I factors are transposable elements of Drosophila melanogaster similar to mammalian LINEs, that transpose by reverse transcription of an RNA intermediate and are responsible for the I–R system of hybrid dysgenesis. There are two categories of strains in this species: inducer, that contain about 15 I elements at the various sites on chromosomal arms, and reactive, that lack active I factors. I elements occur in various Drosophila species. Potentially functional I factors from Drosophila teissieri can transpose when introduced by P-element-mediated transformation in a reactive strain of Drosophila melanogaster. We have studied the properties of Drosophila melanogaster strains into which such an I factor from Drosophila teissieri, named Itei, was introduced. Typical hybrid dysgenesis is produced when males carrying Itei are crossed with reactive females. However, more than one copy of the element seems necessary to produce dysgenic traits, whereas only one I factor of Drosophila melanogaster seems to be sufficient. The copy number of Itei in transformed lines maintained by endogamous crosses increases rapidly and stabilizes at values similar to those observed in inducer strains. As Drosophila teissieri contains much fewer copies than the Drosophila melanogaster strains, this suggests that the copy number of I elements is not simply regulated by sequences present in the element itself.

An aberrant chromosome 1 carrying an inverted fragment with two amplified DNA regions was isolated from wild populations of Mus musculus. Meiotic drive favouring the aberrant chromosome was demonstrated for heterozygous females. Its cause was preferential passage of aberrant chromosome 1 to the oocyte. Genetic analysis allowed us to identify a two-component system conditioning deviation from equal segregation of the homologues. The system consists of a postulated distorter and responder. The distorter is located on chromosome 1 distally to the responder, between the ln and Pep-3 genes, and it acts on the responder when in trans position. Polymorphism of the distorters was manifested as variation in their effect on meiotic drive level in the laboratory strain and mice from wild populations.

An aberrant chromosome 1 with two large homogeneously staining insertions was isolated from wild populations of Mus musculus musculus. The specific features of the aberrant chromosome have been described elsewhere (Agulnik et al. 1990). These include its preferential entry into the oocyte of heterozygous females, increased mortality of homozygotes and decreased fertility of homozygous females. Here we present data indicating that chromatid segregation in heterozygous females depends upon which sperm enters the oocyte before the second meiotic division: meioticdrive is powerful when it is sperm bearing normal chromosome 1, and segregation normalizesduring Mil when it is sperm bearing chromosome 1 with the extra segment. Experimental data are adduced and explanations offered for the observed phenomenon.

An understanding of the physiological and genetic changes which determine the response to selection is critical for both evolutionary theory and to assess the application of new molecular techniques to commercial animal breeding. We investigated an aspect of physiology, growth hormone (GH) metabolism, which might a priori have been expected to play a large part in the response of mouse lines selected for high or low body weight. Disruption of endogenous GH or addition of exogenous GH had similar proportionate effects on body weight in both lines of mice (although differences in body composition arose) suggesting that neither the production of GH nor receptor sensitivity to GH had been altered as a result of selection. This supports a ‘pleiotropic model’ of the response to selection: that many genes with diverse metabolic roles all contribute to the divergent phenotype. This result has significant commercial implications as it suggests that artificial selection, transgenic technology and environmental manipulation may be synergistic rather than antagonistic strategies.

Divergent selection for abdominal bristle number was carried out for 47 generations, starting from a completely homozygous population of Drosophila melanogaster. All lines were selected with the same proportion (20%) but at two different numbers of selected parents of each sex (5 or 25). A significant response to selection was obtained in 25 lines (out of 40). In most cases, it could be wholly attributed to a single mutation of relatively large effect (> 0·3 phenotypic standard deviations). A total number of 30 mutations were detected. In agreement with theory, larger responses in each direction were achieved by those lines selected at greater effective population sizes. A large fraction of mutations were lethals (10/30). Thus, the observed divergence between lines of the same effective size selected in opposite directions was smaller than expected under neutrality. The ratio of new mutational variance to environmental variance was estimated to be(0·52±0·09)×10−3.

Starting from a completely homozygous population of Drosophila melanogaster, lines have been derived and subjected to 47 generations of divergent selection for abdominal bristle number (20 lines selected in each direction) or to 60–67 generations of inbreeding (100 B lines maintained by a single brother-sister mating, 100 C lines maintained by two double first cousin ma tings). In the selected lines, 25 were identified carrying at least 30 mutations affecting bristle number. A large fraction of these mutations (42 %) were lethals. Non-lethal mutations had smaller effects on the trait, were predominantly additive and had no detectable pleiotropic effects on fitness. In the inbred lines, 21 mutations affecting bristles were individually analysed. Deleterious mutations had the largest effects on the trait (irrespective of sign) and showed recessive gene action (complete or incomplete). The rest were predominantly additive and had smaller effects. Thus, both procedures identify a quasi-neutral class of additive mutations which should be close to that responsible for standing variation in natural populations. Moreover, the results indicate a leptokurtic distribution of mutant effects, consistent with a model of natural selection acting on bristles through pleiotropic effects of pertinent loci on fitness. Consequently, neutral additive alleles of considerable effect can be found segregating at intermediate frequencies in natural populations.

We investigate the establishment and spread of new adaptive peaks within Wright's ‘shifting balance’. The third phase of the ‘shifting balance’ involves a kind of group selection, since demes in which a superior peak has been established contain more individuals, and so send out more migrants. We assume that population size, N, increases with mean fitness, , according to the exponential relation, . Here, k is a measure of the weakness of density-dependent regulation, and equals the inverse of the regression of log (fitness) on log(N). In the island model, we find that just as with soft selection (k = 0), two distinct types of behaviour exist: group selection makes no qualitative difference. With low numbers of migrants, demes fluctuate almost independently, and only one equilibrium exists. With large numbers of migrants, all the demes evolve towards the same adaptive peak, and so the whole population can move towards one or other of the peaks. Group selection can be understood in terms of an effective mean fitness function. Its main consequence is to increase the effect of selection relative to drift (Ns), and so increase the bias towards the fitter peak. However, this increased bias depends on the ratio between k and the deme size (k/N), and so is very small when density-dependence is reasonably strong.