The development of eyespot patterns on butterfly wings: morphogen sources or sinks?

ABSTRACT We have studied the development of the eyespot colour pattern on the adult dorsal forewing of the nymphalid butterflies, Bicyclus safitza and B. anynana, by cauterising the presumptive eyespot centres (the foci) on the pupal wing. The effects on pattern depended on age at cautery. Early focal cautery (at 1-12 hours after pupation) usually reduced or eliminated the eyespot, while cautery at a non-focal site usually had no effect. These results resemble those of a previous study on another species but, in addition, we find that a later cautery (at 12-24 hours) had the converse effect of generating pattern, so that focal cautery enlarged the anterior eyespot (but usually not the large posterior eyespot) and nonfocal cautery induced a new ectopic eyespot. The effects of cautery on patterning are more extensive by an order of magnitude than the cell death which is caused, so implicating a long-range mechanism, such as a morphogen gradient, in eyespot development. The focus clearly acts to establish the normal eyespot pattern, but a simple source/diffusion model is not supported by the response to late cautery. We suggest two alternative forms of gradient model in which late damage can mimic and augment the action of a focus. In the Source/Threshold model, the focus is a morphogen source, and cautery can remove the focus but also transiently lowers the response threshold in surrounding cells. In the Sink model, the focus generates the gradient by removing morphogen, and cautery can eliminate the focus but it also causes a transient destruction or leakage of morphogen. These models can explain most features of the results of cautery.