1. Institute of Cell Biology, School of Biological Sciences, University of Edinburgh 1 , Edinburgh EH9 3FF , UK
2. Max Planck Institute for Biology of Ageing 2 , Joseph-Stelzmann-Strasse 9b, Cologne 50931 , Germany
3. Wellcome Centre for Cell Biology, Institute of Cell Biology, School of Biological Sciences, University of Edinburgh 3 , Edinburgh EH9 3BF , UK
4. Institute of Molecular Plant Sciences, School of Biological Sciences, University of Edinburgh 4 , Edinburgh EH9 3BF , UK