1. UCL Institute of Ophthalmology, London EC1V 9EL, UK
2. Nuffield Laboratory of Ophthalmology, Nuffield Department of Clinical Neurosciences, University of Oxford, John Radcliffe Hospital, Oxford OX3 9DU, UK
3. UMBC Department of Biological Sciences, Baltimore, MD 21250, USA
4. School of Animal Biology and Oceans Institute, University of Western Australia, Perth, WA 6009, Australia