The study of metamorphosis and morphogenesis in the Drosophila fruit fly has far-reaching consequences for our fundamental understanding of these processes and wide-spread applications in biomedicine. Understanding the influence of mutations in the planar cell polarity pathway is one important aspect in this respect, and will potentially help in the treatment of diseases. The Drosophila eye consists of small units called ommatidia, shaped as regular hexagons under the balance of cell-cell adhesion and interfacial tension. Such regular hexagonal tessellations are known to obey certain topological and geometrical relations. In particular, the theory of Voronoi tilings shows that only a small magnitude of area variation is allowed before defects occur. Yet, the experiment image shows area variations far greater than this limit. A theory is proposed separating the variation into two independent parts, i.e., systematical and statistical variations. The statistical variation describes the local magnitude of disorder and should follow the variation restriction. The systematical variation describes area changes over large scales. The total variation observed in experiment is the combination of the two variations. This hypothesis is verified both in simulations and in analyzing experimental images of wild-type and mutant Drosophila retinas.
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Systematic and statistical variations of ommatidial area in the Drosophila eye and their effect on eye morphology