Assessment

A book on developmental plasticity needs a chapter on assessment, if only to show that adaptive environmental assessment occurs. Skepticism regarding the ability of nonhuman organisms to assess conditions well enough to make adaptive decisions has a long history in evolutionary biology, and it has been an important barrier to understanding the evolution of adaptive developmental plasticity. It is worth briefly reviewing this history in order to understand certain preconceptions about assessment that still persist. In the nineteenth century, critics of Darwin’s theory of sexual selection (Darwin, 1871) balked at the idea of an “aesthetic sense” in lowly creatures that would enable female choice of mates (representative papers are reprinted and discussed in Bajema, 1984). Later, the barrier persisted for other reasons. Even though naturalists routinely used the condition-appropriate expression of phenotypic traits to support adaptation hypotheses—a practice that assumes adaptive assessment of conditions as it is defined here—theoretically inclined biologists paid little attention to the question of facultatively expressed traits. Part of the difficulty lay in the problem of explaining how adaptive assessment could evolve within the framework of conventional genetics. Theodosius Dobzhansky, one of the twentieth century’s leading evolutionary biologists, acknowledged this unresolved problem in remarks following a lecture by J. S. Kennedy on the phase polyphenisms of migratory locusts (Kennedy, 1961). Dobzhansky referred to the “challenge to a geneticist” of explaining the adaptive switch between the sedentary and the migratory phenotypes of the locusts, which had been shown to be largely independent of genotype. He suggested that an extrachromosomal factor may be involved, a symbiotic microorganism that acts as a “plasmagene” whose multiplication would eventually stimulate phase change. Although Dobzhansky’s proposal was no more preposterous than some of the regulatory devices that have actually been discovered, Kennedy (1961) minced no words in his reply to this suggestion: . . . [W]e need not feel obliged to invoke a second organism to explain [phase polymorphism] unless we are reluctant to concede an important part to the environment as well as to heredity in moulding development. . . .