1. Department of Applied Sciences, Centre for Research in Biosciences, University of the West of England , Bristol BS16 1QY , UK
2. Cell Biology Laboratories, School of Biochemistry, University of Bristol , Bristol BS8 1TD , UK
3. Department of Physiology and Trinity College Institute for Neuroscience, Trinity College Dublin , Dublin 2 , Ireland
4. Bristol Medical School, Faculty of Health Sciences, University of Bristol , Bristol BS1 3NY , UK