Adaptability of experimental populations of Drosophila melanogaster in directed and stabilizing selection]

Stabilizing selection for a set of morphometric wing traits was combined with directional selection for increased expression of radius incompletus (ri) mutation of Drosophila melanogaster. Three experimental regimens were used: directional and stabilizing selection (stabilized lines); directional selection (unstabilized lines); no selection (control lines). The dynamics of two fitness-associated parameters (coefficient of fluctuating asymmetry and population size) was registered during the experiment. At the end of experiment, selection competitive ability test was carried out. The competitive indices did not significantly differ under various treatments, while fluctuating asymmetry was significantly lower in stabilized than in unstabilized lines. Average population size was greater in stabilized lines as compared to the unstabilized ones. The possible causes for these differences are discussed.     

Interaction of directional and stabilizing selection for wing characteristics in Drosophila melanogaster

The effects of interaction of artificial directional and stabilizing selection were studied using a recessive mutation of Drosophila melanogaster radius incompletus which causes interruption of the second longitudinal wing vein (L2). The character under directional selection was the length of the proximal segment of L2; stabilizing selection was conducted for a complex of morphometric wing characters. Three experimental designs in three replications were used: 1) directional and stabilizing selection; 2) directional selection; 3) unselected controls. The dynamics and variation of the character under directional selection was shown to be different in the stabilized and unstabilized lines. Coefficient of variation which increased in the unstabilized lines remained at the same level in the stabilized lines and in the controls. The results are in agreement with those of the previous experiment, where directional selection was conducted for a behavioral trait, which suggests general character of the relationships observed.     

The effect of directed and stabilizing selection on the population size of Drosophila melanogaster

Population size as a measure of population fitness was assayed in laboratory lines of Drosophila melanogaster subjected to directional selection for positive photoactivity and stabilizing selection for a complex of metric wing characters. Three experimental designs were used: directional and stabilizing selection (DS); directional selection (D); unselected controls (C). Population size increased under all three experimental designs. The population size of DS and C lines as well as the rate of increase of the DS lines were higher as compared to the D lines. The results suggest that stabilizing selection maintains population fitness at the higher level.     

Evolutionary genetics of Drosophila ananassae. I. Effect of selection on body size and inversion frequencies

Adaptive importance of inversion polymorphism has been discussed in Drosophila species at several levels but no study has been carried out demonstrating the individual and combined effects of polymorphic inversions on the fitness of flies through bi‐directional selection. Therefore, artificial bi‐directional selection for thorax length in Drosophila ananassae was carried out for 10 generations. Both, Tukey test for selection difference and regression coefficients of offspring on mid‐parent are highly significant. The realized heritability (h²) in males of both high and low selection lines is more or less similar but is more pronounced in low line females, which suggests the asymmetrical response. This asymmetry in selection is discussed in the light of evidence provided by the study of chromosome inversion frequencies in different selection lines at different generations of selection. Interestingly, chromosome inversion frequency changes towards homozygosity for different gene arrangements in different selection lines. Tests of correlations at G₆ and G₁₀ among different gene arrangements as well as with mean thorax length suggest that 2L‐ST gene arrangement is negatively correlated, while 3L‐ST gene arrangement is positively correlated with thorax length. Furthermore, the present study shows the significant effects of 3L‐ST and 2L + 3L (positive correlation) on thorax length, while 3R‐ST and 2L + 3R show significant effect (negative correlation) on thorax length, which was not evident in the previous study. Present results also suggest how polymorphic inversions and their combinations affect the body size differently in different selection lines. These results suggest that thorax length in D. ananassae is under polygenic control and inversion polymorphism plays crucial role in maintaining body size by modifying genotypic frequency under various selection pressures.     

Molecular changes in the maize composite EPS12 during selection for resistance to pink stem borer

The pink stem borer (Sesamia nonagrioides Lefèvbre) is the most important pest of maize (Zea mays L.) throughout the Mediterranean area. The maize composite EPS12 has been chosen as the base population for a breeding program based on its resistance to pink stem borer, with the main selection criterion being resistance to stem tunneling. Yield was taken as a secondary selection criterion to avoid any unwanted negatively correlated response on this character. The aims of investigation were: (1) to monitor the effects of selection for resistance to pink stem borer on allele frequency at 70 simple sequence repeat (SSR) markers and their impact on the genetic structure of EPS12 and (2) to identify loci at which allelic frequencies changed significantly due to directional selection. Genetic diversity was reduced during the selection process (as expected since random genetic drift as well as selection could reduce genetic variability), but not significantly so. Although the loss of genetic variation was generally consistent with that expected in a model in which random genetic drift acts alone on neutral alleles, the changes observed in the frequency of five alleles were significantly greater than expected. Further, the linear trend of the departure from the random genetic drift model was significant for some allelic versions of two SSR markers, umc1329 and phi076; directional selection was therefore acting on these loci. The significant effect of directional selection on those markers suggests the presence of quantitative trait loci (QTLs) for tunnel length and/or for yield under artificial infestation with Sesamia nonagrioides on the long arm of chromosome 4.     

Apparent directional selection by biased pleiotropic mutation

Pleiotropic effects of deleterious mutations are considered to be among the factors responsible for genetic constraints on evolution by long-term directional selection acting on a quantitative trait. If pleiotropic phenotypic effects are biased in a particular direction, mutations generate apparent directional selection, which refers to the covariance between fitness and the trait owing to a linear association between the number of mutations possessed by individuals and the genotypic values of the trait. The present analysis has shown how the equilibrium mean value of the trait is determined by a balance between directional selection and biased pleiotropic mutations. Assuming that genes act additively both on the trait and on fitness, the total variance-standardized directional selection gradient was decomposed into apparent and true components. Experimental data on mutation bias from the bristle traits of Drosophila and life history traits of Daphnia suggest that apparent selection explains a small but significant fraction of directional selection pressure that is observed in nature; the data suggest that changes induced in a trait by biased pleiotropic mutation (i.e., by apparent directional selection) are easily compensated for by (true) directional selection.     

Phenotypic selection on flowering phenology and pollination efficiency traits between Primula populations with different pollinator assemblages

Floral traits have largely been attributed to phenotypic selection in plant–pollinator interactions. However, the strength of this link has rarely been ascertained with real pollinators. We conducted pollinator observations and estimated selection through female fitness on flowering phenology and floral traits between two Primula secundiflora populations. We quantified pollinator‐mediated selection by subtracting estimates of selection gradients of plants receiving supplemental hand pollination from those of plants receiving open pollination. There was net directional selection for an earlier flowering start date at populations where the dominant pollinators were syrphid flies, and flowering phenology was also subjected to stabilized quadratic selection. However, a later flowering start date was significantly selected at populations where the dominant pollinators were legitimate (normal pollination through the corolla tube entrance) and illegitimate bumblebees (abnormal pollination through nectar robbing hole which located at the corolla tube), and flowering phenology was subjected to disruptive quadratic selection. Wider corolla tube entrance diameter was selected at both populations. Furthermore, the strength of net directional selection on flowering start date and corolla tube entrance diameter was stronger at the population where the dominant pollinators were syrphid flies. Pollinator‐mediated selection explained most of the between‐population variations in the net directional selection on flowering phenology and corolla tube entrance diameter. Our results suggested the important influence of pollinator‐mediated selection on floral evolution. Variations in pollinator assemblages not only resulted in variation in the direction of selection but also the strength of selection on floral traits.     

Genetic heterogeneity and accelerated responses to directional selection in Drosophila

Summary Strains set up from single inseminated founder females of D. melanogaster from the same population have been previously shown to differ genetically for the incidence of flies with more than four scutellar chaetae (additional chaetae). Therefore the base population is polymorphic for this trait. This was exploited by carrying out directional selection in lines derived from those strains initially having a high mean chaeta number. This led to far more rapid responses to selection than were obtained in lines derived from strains with lower means, such that in one line a continuous accelerated response was observed for the first 12 generations of selection. A correlated response was found for sternopleural chaeta number at generation 19 of selection, showing that at least some genes may affect both traits.     

Action of natural selection in lines of Drosophila selected for a new level of canalization

Summary Four lines of Drosophila melanogaster previously selected for a stabilized phenotype of two extra dorsocentral bristles were examined for 20 generations of canalizing selection and relaxation of selection. A substantial frequency of flies with either two anterior or two posterior extra bristles was maintained in the relaxed lines. These patterns were the only ones tolerated by natural selection, i.e., the only symmetric ones. It was concluded that anterior and posterior dorsocentral bristles are two independent development structures, and the results are discussed in relation to two proposed genetic systems for bristle determination.     

Effect of directional selection for body size on fluctuating asymmetry in certain morphological traits in <Emphasis Type="Italic">Drosophila ananassae</Emphasis>

Variation in the subtle differences between the right and left sides of bilateral characters or fluctuating asymmetry (FA) has been considered as an indicator of an organism’s ability to cope with genetic and environmental stresses during development. However, due to inconsistency in the results of empirical studies, the relationship between FA and stress has been the subject of intense debate. In this study, we investigated whether stress caused by artificial bidirectional selection for body size has any effect on the levels of FA of different morphological traits in Drosophila ananassae. The realised heritability (h²) was higher in low-line females and high-line males, which suggests an asymmetrical response to selection for body size. Further, the levels of FA were compared across 10 generations of selection in different selection lines in both sexes for sternopleural bristle number, wing length, wing-to-thorax ratio, sex comb-tooth number and ovariole number. The levels of FA differed significantly among generations and selection lines but did not change markedly with directional selection. However, the levels of FA were higher in the G10 generation (at the end of selection) than G0 (at the start of selection) but lower than the G5 generation in different selection lines, suggesting that the levels of FA are not affected by the inbreeding generated during the course of selection. Also, the levels of FA in the hybrids of high and low lines were significantly lower than the parental selection lines, suggesting that FA is influenced by hybridisation. These results are discussed in the framework of the literature available on FA and its relationship with stress.     

Effect of cryoprotectant and genetic selection for body fat content on embryonic cryosurvival in mice

Lines of mice selected for high (HF) or low (LF) 12-week epididymal fat pad weight as a percentage of body weight were used to investigate the effects of genotype, two cryoprotectants [glycerol (GLY) and propylene glycol (PG)] and genotype x cryoprotectant interaction on cryosurvival of four and eight-cell embryos. Embryos were collected from selection lines and reciprocal crosses of selection lines (HFLF and LFHF) and frozen by established slow-cool methods. Embryos were thawed for 40s at room temperature and then placed in a 37° C waterbath for 1 min. Cryoprotectant was diluted from embryos with either 0.5 M sucrose (GLY-treated) or 1.0 M sucrose (PG-treated). Post-thaw survival was measured as the percentage of embryos developing to 36 h (PTS36), 48 h (PTS48) and hatched blastocyst (PTSHB), respectively. Non-frozen controls were cultured concurrently with frozen embryos. No significant genotype or genotype x cryoprotectant interaction effects were found. Results of the embryo freezing study indicated that selection for high or low fat content did not affect the ability of embryos to survive cryopreservation. There was no indication of embryo heterosis for post-thaw survial. Embryos frozen with GLY survived the freeze-thaw stress significantly better than those frozen in PG (P < 0.05). In vitro development of non-frozen controls at 36 and 48 h did not vary significantly among lines, but in vitro development was significantly different among lines at the hatched blastocyst stage (P < 0.05). Linear contrasts showed that the embryonic genome was responsible for differential in vitro development at the hatched blastocyst stage between these selected lines (HF > LF; P < 0.05); asymmetric response also occurred in that both HF and LF exceeded the unselected control line (P < 0.05).The research reported in this publication was funded by the North Carolina Agricultural Research Service (NCARS), Raleigh, NC 27695-7643. Use of trade names in this publication does not imply endorsement by the NCARS of the products named, nor criticism of similar ones not mentioned Present address: Department of Animal Science, University of Tennessee, Knoxville, TN 37901, USA     

Viability selection on prey morphology by a generalist predator

Prey use their locomotory capacity to escape predators, and there should thus be strong viability selection on locomotory morphology of prey. We compared feather morphology of wood pigeons Columba palumbus killed by goshawks Accipiter gentilis with that of survivors to quantify directional and quadratic selection on primary and rectrix feathers. The goshawk is mainly a predator attacking by surprise, leaving wood pigeons with an ability to accelerate fast at a selective advantage. There was directional selection for light primary feathers with a narrow calamus. In addition, there was directional selection for increased area of rectrices. These patterns of natural selection were confirmed in multivariate analyses of selection that showed selection for light primary feathers with a large area and narrow calamus and for a large area of rectrix feathers. These results provide evidence of selection on different aspects of feather morphology directly related to flight performance and thus escape ability from predators.     

Contrasted patterns of selection since maize domestication on duplicated genes encoding a starch pathway enzyme

Maize domestication from teosinte (Zea mays ssp. parviglumis) was accompanied by an increase of kernel size in landraces. Subsequent breeding has led to a diversification of kernel size and starch content among major groups of inbred lines. We aim at investigating the effect of domestication on duplicated genes encoding a key enzyme of the starch pathway, the ADP-glucose pyrophosphorylase (AGPase). Three pairs of paralogs encode the AGPase small (SSU) and large (LSU) subunits mainly expressed in the endosperm, the embryo and the leaf. We first validated the putative sequence of LSU_leaf through a comparative expression assay of the six genes. Second, we investigated the patterns of molecular evolution on a 2 kb coding region homologous among the six genes in three panels: teosintes, landraces, and inbred lines. We corrected for demographic effects by relying on empirical distributions built from 580 previously sequenced ESTs. We found contrasted patterns of selection among duplicates: three genes exhibit patterns of directional selection during domestication (SSU_end, LSU_emb) or breeding (LSU_leaf), two exhibit patterns consistent with diversifying (SSU_leaf) and balancing selection (SSU_emb) accompanying maize breeding. While patterns of linkage disequilibrium did not reveal sign of coevolution between genes expressed in the same organ, we detected an excess of non-synonymous substitutions in the small subunit functional domains highlighting their role in AGPase evolution. Our results offer a different picture on AGPase evolution than the one depicted at the Angiosperm level and reveal how genetic redundancy can provide flexibility in the response to selection.     

Non-linear selection response in Drosophila: a strategy for testing the rare-alleles model of quantitative genetic variability

Quantitative genetic theory predicts that variation due to rare alleles at many loci will generate a transient acceleration in the response to directional selection. We have tested this prediction by constructing experimental lines ofDrosophila melanogaster that carry positively selected ethanol resistance alleles at low frequencies, and then subjecting the lines to directional selection for ethanol resistance. Approximately 468,000 files were subjected to artificial selection over 30 generations. The predicted non-linear selection responses were observed in all experimental lines and replicates, on three genetic backgrounds. In contrast, un-selected controls and lines carrying random alleles at low frequencies on the same genetic backgrounds exhibited linear selection responses. These results demonstrate that non-linearities due to rare alleles are detectable and repeatable, provided that experiments are done on a sufficiently large scale. The results suggest that it may be possible to test for rare-alleles as a component of naturally occurring genetic variation by careful examination of selection response curves.     

Wolbachia infection and incompatibility dynamics in experimental selection lines

High and low levels of Wolbachia-induced cytoplasmic incompatibility (CI) were selected for in the parasitic wasp Nasonia vitripennis, in the single-infected strain Ti277. After nine generations of selection, males from lines selected for high incompatibility level (HI lines) were significantly more incompatible with uninfected females (AsymC) than the maternal strain. The reverse response, a full compatibility with AsymC, was observed in eight out of 12 lines selected for low incompatibility (LO lines), correlated with loss of Wolbachia infection. Bacterial density estimates in the eggs of some HI lines increased significantly. The procedure for line maintenance resulted in introgression of AsymC nuclear genome into the Ti277 background. Significant changes of CI level and bacterial density due to the introgression were also observed in the control lines, possibly reflecting an effect of host genotype on bacterial density and CI. After selection had been relaxed for six generations, bacterial density in the five high-infected HI lines declined back to a level comparable to the other lines. The data are consistent with the ‘bacterial dosage’ model, but with an upper threshold of bacterial infection above which there is no correlation between infection level and CI level. We further investigate the maternal transmission of bacterial density by a mother–daughter regression on bacterial density. The pattern observed is consistent with a density dependent regulation of bacterial numbers around an ‘equilibrium’ density, independent of any effects of CI. The equilibrium value is likely to be determined by both bacterial strain and host genotype.     

Relative efficiency of selection methods to improve a ratio of two traits in Tribolium

Summary Three generations of upward selection for the egg mass/adult weight ratio were carried out in Tribolium castaneum. The experiment involved five lines: E — selected for increased egg mass; W — selected for decreased adult weight; R — selected for egg mass/adult weight directly; L — linear index selected with economic weights of m ₂: -m ₁ egg mass to adult weight (m ₁ and m ₂ are the means for adult weight and egg mass, respectively); NL — nonlinear index selected. Adult weight (7 days after adult emergence) and egg mass (between 7 and 11 days) were measured. The NL, E, and L lines had the greatest observed responses for the ratio; the R and W lines were not effective in improving the egg mass/adult weight ratio. It was expected that the NL line would be superior to the E, L, and R lines, and that the W line would respond the least. Observed response was significant for egg mass in the NL line, and for adult weight in the E, W, and R lines. Strong selection to increase egg mass seems to represent the optimal criterion for the ratio to be improved. The usefulness of nonlinear indices as selection criteria to improve a nonlinear trait, previously found to be optimal for a trait defined as the product of two component traits, appears to hold also for the selective improvement of the ratio of two traits. Serious limitations expected for direct selection of the ratio have been confirmed in this experiment.     

Evidence for complex selection on four‐fold degenerate sites in Drosophila melanogaster

We considered genome‐wide four‐fold degenerate sites from an African Drosophila melanogaster population and compared them to short introns. To include divergence and to polarize the data, we used its close relatives Drosophila simulans, Drosophila sechellia, Drosophila erecta and Drosophila yakuba as outgroups. In D. melanogaster, the GC content at four‐fold degenerate sites is higher than in short introns; compared to its relatives, more AT than GC is fixed. The former has been explained by codon usage bias (CUB) favouring GC; the latter by decreased intensity of directional selection or by increased mutation bias towards AT. With a biallelic equilibrium model, evidence for directional selection comes mostly from the GC‐rich ancestral base composition. Together with a slight mutation bias, it leads to an asymmetry of the unpolarized allele frequency spectrum, from which directional selection is inferred. Using a quasi‐equilibrium model and polarized spectra, however, only purifying and no directional selection is detected. Furthermore, polarized spectra are proportional to those of the presumably unselected short introns. As we have no evidence for a decrease in effective population size, relaxed CUB must be due to a reduction in the selection coefficient. Going beyond the biallelic model and considering all four bases, signs of directional selection are stronger. In contrast to short introns, complementary bases show strand specificity and allele frequency spectra depend on mutation directions. Hence, the traditional biallelic model to describe the evolution of four‐fold degenerate sites should be replaced by more complex models assuming only quasi‐equilibrium and accounting for all four bases.     

Phenotypic selection and response to selection in Lobularia maritima: importance of direct and correlational components of natural selection

By using selection differentials, gradients and structural equation modelling (SEM), I have quantified the phenotypic selection acting on Lobularia maritima (Cruciferae) flower size, display, colour and density, using data on lifetime female fitness. Furthermore, by analysing the resulting F₁ generation in field and greenhouse conditions, I estimated the actual intergenerational change in the value of these traits. Both pollinators preferred plants with many and large flowers. Strong directional selection for increased flower display was found in all years of the study, regardless of the technique used. Indirect selection due to a high significant correlation with flower display occurred on flower colour and size. SEM showed that pollinators played only a minor role in this observed phenotypic selection. The analysis of the phenotypes of F₁ plants showed that flower display actually increased across generations. In addition, white flowers were significantly more frequent in the offspring population than in the parental one, mostly due to the association between flower display and white coloured flowers. This suggests that both direct and indirect selection can play a role in the evolution of correlated traits in this crucifer.     

Genetically idiosyncratic responses of Drosophila melanogaster populations to selection for improved learning ability

To what extent is adaptive evolution over short timescales repeatable? To address this question, we studied the performance of crosses between replicate Drosophila melanogaster lines previously subject to selection for improved learning response in the context of oviposition substrate choice. Of the 10 pairwise F₁ crosses among the five selection lines, four performed in the original learning assay similarly to the parental lines, whereas the remaining six showed learning scores significantly below the average of the parental lines. In particular, four F₁ crosses (three involving the same line) showed no detectable learning, on a par with unselected control lines. This indicates that the response to selection in some lines involved allelic substitutions at different loci. Additional assays of crosses between two selection lines indicated that the loss of performance in hybrids generalized to another type of learning assay, and held for both short‐ and long‐term memory. Joint analysis of first‐ and second‐generation crosses between these two lines supported the hypothesis that the response to selection in these different lines was based on the spread of recessive alleles at different loci. These results show that the evolutionary trajectories of populations of the same origin subject to uniform selection may sometimes diverge over very short evolutionary timescales.     

Haplo-diploid gene expression and pollen selection for tolerance to acetochlor in maize

The objectives of this research were to determine if genes controlling the reaction to the herbicide acetochlor in maize (Zea mays L.) are active during both the haploid and the diploid phases of the life cycle and if pollen selection can be utilized for improving sporophytic resistance. Pollen of eight inbred lines, previously characterized through sporophytic analysis for the level of tolerance to acetochlor, showed a differential reaction to the herbicide forin vitro tube length; moreover, such pollen reactions proved to be significantly correlated (r =0.786^*,df=6) with those of the sporophytes producing the pollen. Pollen analysis of two inbred lines (i.e. Mo17, tolerant, and B79, susceptible) and their single cross showed that thein vitro pollen-tube length reaction of the hybrid was intermediate between those of two parents. An experiment on pollen selection was then performed by growing tassels of Mo17xB79 in the presence of the herbicide. Pollen obtained from treated tassels showed a greater tolerance to acetochlor, assessed asin vitro tube length reaction, than pollen obtained from control tassels. Moreover, the backcross [B79 (Mo17xB79)] sporophytic population obtained using pollen from the treated tassels was more tolerant (as indicated by the fresh weight of plants grown in the presence of the herbicide) than was the control backcross population. The two populations did not differ when grown without the herbicide. These findings indicate that genes controlling the reaction to acetochlor in maize have haplodiploid expression; consequently, pollen selection can be applied for improving plant tolerance.