Asymmetrical response to selection for rate of development inDrosophila subobscura

Abstract

Starting from a geographically hybrid foundation population ofDrosophila subobscura, selection for fast and for slow development has been practised without inbreeding on a diet with an unusually high level of protein. Realized heritabilities in the fast and slow lines were + 0·063 ± 0·029 and + 0·186 ± 0·031 respectively. A half-sib analysis of the foundation population and full-sib analyses of the first two and the last two selected generations were carried out. Hybrids between the two lines were approximately intermediate between their parents.Two types of genetic explanation of the asymmetrical response are discussed. The first assumes directional dominance of alleles for fast development. Such an assumption can explain the asymmetrical response, but runs into difficulties in explaining the nature of the genetic variance in the selected populations and the intermediacy of the hybrids between the two lines.A second assumption, which appears to fit the facts better, is that there exists a ‘developmental barrier’ preventing development at a rate appreciably faster than that of the foundation population. In physiological terms this implies that more rapid development requires a more profound modification of the population than could be achieved by a few generations of selection. In genetic terms, it implies epistatic interactions between genes at different loci: gene substitutions at a given locus which increase development rate on a genetic background causing slow development have little or no effect on a genetic background causing rapid development. In other words, there is a law of diminishing returns as more and more alleles for fast development are accumulated in the genotype. It is suggested that genetic situations of this kind may be common in populations which have been exposed to directional selection for a long time in reasonably large populations, either in nature or in domestication.