1. Wolfson Centre for Mathematical Biology, Mathematical Institute, University of Oxford, Andrew Wiles Building, Radcliffe Observatory Quarter, Woodstock Road, Oxford OX2 6GG, UK
2. Department of Civil, Environmental and Geomatic Engineering, Faculty of Engineering Science, University College London, Chadwick Building, Gower Street, London WC1E 6BT, UK
3. Oxford Centre for Integrative Systems Biology and Department of Biochemistry, University of Oxford, South Parks Road, Oxford OX1 3QU, UK