Publications [#229537] of H. Frederik Nijhout
Papers Published
- Nijhout, HF, Symmetry systems and compartments in Lepidopteran wings: The evolution of a patterning mechanism,
Development, vol. 120 no. SUPPL.
(January, 1994),
pp. 225-233
(last updated on 2024/04/23)Abstract:
The wing patterns of butterflies are made up of an array of discrete pattern elements. Wing patterns evolve through changes in the size, shape and color of these pattern elements. The pattern elements are arranged in several parallel symmetry systems that develop independently from one another. The wing is further compartmentalized for color pattern formation by the wing veins. Pattern development in these compartments is largely independent from that in adjacent compartments. This two-fold compartmentalization of the color pattern (by symmetry systems and wing veins) has resulted in an extremely flexible developmental system that allows each pattern element to vary and evolve independently, without the burden of correlated evolution in other elements. The lack of developmental constraints on pattern evolution may explain why butterflies have diverged so dramatically in their color patterns, and why accurate mimicry has evolved so frequently. This flexible developmental system appears to have evolved from the convergence of two ancient patterning systems that the butterflies inherited from their ancestors. Mapping of various pattern types onto a phylogeny of the Lepidoptera indicates that symmetry systems evolved in several steps from simple spotting patterns. Initially all such patterns were developmentally identical but each became individuated in the immediate ancestors of the butterflies. Compartmentalization by wing veins is found in all Lepidoptera and their sister group the Trichoptera, but affects primarily the ripple patterns that form the background upon which spotting patterns and symmetry systems develop. These background pattern are determined earlier in ontogeny than are the symmetry systems, and the compartmentalization mechanism is presumably no longer active when the latter develop. It appears that both individuation of symmetry systems and compartmentalization by the wing veins began at or near the wing margin. Only the butterflies and their immediate ancestors evolved a pattern formation mechanism that combines the development of a regular array of well-differentiated symmetry systems with the mechanism that compartmentalizes the wing with respect to color pattern formation. The result was an uncoupling of symmetry system development in each wing cell. This, together with the individuation of symmetry systems, yielded an essentially mosaic developmental system of unprecedented permutational flexibility that enabled the great radiation of butterfly wing patterns.