1. The Walter and Eliza Hall Institute of Medical Research, Parkville, Victoria 3052, Australia
2. Department of Medical Biology, University of Melbourne, Parkville, Victoria 3010, Australia
3. Department of Biological Sciences, University of Cape Town, RSA-7700 Rondebosch, South Africa
4. Department of Microbiology and Immunology, University of Melbourne at the Peter Doherty Institute for Infection and Immunity, Parkville, Victoria 3010, Australia