Nijhout H F
J Embryol Exp Morphol. 1984 Jun;81:287-305.
When young pupae of Vanessa cardui, V. virginiensis and Precis coenia (Lepidoptera: Nymphalidae) are exposed to severe coldshock many develop aberrant adult wing patterns. For each species, a synchronous cohort of experimental animals always develops a broad range of aberrant pattern morphologies but these can always be arranged in a single unbranched morphological series. When such phenotypic series are compared, between species and between wing surfaces within a species (each wing surface usually bears a different colour pattern), many parallel modifications and trends become evident. These parallelisms reveal certain homologies of pattern elements and suggest that a common physiology underlies the development of a considerable diversity of normal and aberrant colour patterns. The case is made that the phenotypic series produced may represent either a series of sequential stages in colour pattern determination or, a series of quantitatively different 'interpretations' of an established gradient system. Colour pattern morphoclines reveal which patterns are developmentally 'adjacent' to one another and may therefore prove useful in elucidating the evolution of patterns.
当荨麻蛱蝶、维吉尼亚蛱蝶和苎麻珍蝶（鳞翅目：蛱蝶科）的幼蛹遭受严重冷休克时，许多会发育出异常的成虫翅型。对于每个物种而言，一组同步的实验动物总会发育出广泛的异常形态，但这些形态总能排列成单一的无分支形态序列。当比较不同物种之间以及同一物种内不同翅面（每个翅面通常具有不同的颜色图案）的此类表型序列时，许多平行的变化和趋势就会变得明显。这些平行关系揭示了图案元素的某些同源性，并表明一种共同的生理机制是正常和异常颜色图案多样性发展的基础。有观点认为，所产生的表型序列可能代表颜色图案确定过程中的一系列连续阶段，或者是既定梯度系统的一系列数量上不同的“解释”。颜色图案形态渐变群揭示了哪些图案在发育上彼此“相邻”，因此可能有助于阐明图案的进化。
