Population structure in artificial selection programmes: simulation studies.
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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) x 10(-3).
Selective sweeps of variation caused by fixation of major genes may have a dramatic impact on the genetic gain from background polygenic variation, particularly in the genome regions closely linked to the major gene. The response to selection can be restrained because of the reduced selection intensity and the reduced effective population size caused by the increase in frequency of the major gene. In the context of a selected population where fixation of a known major gene is desired, the question arises as to which is the optimal path of increase in frequency of the gene so that the selective sweep of variation resulting from its fixation is minimized. Using basic theoretical arguments we propose a frequency path that maximizes simultaneously the effective population size applicable to the selected background and the selection intensity on the polygenic variation by minimizing the average squared selection intensity on the major gene over generations up to a given fixation time. We also propose the use of mating between carriers and non-carriers of the major gene, in order to promote the effective recombination between the major gene and its linked polygenic background. Using a locus-based computer simulation assuming different degrees of linkage, we show that the path proposed is more effective than a similar path recently published, and that the combination of the selection and mating methods provides an efficient way to palliate the negative effects of a selective sweep.
Previous divergent selection for nest-building behavior at 22 +/- 1 degrees C resulted in a 40-fold difference between the high and the low lines in amount of cotton used to build a nest. Correlated responses to selection indicated positive genetic correlations with body weight, nest-building at 4 +/- 1 degrees C, and litter size and negative genetic correlations with food consumption. At generation 46, the replicate high-selected (High 1 x High 2), randomly bred control (Control 1 x Control 2), and low-selected (Low 1 x Low 2) lines were crossed and the F1 showed significant heterosis for nest-building behavior. Regression of the F3 on the F2 generation gave heritability estimates of 0.16 +/- 0.10 for the high and 0.07 +/- 0.10 for the low cross, revealing a potential to break the selection limit (at least in the high direction), which had been reached at about 20 generations of selection. Indeed, renewed selection resulted in responses in both the high and the low directions of nesting, yielding realized heritabilities of 0.29 +/- 0.02 and 0.30 +/- 0.004, respectively. Replicated renewed selection, using the F3 generation as the base population, in the high direction of nesting resulted in correlated increases in nest-building at 4 +/- 1 degrees C, litter size, and food consumption. Body weight did not change. The positive correlation with food consumption is opposite in sign compared to the original selection experiment. This indicates that the evolutionary potential of a population to adapt to a changing environment not only depends on its current genetic variability in one adaptive trait, but may be constrained by genetic correlations changing over the course of selection.
Replicated within-family selection for increased voluntary wheel running in outbred house mice (Mus domesticus; Hsd:ICR strain) was applied with four high-selected and four control lines (10 families/line). Mice were housed individually with access to activity wheels for a period of 6 days, and selection was based on the mean number of revolutions run on days 5 and 6. Prior to selection, heritabilities of mean revolutions run per day (rev/day), average running velocity (rpm), and number of minutes during which any activity occurred (min/day) were estimated by midparent-offspring regression. Heritabilities were 0.18, 0.28, and 0.14, respectively; the estimate for min/day did not differ significantly from zero. Ten generations of selection for increased rev/day resulted in an average 75% increase in activity in the four selected lines, as compared with control lines. Realized heritability averaged 0.19 (range, 0.12-0.24 for the high-activity lines), or 0.28 when adjusted for within-family selection. Rev/day increased mainly through changes in rpm rather than min/day. These lines will be studied for correlated responses in exercise physiology capacities and will be made available to other researchers on request.
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A simple method is presented for determining the effect on the genetic variance of a quantitative trait, of selection followed by various degrees of assortative mating.
Quantitative characters are often said to evolve rather slowly, taking many generations to exhibit appreciable differences among populations. We tested this notion experimentally by performing bi-directional selection on corolla diameter of plants from a wild population of Phlox drummondii for three generations. By monitoring flower size, tube length and stigma-anther proximity of flowers, we obtained the direct and indirect responses to selection, and calculated genetic correlations, realized and narrow sense heritabilities using offspring-mother regression. Realized heritability of flower size was high (0.83), whereas genetic correlations among traits were weak or not significant. The per-generation average of the response in corolla diameter was about 5%. We found that P. drummondii has a great capacity to respond rapidly to selection, and this capacity may be in part responsible for the observed high degree of differentiation within the species. We also concluded that rapid evolution of morphological floral traits is possible.
The tropical butterfly Bicyclus anynana shows adaptive phenotypic plasticity in response to wet-dry seasonality. The wet season form (WSF) has a conspicuous wing pattern with large eyespots, whereas the dry season form (DSF) lacks eyespots and therefore has a more cryptic appearance. Temperature is the main factor controlling this difference: rearing larvae at a low (<19 degrees C) temperature in the laboratory results in the DSF, whereas rearing at a high (>23 degrees C) temperature induces the WSF. We applied truncation selection in opposite directions in successive generations reared at two alternating temperatures (18.5 degrees C and 23.5 degrees C) to increase (for two High Plasticity (HP) lines), and decrease (for two Low Plasticity (LP) lines) wing pattern plasticity. Plasticity was assessed by partitioning full-sib families over four rearing temperatures (18.5 degrees C, 20.5 degrees C, 21.5 degrees C and 23.5 degrees C). Several wing pattern elements were measured for which the first principal component (PC1) provides a useful summary. The slopes of reaction norms for PC1 were significantly steeper in the HP lines than in the LP lines; however, the selection lines did not always differ significantly from the unselected stock. The results of crosses between the replicates of the selection lines gave no indication for effects of inbreeding. We argue that high, positive genetic correlations across temperatures retard a response to selection in opposite directions in different environments. This is discussed with respect to potential evolutionary constraints in natural populations in these butterflies.
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