of Oxford, South Parks Road OX1 3QU, UK
3. Center for Developmental Biology, RIKEN, Minatojima-minami 2-2-3, Kobe 650-0047, Japan.