1. Division of Plant and Crop Sciences, School of Biosciences, University of Nottingham, Sutton Bonington Campus , Loughborough, LE12 5RD, UK
2. Department of Biology, University of Fribourg , Fribourg CH-1700, Switzerland
3. Centre for Mathematical Medicine and Biology, School of Mathematical Sciences, University of Nottingham , Nottingham NG7 2RD, UK