Structure of butterfly scales: patterning in an insect cuticle.

All butterfly and moth scales and bristles are made of non-living insect cuticle. Each is the product of a single epithelial cell, and all share the same basic architecture. However, some are highly specialized, and their cuticle is further elaborated into stacks of thin-films, lattices, or other minute structures, many of which first came to our attention because they interact with light to produce structural colors. The scale cell forms the scale by extruding a projection of itself and secreting around it the outer epicuticle, a thin cuticular envelope which will form the outer-most layer of the scale. The inner layers of cuticle, collectively called the procuticle, are secreted thereafter and go on to form the lattices, pillars, or other internal structures of the scale. We believe that the pattern-forming mechanisms used by the cell to shape the cuticle into its finished form include elastic buckling of the outer epicuticle to produce external folds, and "masking" of certain areas of the original epicuticular envelope to produce thin spots which will break through to become windows. Varied though they be, all insect cuticular patterns have common basic elements, which suggests that our findings may be generalized to other highly patterned insect cuticles, particularly those formed by single cells.