Increased Selection Response in Larger Populations. II. Selection for Ethanol Vapor Resistance in Drosophila Melanogaster at Two Population Sizes

The effect of large population size on selection response was investigated using Drosophila melanogaster, with four "small" lines of 160 selected parents/generation compared to two "large" lines of 1,600 selected parents/generation. All lines were selected under similar conditions at a selection intensity of approximately 0.55 standard deviations, for 65 generations, for increased ethanol vapor resistance (measured in minutes required to become anesthetized). Two unselected control lines of 320 parents/generation were also maintained. A significant effect of population size was found. The final treatment means and standard errors were: 27.91 +/- 1.28 min (two "large" lines); 19.40 +/- 1.54 min (four "small" lines); and 4.98 +/- 0.35 min (two control lines). To estimate the mutation rate for the trait, two isogenic lines of about 400 selected parents were selected for 29 generations. The mean increase in additive genetic variance per generation was 0.0009 times the initial environmental variance of the outbred lines. This is comparable to other reported mutation rates. Mutation can explain part of the difference in evolved resistance between treatments, but it appears that even at rather large population sizes, a large difference in long-term response can be obtained in larger outbred lines, from more complete utilization of the initial genetic variation.     

The Response to Artificial Selection from New Mutations in Drosophila Melanogaster

Twenty generations of divergent selection for abdominal bristle number were carried out 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 detected in eight lines (out of 40) and, in most cases, it could be wholly attributed to a single mutation of relatively large effect (0.5-2 phenotypic standard deviations). The ratio of new mutational variance to environmental variance was estimated to be (0.33 +/- 0.11) X 10(-3). The distribution of mutant effects was asymmetrical, both with respect to bristle number (85% of it was negative) and to fitness (most detected bristle mutations were lethal or semilethal). Moreover, this distribution was leptokurtic, due to the presence of major genes. Gene action on bristles ranged from additive to completely recessive, no epistatic interactions being found. In agreement with theory, larger responses in each direction were achieved by those lines selected at greater effective population sizes. Furthermore, the observed divergence between lines selected in opposite directions was proportional to their effective size, as predicted for mutations of large effect.     

Effects of Population Size and Selection Intensity on Correlated Responses to Selection for Postweaning Gain in Mice

Correlated responses to selection for postweaning gain in mice were studied to determine the influence of population size and selection intensity. Correlated traits measured were three-, six- and eight-week body weights, litter size, twelve-day litter weight, proportion infertile matings and two indexes of reproductive performance. In general, the results agreed with observations made on direct response: correlated responses in the body weight traits and litter size increased as (1) selection intensity increased and (2) effective population size increased. Correlated responses in the body weight traits and litter size were positive in the large population size lines (16 pairs), as expected from the positive genetic correlation between these traits and postweaning gain. However, several negative correlated responses were observed at small population sizes (one and two pairs). Within each level of selection intensity, traits generally associated with fitness tended to decline most in the very small populations (one and two pairs) and in the large populations (16 pairs) for apparently different reasons. The fitness decline at the small effective population sizes was attributable to inbreeding depression. In contrast, it was postulated that the fitness decline at the large effective population size was due to selection moving the population mean for body weight and a trait positively correlated genetically with body weight (i.e., percent body fat) away from an optimum.     

Selection for nursing ability and adult weight in mice

Three selection treatments were conducted for 12 generations in each of two base populations (P and Q): (1) increased nursing ability of the mother (n12), as measured by mean 12-day weight of eight young within a crossfostering set (M(P) and M(Q) lines), (2) increased adult (42-day) body weight of the offspring (w42) (W(P) and W( Q) lines), and (3) performance combining the two traits (n12 and w42) into a selection index (B(P) and B(Q) lines). Lines C( P) and C(Q) were maintained as unselected controls in each population. In each line-generation subclass, 92 single-pair matings were made and the offspring assigned to balanced crossfostering sets of four dams each. Regression coefficients of mean performance (in grams) on generations were 0.080 +/-0.029 and 0.054 +/- 0.031 for n12 in M(P) and M(Q), and 0.680 +/- 0.039 and 0.868 +/- 0.051 for w42 in W(P) and W(Q), respectively. The B(P) and B(Q) lines showed genetic gains in n12 (0.090 and 0.053, respectively) and w42 (0.576 and 0.696) intermediate between the performance of M(P) and W(P), and M(Q) and W(Q), respectively, except for n12 of B(Q). Realized heritabilities for n12 were 0.16 +/- 0.05 and 0.11 +/- 0.06 and those for w42 were 0.40 +/- 0.02 and 0.43 +/- 0.03 for P and Q, respectively. The realized genetic correlations between n12 and w42 were 0.70 +/- 0.07 and 0.73 +/- 0.08 in P and Q, respectively. The ratios of the predicted to observed responses in M(P), B(P) and B(Q) were 0.99, 1.03 and 0.89, respectively. However, the predicted and observed responses differed in M( Q), W(P) and W(Q); the ratios were 1.29, 0.65 and 0.65, respectively. The observed combined responses for n12 and w42 in the index lines (B(P) and B(Q)) were smaller than the optimum expected from index selection. A possible cause was that the estimated genetic correlations (0.22 +/- 0.16 and -0.17 +/- 0.16 for B(P) and B( Q), respectively) and heritabilities (0.39 +/- 0.03 and 0.28 +/- 0.02, respectively) for w42 that were used to construct the selection index were smaller than the respective realized parameters.     

Genetic Divergence under Uniform Selection. II. Different Responses to Selection for Knockdown Resistance to Ethanol among DROSOPHILA MELANOGASTER Populations and Their Replicate Lines

We have tested the hypothesis that genetic differences among conspecific populations may result in diverse responses to selection, using natural populations of Drosophila melanogaster. Selection for ethanol tolerance in a tube measuring knockdown resistance was imposed on five West Coast populations. In 24 generations the selected lines increased their mean knockdown times, on average, by a factor of 2.7. An initially weak latitudinal cline was steepened by selection. The two southernmost populations showed the same increases in the selected character, but differed consistently in their correlated responses in characters related to ethanol tolerance. This result indicates that the populations responded to selection by different genetic changes. Selection decreased female body weight and increased resistance to acetone, suggesting components of the response unrelated to ethanol metabolism. The Adhs allele was favored by selection in all populations at the onset, but increased in frequency only in the selected lines of the southernmost population. There was a correlation between latitude and Adh frequency changes, suggesting that fitnesses of the Adh alleles were dependent on the genetic background. Genetic background also had a large effect on the loss of fitness due to selection. Genetic drift between replicate lines caused more variation in selection response than initial genetic differences between populations. This result demonstrates the importance of genetic drift in divergence among natural populations undergoing uniform selection, since the effective population sizes approached those of small natural populations. Drift caused greater divergence between selected replicates than control replicates. Implications of this result for the genetic model of selection response are discussed.     

Selection for starvation resistance in Drosophila melanogaster : physiological correlates,enzyme activities and multiple stress responses

Correlated responses to artificial selection for stress tolerance can provide insight into underlying genetic variation and the physiological basis of stress resistance . Lines of Drosophila melanogaster held in the absence of food or with an unsuitable resource, specifically decomposing lemon, responded to selection by becoming starvation resistant. The lemon-selected lines also adapted by evolving a resource-based induction response. Compared to control lines, the selected lines tended to store more lipid, develop slower and have a larger body size. Additional responses included resistance to desiccation and acetone fumes, suggesting multiple stress resistance is a correlated result of selection for starvation resistance. The specific metabolic rate was lower in the starvation selected lines and enzyme activities changed in response to selection. In particular, enzyme activities indirectly associated with lipid biogenesis increased in both types of selected lines. The correlated responses to the two selection regimes were sufficiently consistent to indicate a common basis for starvation resistance. Specific responses to starvation selection appeared to oppose the short-term phenotypic responses to starvation. Thus, a common response to stress selection may be to ameliorate the immediate physiological impact of the stress factor.     

Effects of Population Size and Selection Intensity on Short-Term Response to Selection for Postweaning Gain in Mice

The effects of population size and selection intensity on the mean response was examined after 14 generations of within full-sib family selection for postweaning gain in mice. Population sizes of 1, 2, 4, 8 and 16 pair matings were each evaluated at selection intensities of 100% (control), 50% and 25% in a replicated experiment. Selection response per generation increased as selection intensity increased. Selection response and realized heritability tended to increase with increasing population size. Replicate variability in realized heritability was large at population sizes of 1, 2 and 4 pairs. Genetic drift was implicated as the primary factor causing the reduced response and lowered repeatability at the smaller population sizes. Lines with intended effective population sizes of 62 yielded larger selection responses per unit selection differential than lines with effective population sizes of 30 or less.     

Selection for High Adult Body Weight in Drosophila Populations with Different Structures

Selection for high adult body weight in Drosophila melanogaster was practiced for 18 generations in three selection lines. These lines were genetically similar and of equal size but different in population structure. One line represented a large mass-selected, random-mating population, while the other two lines simulated large populations that had been subdivided into partial isolates or demes. Mass selection and random mating occurred within each deme. These two subdivided lines were different only in the rate of effective migration among the demes (5% and 10%). Selection intensities of approximately 20% were applied to these populations. A fourth line served as a random mating control. Heritability of adult body weight in the base population was estimated to be 0.58± 0.22. The results indicate that significantly greater responses were achieved in the subdivided lines than in the large mass-selected line, in spite of the fact that larger selection differentials were applied to the latter. No significant differences in response were observed between the two subdivided lines. Wright (1930, 1931) postulated that selection would be most efficient in subdivided populations with limited interdeme migration. The present findings appear to support this theory.     

Direct Response to Selection for Postweaning Gain in the Rat

The effectiveness of selection for 3–9-week gain was examined in a population of rats with a history of past selection for high 3–9-week gain. Lines were selected for high (U line) and low (D line) 3–9-week gain with two replicates of each line. Two randomly selected lines were also kept, one originating from the same base population as the two selected lines (R line) and the other originating from a population that had been randomly mated for the previous 27 generations (C line). Two replicates of each of these lines were kept. After seven generations of selection, a randomly selected line (relaxed line) was formed from each of the two upward- and each of the two downward-selected lines. Results have been presented for 13 generations of selection. The environmental trend for 3–9-week gain, as indicated by the randomly selected R and C lines, was consistently negative in all four lines. Realized heritabilities calculated by deviating the response to selection from the trend in the R or C lines resulted in non-significantly higher values in the D lines than the U lines. Six generations of relaxation of selection indicated no effect of natural selection in the U lines or the D lines. The relative magnitude of the drift, error and common environmental variances were estimated by the methods given by Hill (1971). The estimates of these parameters then led to calculation of the degree of bias in the sampling variances of the realized heritability estimates. As was predicted by Hill (1971), estimates of the variance of realized heritabilities obtained by using standard regression techniques were less than those obtained using Hill''s formulae. The results are discussed in relation to other similar studies with rats and mice.     

Evolutionary Endocrinology of Juvenile Hormone Esterase in Gryllus Assimilis: Direct and Correlated Responses to Selection

Hemolymph juvenile hormone esterase (JHE) activity on the third day of the last stadium in the cricket, Gryllus assimilis, exhibited a significant response to selection in each of six replicate lines. Mean realized heritability was 0.26 +/- 0.04. The response was due to changes in whole-organism enzyme activity as well as to changes in the proportion of enzyme allocated to the hemolymph compartment. In vivo juvenile hormone metabolism differed between some lines selected for high vs. low enzyme activity. Only minimal differences were observed between lines with respect to hemolymph protein concentration or whole-cricket activity of juvenile hormone epoxide hydrolase, the other major JH-degrading enzyme. Dramatic correlated responses to selection, equal in magnitude to the direct response, were observed for JHE activity on each of three other days of the last juvenile stadium. In contrast, no correlated responses in JHE activity were observed in adults. This indicates that JHE activities throughout the last stadium will evolve as a highly correlated unit independent of adult activities and the evolution of endocrine mechanisms regulating juvenile development can be decoupled from those controlling adult reproduction. This study represents the first quantitative-genetic analysis of naturally occurring endocrine variation in an insect species.     

Polygenic Mutation in Drosophila Melanogaster: Genetic Analysis of Selection Lines

We have conducted genetic analyses of 12 long-term selection lines of Drosophila melanogaster derived from a highly inbred base population, containing new mutations affecting abdominal and sternopleural bristle number. Biometric analysis of the number of effective factors differentiating the selected lines from the base inbred indicated that with the exception of the three lines selected for increased number of abdominal bristles, three or more mutations contributed to the responses of the selection lines. Analysis of the chromosomal distribution of effects revealed that mutations affecting abdominal bristle number occurred on all three major chromosomes. In addition, Y-linked mutations with effects ranging from one to three bristles occurred in all three lines selected for decreased number of abdominal bristles, as well as in one line selected for increased abdominal bristle number. Mutations affecting sternopleural bristle number were mainly on the X and third chromosomes. One abdominal and one sternopleural selection line showed evidence of a segregating lethal with large effects on bristle number. As an indirect test for allelism of mutations occurring in different selection lines, the three lines selected in the same direction for the same trait were crossed in all possible combinations, and selection continued from the F(2) hybrids. Responses of the hybrid lines usually did not exceed those of the most extreme parental lines, indicating that the responses of the parental lines may have been partly due to mutations at the same loci, although other interpretations are possible.     

Long-Term Selection Response for 12-Day Litter Weight in Mice

Long-term selection for increased 12-day litter weight in two replicate lines (W₂, W₃) of mice resulted in an apparent selection limit at about 17 generations. Quadratic polynomial and exponential models were fitted to the data in order to estimate the plateaued response and half-life of the selection process. Using the polynomial results, the half-life estimates were 4.5 and 8.6 generations for W₂ and W₃, respectively. The plateaued responses were 5.1 and 5.8 g which, when expressed in phenotypic standard deviation units, became 1.1 and 1.3. The exponential model provides similar estimates. A negative association between 12-day litter weight and fitness was not considered to be an adequate explanation for the plateau since there was no decrease in fertility of the selected lines. Evidence that exhaustion of genetic variability was not the cause of the plateau came from the immediate response to reverse selection. It was proposed that the plateau may be due to a negative genetic correlation between direct and maternal genetic effects, which would be expected to occur after many generations of selection. There were positive correlated responses in both replicates for adult body weight, which was in agreement with the positive genetic correlation between preweaning and postweaning body weight. The expected positive correlated response for number born was realized in only one of the replicates.     

Selection for increased abdominal bristle number in Drosophila melanogaster with concurrent irradiation

Summary Replicate lines, each, with one hundred pairs of parents selected at 50% intensity, were derived from an inbred line. For twenty generations three lines were selected without irradiation and five with 1000 r X-rays per generation given to both females and males. After adjustment for level of crowding in the cultures, the final mean was 1.3 bristles higher in females and 1.0 bristles higher in males in the irradiated lines than in the unirradiated lines. In terms of phenotypic standard deviations in each sex in the base population, these total responses were 0.74 and 0.59 respectively. Radiation can induce mutations useful in increasing responses in selection programmes, but the average response attributable to radiation is small, and heterogeneity between replicate lines is to be expected.This work was carried out while B. H. was on leave from the New South Wales Department of Agriculture, and held a Commonwealth Research Studentship. Our thanks to Robin Hall and Tricia Brown for technical assistance, to Dr. B. L. Sheldon, C.S.I.R.O. Division of Animal Genetics, for providing the inbred line, and to the Radio-Therapy Unit, Royal Prince Alfred Hospital, for the use of their equipment.     

Selection for increased abdominal bristle number in Drosophila melanogaster with concurrent irradiation

Summary Replicate lines, each initially with one hundred pairs of parents selected at 50% intensity, were derived from the Canberra strain. In later generations population size was reduced and selection intensity increased. Three lines were selected without irradiation and five with 1000 r X-rays per generation for thirty generations. Selection was continued until generation 66. Long-term responses were similar in unirradiated and irradiated lines, and there was evidence that genes with large effects influenced response patterns in both groups of lines.This work was carried out while B.H. was on leave from the New South Wales Department of Agriculture, and held a Commonwealth Research Studentship. Robin Hall and Tricia Brown provided much appreciated technical assistance, and we are indebted to the Radio-Therapy Unit, Royal Prince Alfred Hospital, for the use of their equipment.     

Selection for egg production on part records

Summary Responses from four generations of index selection for egg production to 280 days of age in four White Leghorn populations have been presented. A pedigreed randombred population derived from one of the lines was reared with the selected lines to measure the environmental trend. The magnitude of total as well as average response although varying from population to population was positive in all the lines studied. Close correspondence between predicted and realized gains indicated that natural selection, genotype environmental interactions and environmental fluctuations were unimportant during the course of selection. Realized heritabilities agreed fairly well with the estimated heritabilities in at least three out of four populations studied. Probable reasons for variable and insufficient response were investigated.     

Studies on the Scutellar Bristles of DROSOPHILA MELANOGASTER. II. Long-Term Selection for High Bristle Number in the Oregon Rc Strain and Correlated Responses in Abdominal Chaetae

Results are presented of 135 generations of selection for high scutellar bristle number in two lines M and M3 derived from the same original mating of one female with 5 bristles by one male with 4 bristles, the latter being the wild-type canalised phenotype. Results are also given of two relaxed lines per line and of a reselection line M2 derived from the first relaxed line of line M which had regressed almost to base population level. The effect of introducing the sc(1) allele into the M and M3 selected backgrounds was studied at generations 39-44. At the end of selection the effect of an extra dose of sc(+) was also studied in males of all selected backgrounds. The correlated responses in abdominal bristles were followed in all lines.-Considering their common origin, the selection lines differed markedly in pattern of scutellar response and in most other aspects observed, namely correlated responses in abdominals and p.c. scutellars, sex differences, and behaviour on relaxation. Selection limits for scutellar bristles in lines M and M2 were equal to or greater than the most extreme reported in the literature.-The probit span of the canalised 4 bristle class decreased in each selection line as the mean scutellar bristle number increased, and increased again in the relaxed lines as the mean bristle number decreased. In the context of an hypothesis that canalisation at 4 bristle is due to regulation of the scute locus, this result is now interpreted as being due mainly to selection for poor regulators of sc(+), in contrast to a previous interpretation that only the minor gene background was altered by selection, the canalisation (regulation) genotype not being affected.-Introducing the sc(1) allele into the selected backgrounds M and M3 showed a reduced effect on sc(1) flies compared with sc(+) flies, and an interaction of sc(1) and sc(+) with selected background. sc(1) flies had about the same number of bristles in both backgrounds though the mean of sc(+) flies in line M was about 3sigma higher than in line M3. Dominance of sc(+) to sc(1) was reduced slightly in M3. However, the effect of an extra dose of sc(+) at the end of selection was about the same as in unselected in all lines, so the first or dominance level of regulation of the scute locus was not significantly affected by selection, though the second or canalisation level of regulation was.-A large positive correlated response in abdominal bristles occurred in all lines. The response in line M was about twice that in M2 and M3 and was in fact as large as can be obtained from direct selection on abdominals. In line M some genes may have been selected with a proportionately greater effect on abdominals than on scutellars. This is supported by the further observation in line M that the abdominal scores of flies with particular scutellar bristles scores increased as the scutellar mean increased. An attempt was made to apply to these results Rendel's (1962) model of competition between scutellars and abdominals for common bristle-making resources. This could not be done satisfactorily mainly because the assumptions in the model about the similarity of effects in scute and wild-type flies were not met in the present material.     

Selection for components related to body composition in mice: correlated responses

Summary Correlated responses were estimated in each of two replicate lines of mice selected within full-sib families for high (HF) or low (LF) 12-week epididymal fat pad weight as a percentage of body weight (epididymal fat pad percentage), or high (HL) or low (LL) 12-week hind carcass weight as a percentage of body weight (hind carcass percentage). Two replicate control lines (RC) were maintained. Correlated traits were measured in each of the 10 generations of selection. Realized (r_{G R}) and offspring-sire genetic correlations generally were in agreement. In HF and LF, 3–6 week postweaning gain (r_{G R} = 0.36±0.04) and feed intake (r_{G R} = 0.50±0.13) had positive correlated responses, but feed efficiency and feed intake/metabolic body size did not change. Positive correlated responses were found for subcutaneous fat pad percentage, body weight-adjusted subcutaneous fat pad weight and fat percentage in the hind carcass (r_{G R}'s were 1.04±0.13, 0.93±0.13 and 0.90±0.08). In the hind carcass, fat-free dry (protein + ash) percentage showed a small negative correlated response, and fat-free dry weight did not change. In HL and LL, the correlated responses for the above traits were generally opposite to those observed in HF and LF. Litter size, percentage of infertile matings, and preweaning mortality showed no consistent correlated responses in any of the lines, but an index of fitness combining the three traits showed a decrease in all four selection treatments.Paper no. 11057 of the Journal Series of the North Carolina Agricultural Research Service, Raleigh, 27695-7601. The use of trade names in this publication does not imply endorsement by the North Carolina Agricultural Research Service of the products named, nor criticisms of similar ones not mentioned     

Correlated Responses to Selection for Postweaning Gain in the Rat

Evidence for correlated responses to selection was investigated in lines of rats selected for 13 generations for high (U line) and low (D line) 3-9-week gain in comparison with random-bred control lines (R and C lines). The increase in 3-9-week gain in the U lines was shown to be due largely to an increase in 9-week weight, although 3-week weight also increased in these lines. In the D lines, where a marked decrease in 3-9-week gain was observed, this was found to be due to a large decrease in 9-week weight and no detectable change in 3-week weight. The average 2-week litter weight, a measure of the lactational performance of the dam, was significanly greater in the U lines than in the D lines. Selection for 3-9-week gain in these lines of rats led to changes of litter size at birth in the same direction as that of selection. This resulted in a significantly higher litter size in the U lines than in the D lines. The number of rats alive 2 and 9 weeks of age and the percentage of mated females pupping were similar in the U and D lines but lower in these lines than the random bred C lines, providing evidence for a reduction of "fitness" in the selected lines. Carcass composition was studied for all lines at the 11th generation of selection. Carcass composition, in terms of water, fat, ash and protein, was similar in the R and C lines. The U lines had more water and lesss fat than the R or C line. The D lines had similar carcass composition to the R and C lines. It is suggested that these selected and random-bred lines of rats are potentially useful animals to investigate further the developmental and physiological mechanisms which control growth.     

Response to selection for litter size in Danish Landrace pigs: a Bayesian analysis

A replicated selection experiment aimed at increasing litter size (total number of pigs born per litter) in Danish Landrace pigs was conducted from 1984 to 1991. The experiment included two selection and two control lines. In each generation, 30 and 14 first litters were produced in selection and control lines, respectively, and dams produced two litters. Each replicate, consisting of one selection and one control line, was founded from 60 families chosen randomly from the population at large. Family selection was practiced, and the criterion was the predicted breeding value for litter size computed using a repeatability animal model, and taking into account all available information. The data consisted of 947 records from 523 dams (424 dams had two litters) representing five cycles of selection of increased litter size. Data were analyzed from a Bayesian perspective, based on marginal posterior distributions of genetic parameters of interest. Marginalization was achieved using Gibbs sampling, with a single chain length of 1 205 000. After discarding the first 5 000 iterations, a sample was drawn every ten iterations, so 120 000 samples in total were saved. Densities were estimated and plotted, and summary statistics were computed from the estimated densities. The posterior means (± standard error) of heritability and repeatability were 0.22 ± 0.06 and 0.32 ± 0.05, respectively. These point estimates of genetic parameters were within the range of literature values, although on the high side. The posterior mean (± standard error) of genetic response to selection, defined as the difference between the mean breeding values of the selected lines and that of the base population, was 1.37 ± 0.43 pigs after five cycles of selection. The regression (through the origin) of breeding values in the selected lines on generation was 0.25 ± 0.08 pigs. Several informative priors constructed from information obtained with field data in this population were used to examine their influence on inferences. The priors were influential because of the relatively small scale of the experiment. An analysis excluding data from one of the control lines gave smaller genetic variance and heritability, and a smaller response to selection. However, it appears that selection for litter size is effective, but that the true rate of response is probably smaller than data from this experiment suggest.     

Simultaneous selection on two fitness-related traits in the butterfly Bicyclus anynana

Abstract. Theory about the role of constraints in evolution is abundant, but few empirical data exist to describe the consequences a bias in phenotypic variation has for micro evolution. Responses to natural selection can be severely hampered by a genetic correlation among a suite of traits. Constraints can be studied using antagonistic selection experiments, that is, two-trait selection in opposition to this correlation. The two traits studied here were development time and wing pattern (eyespot size) in the butterfly Bicyclus anynana , both of which have a clear adaptive significance. Rates of response were higher for eyespot size than for development time, but were independent of the concurrent selection (either in the same direction as the correlation or perpendicular to it). Regimes differed in both traits in all directions after 11 generations of selection. The uncoupling lines had higher relative responses than the synergistic lines in development time and equal relative responses in eyespot size. The patterns for eyespot size (reaction norms) were consistent across different rearing temperatures. Differences in lines selected for fast and slow development time were more pronounced at lower temperatures, irrespective of the direction of joint wing pattern selection. Furthermore, correlated responses in pupal weight and growth rate were observed; lines selected for a slower development had higher pupal weights, especially at lower temperatures. The response of the uncoupling lines was not hampered by a lack of selectable genetic variation, and the relative response in the development time was larger than expected based on response in the coupled direction and quantitative genetic predictions. This suggests that the structure of the genetic architecture does not constrain the short-term, independent evolution of both wing pattern and development time.