Structure and development of iridescent butterfly scales: Lattices and laminae

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Abstract
Iridescent butterfly scales are structurally colored, relying upon the interaction of light with detailed architecture to produce their color. In some iridescent scales, the reflective elements are contained within the body of the scale and come in two basic forms, lattices that produce diffraction colors (analogous to those produced by opal), and stacks of laminae that produce thin-film interference colors (analogous to those produced by soap or oil films). Both structures are remarkably complex and precise, yet each is only part of the total edifice built by the cell that makes the scale. To understand better how a cell can produce lattices or thin-film laminae, I studied the development of iridescent scales from two lycaenid butterflies. The presence of diffraction and thin-film scales in the same family (and in some cases on the same individual) suggests that the two types must be developmentally related; yet these results yield no clear explanation as to how. The diffraction lattice appears to be shaped within the boundaries of the scale cell by means of a convoluted series of membranes in which the smooth endoplasmic reticulum plays an important part. The thin-film interference laminae appear to result from the condensation of a network of filaments and tubes secreted outside the boundaries of the cell. This paper outlines the developmental histories of both types of scale and discusses the developmental implications of the mechanisms by which they form.