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Word Count: 248 Text Word Count: 3987 Number of Figures: 6 D ise as e M od el s & M ec ha ni sm s D M M Ac ce pt ed m an us cr ip t © 2013. Published by The Company of Biologists Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial Share Alike License (http://creativecommons.org/licenses/by-nc-sa/3.0), which permits unrestricted non-commercial use, distribution and reproduction in any medium provided that the original work is properly cited and all further distributions of the work or adaptation are subject to the same Creative Commons License terms. http://dmm.biologists.org/lookup/doi/10.1242/dmm.012591 Access the most recent version at DMM Advance Online Articles. Posted 4 July 2013 as doi: 10.1242/dmm.012591