Surprisingly little is known about the genetic architecture of body size in natural populations of Drosophila melanogaster. Using both generation means and triple-test-cross analyses, we investigated the genetic architecture of wing size (an indicator of body size) and wing shape in a naturally occurring body size cline. For wing size, we found signi®cant epistatic genetic variance and evidence of past directional selection for increased body size. While wing shape also exhibits signi®cant epistatic genetic variance, there was no indication of directional selection, suggesting instead a history of optimizing selection. Our results support the idea that epistatic variance may be more common in natural populations than was once suspected. Also, our results suggest substantial directional selection on wing size but not shape.Keywords: directional selection, Drosophila melanogaster, epistasis, genetic architecture, wing shape, wing size.
IntroductionBody size is of central importance in evolution and ecology, and it has been study extensively in both arti®cial and natural environments. Numerous allometric relationships between life history, physiological and behavioural traits and body size have been reported across species (e.g. Schmidt-Nielsen, 1984). Various trade-os aecting body size have been identi®ed (Stearns, 1992). Although arti®cial selection experiments provide numerous insights into aspects of the evolution of body size, the generality of the ®ndings is necessarily limited. Attempts to understand the how and why of evolution must eventually involve studies of natural populations.In Drosophila, in particular, surprisingly little is known about the quantitative genetics of body size of natural populations, apart from its high heritability (Coyne & Beecham, 1987; Prout & Barker, 1989; Ruiz et al., 1991; Thomas & Barker, 1993). Our intention in this study was to answer some additional and important questions regarding the evolution of body size in Drosophila melanogaster in a natural body-size cline. First, using wing area as a measure of body size, we questioned the importance of epistasis in population divergence. The answer is relevant not only to the shifting balance theory of evolution, but also to questions regarding the evolution of mating systems and conservation genetics (see Whitlock et al., 1995; Fenster et al., 1997). Secondly, what sort of selection (either directional or optimizing) is likely to be the predominant form of natural selection acting on the trait? Body size in D. melanogaster has an intermediate optimum value, largely determined by correlations with fecundity, development time and larval survival (Ro, 1981). Thus, although appearing to be under optimizing selection, the optimizing selection on body size is only apparent (Falconer, 1989): it is not clear what type of selection acts directly on genes determining body size within the constraints imposed by correlated characters.In addition to wing area, we also analysed wing shape. Although a number of investigators have documented ...