Asymmetrical responses to automatic selection for body size in Drosophila melanogaster

Summary An apparatus has been made for the automatic selection of Drosophila for body size, operating on the principle of a fractionating sieve. The measurements of individual flies by this method were approximately normally distributed and the repeatability of measurements on successive days was 0.5. A two-way selection experiment for this character was carried out with two replicates for ten generations. The realised heritability for the measured score was 0.14±0.02 for high score and 0.20±0.02 when it was for low score. The correlated response in body weight was asymmetrical, the change downwards being much greater than that upwards. There was a clear divergence in activity measurements between the lines selected in the two directions but no clear trends in fertility. Examination of the selected lines after eleven generations showed that the relationship between score and body weight was clearly different in the lines selected in the two directions and was non-linear in both.It is suggested that the response in activity observed as a consequence of selection for score is partly due to the direct response for activity and partly to a correlated response because of a negative genetic correlation between body size and activity. The observed non-linear relationship between score and body weight observed within generations may be a direct cause of the asymmetry of direct and correlated responses which may also have a parallel in other situations.     

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.     

Mutational variance for pupa weight in Tribolium castaneum

Summary Directional selection for heavier pupa weight in Tribolium castaneum was practiced for 18 generations in two replicates of an inbred line, each separately maintained in small population cages for more than 90 generations. Mutational variance was estimated in two ways, based on Hill's (1982a) prediction equation for response to directional selection where an equilibrium state between effective population size and variation created by new mutation is assumed. Estimates of mutational variance based on response to selection in a selected population and from a sire-offspring regression analysis in an unselected control population were in strong agreement within each replicate population. Significant differences between the two replicates were observed. Estimates of the ratio of mutational variance to environmental variance ranged from 0.0002 to 0.0012, depending upon the assumptions made about effective population sizes maintained in the two replicate lines. Estimates of realized heritability from the 18 generations of selection were 0.23±0.02 and 0.12±0.02 in the two replicates. The results support the hypothesis that mutation may have played a significant role in supplying useful genetic variation for long-continuing response to selection for this trait in experiments reported earlier.     

Analysis of response to 20 generations of selection for body composition in mice: fit to infinitesimal model assumptions

Data were analysed from a divergent selection experiment for an indicator of body composition in the mouse, the ratio of gonadal fat pad to body weight (GFPR). Lines were selected for 20 generations for fat (F), lean (L) or were unselected (C), with three replicates of each. Selection was within full-sib families, 16 families per replicate for the first seven generations, eight subsequently. At generation 20, GFPR in the F lines was twice and in the L lines half that of C. A log transformation removed both asymmetry of response and heterogeneity of variance among lines, and so was used throughout. Estimates of genetic variance and heritability (approximately 50%) obtained using REML with an animal model were very similar, whether estimated from the first few generations of selection, or from all 20 generations, or from late generations having fitted pedigree. The estimates were also similar when estimated from selected or control lines. Estimates from REML also agreed with estimates of realised heritability. The results all accord with expectations under the infinitesimal model, despite the four-fold changes in mean. Relaxed selection lines, derived from generation 20, showed little regression in fatness after 40 generations without selection.     

Direct and correlated responses to selection for increased postweaning gain in mice

Summary Mass selection for increased body weight gain from 21 to 42 days of age was practiced for 12 generations in four replicate lines of ICR-albino mice. Response to selection averaged 0.56±0.03 g. per generation. This response represented an increase of 7.0 genetic standard deviation units and 3.4 phenotypic standard deviation units in 12 generations. The realized heritability pooled over the four replicates was 0.24±0.02. Sizable positive correlated responses were found for 42 and 56-day weight and gain from 42 to 56 days. Much smaller positive correlated responses were noted for 12-day litter weight and 12-day individual weight. Neither litter size nor weaning weight were significantly altered by selection for increased postweaning gain. Reproductive efficiency measured as percent fertile matings declined significantly in the selected lines.Supported in part by a grant from the Virginia Agricultural Foundation.Published with the approval of the Director of the Research Division, Virginia Polytechnic Institute and State University.     

Long-term selection for 8-week body weight in chickens — direct and correlated responses

Summary A bidirectional selection experiment for 8-week body weight in chickens was conducted. In addition to 27 generations of selection, random samples were chosen from each selected line in generations 6, 13, 19 and 26 to initiate lines in which selection was relaxed. Genetic change was evident in the selected high-weight line through the first 75% of the study after which response in the direction opposing selection occurred. Selection for low body weight resulted in considerable reduction in body size, particularly in the last quarter of the study. Correlated responses evaluated were body weight at several ages, conformation, age at onset of lay, various reproductive and egg quality traits and ratio (female: male) of some traits. Data from lines where selection was relaxed indicated that natural selection opposed artificial selection with the effects greater in later generations.     

Correlated response in male and female sterility to selection for pupa weight in Tribolium castaneum

Summary The correlated responses in male and female sterility to 50 generations of individual selection for pupa weight in Tribolium were analyzed. Two replicate lines (S-lines) were selected for heavier pupa weight and stabilizing selection for pupa weight was practiced in two replicate control lines (C-lines). There was close agreement between replicates in both sets of lines for direct and correlated responses. The rate of inbreeding has been constant for all lines (approximately 0.5% per generation).Regression of generation means for pupa weight on generation of selection indicated a significant linear regression in the direct response for both lines. The linear increases of 46 and 55 g. per generation in the S-lines accounted for 98% of the variation among generations and the linear decreases of 5 and 10 g. per generation in the C-lines accounted for 70–90% of the variation in the generation means.Maximum likelihood estimators were used to calculate the frequency of male and female sterility for each generation and line. Average sterility in the base population ranged from about 4 to 12% for both sexes. Polynomial regressions of percent sterility on generation of selection showed that quadratic and higher order regressions were occasionally significant but accounted for a relatively small fraction of the total variation. In the two S-line replicates, linear regression coefficients of percent sterility on generation number were 0.16±.09 and 0.20±.07 for males and 0.72±.08 and 0.54±.08 for females, suggesting a larger correlated response in female than in male sterility. In the C-lines, linear regression coefficients were 0.02±.08 and –.12±.05 for males in the two replicates and –.05±.05 and –.05±.05 for females. Estimates of realized genetic correlations between pupa weight and sterility in the S-lines ranged from 0.04 to 0.14 for males and from 0.14 to 0.37 for females when the heritability of sterility was allowed to take on values from 0.05 to 0.25.Supported by NSF Grants G-1238 and GB-5987, NIH Grant GM-16074 and NIH Fellowship 1 FO2 GM4 5130-01.     

The influence of long-term melatonin administration on basic physiological and metabolic variables of young Wistar:Han rats

The question of effects of long-term melatonin (MEL) administration have not yet been explained sufficiently, especially its metabolic consequences in young persons and animals. The aim of the present study was to analyze the effects of MEL given during prolonged time (for 3 months) and chronically (for 6 months) at the dose of 4 μg/mL of tap water, on the selected metabolic and hormonal parameters in young female and male Wistar:Han (WH) rats. The weights of selected organs, tissues, body weight gains and food and water intake were registered. Six weeks aged rats were adapted to standard housing conditions and light regimen L:D=12:12 h, fed standard laboratory diet and drank tap water (controls) or MEL solution ad libitum; finally they were sacrificed after overnight fasting. Prolonged MEL administration decreased serum glucose concentration and increased triacylglycerol and malondialdehyde concentration/content in the liver in females. In males MEL increased concentrations of serum phospholipids, corticosterone and liver malondialdehyde. MEL treatment reduced the body weight in both sexes and weight of epididymal fat in males, without any alterations of food and water intake. Chronic MEL administration reduced serum glucose concentration and increased concentration/content of glycogen, triacylglycerol and cholesterol in the liver and glycogen concentration/content in heart muscle in males. In females, the significant rise of serum corticosterone concentration and liver malondialdehyde content was recorded. MEL significantly increased liver weight and decreased thymus weight in males. MEL administration increased temporarily water intake in males, body and epididymal fat weights were similar to that in controls. Body weight of MEL drinking females was reduced in the 1^st half of experiment only; the food and water intake did not differ from control group. The response in WH rats on MEL was more prominent as in the Sprague-Dawley strain (our previous studies). Male rats were generally more affected, probably due to higher daily and total consumption of melatonin.     

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.     

ARTIFICIAL SELECTION ON HORN LENGTH-BODY SIZE ALLOMETRY IN THE HORNED BEETLE ONTHOPHAGUS ACUMINATUS (COLEOPTERA: SCARABAEIDAE)

Males of the horned beetle Onthophagus acuminatus Har. (Coleoptera: Scarabaeidae) exhibit horn length dimorphism due to a sigmoidal allometric relationship between horn length and body size: the steep slope of the allometry around the inflection of the sigmoid curve separates males into two groups; those larger than this inflection possess long horns, and those smaller than this inflection have short horns or lack horns. I examined the genetic basis of the allometric relationship between horn length and body size by selecting males that produced unusually long horns, and males that produced unusually short horns, for their respective body sizes. After seven generations of selection, lines selected for relatively long horns had significantly longer horn lengths for a given body size than lines selected for relatively short horns, indicating a heritable component to variation in the allometry. The sigmoidal shape of the allometry was not affected by this selection regime. Rather, selected lines differed in the position of the allometry along the body size axis. One consequence of lateral shifts in this allometric relationship was that the body size separating horned from hornless males (the point of inflection of the sigmoid curve) differed between selection lines: lines in which males were selected for relatively long horns began horn production at smaller body sizes than lines selected for relatively short horns. These results suggest that populations can evolve in response to selection on male horn length through modification of the growth relationship between horn length and body size.     

The lipoxygenase-dependent oxygenation of lipid body membranes is promoted by a patatin-type phospholipase in cucumber cotyledons

Oilseed germination is characterized by the mobilization of storage lipids as a carbon and energy source for embryonic growth. In addition to storage lipid degradation in germinating oilseeds via the direct action of a triacylglycerol lipase (TGL) on the storage lipids, a second degradation pathway that is dependent on a specific lipid body trilinoleate 13-lipoxygenase (13-LOX) has been proposed in several plant species. The activity of this specific 13-LOX leads first to the formation of ester lipid hydroperoxides. These hydroperoxy fatty acids are then preferentially cleaved off by a TGL and serve as a substrate for glyoxysomal β-oxidation. As a prerequisite for triacylglycerol (TAG) mobilization, a partial degradation of the phospholipid monolayer and/or membrane proteins of the oil body has been discussed. Evidence has now been found for both processes: partial degradation of the proteins caleosin and oleosin was observed and simultaneously a patatin-like protein together with transient phospholipase (PLase) activity could be detected at the oil body membranes during germination. Moreover, in vitro experiments with isolated oil bodies from mature seeds revealed that the formation of 13-LOX-derived lipid peroxides in lipid body membranes is increased after incubation with the purified recombinant patatin-like protein. These experiments suggest that in vivo the degradation of storage lipids in cucumber cotyledons is promoted by the activity of a specific oil body PLase, which leads to an increased decomposition of the oil body membrane by the 13-LOX and thereby TAGs may be better accessible to LOX and TGL.     

Selection response for part period egg number and egg mass in chickens — a comparison

Summary Two sublines from each of four flocks of White Leghorn were subjected to two cycles of selection for part year egg number and part year egg mass. A family index (combined selection) was the selection criterion for each of the two traits. The generation means of an unselected pedigreed randombred control, bred and maintained along with the selected sub-lines, were used for correcting short term environmental trends. The sublines selected for a common selection criterion were considered as replicates. The control corrected generation means, when averaged over replicates, gave the mean direct and correlated responses for the selected and different unselected traits. The average genetic change per generation was 2.16 eggs in egg number selected sublines and 146 g of egg mass in egg mass selected sublines. The direct response marginally exceeded the correlated response for both part period egg number and egg mass seen in the corresponding sublines. Except for egg weight the correlated responses for different unselected traits were in the same direction in both egg number and egg mass selected sublines. While the egg weight in egg number lines did not change, its response was positive in egg mass selected lines. A comparison between the lines revealed that the egg mass selected lines matured later and laid heavier but slightly less numerous eggs than the egg number selected sub-lines.     

Artificial selection on larval growth curves in Tribolium: correlated responses and constraints

Body size is often constrained from evolving. Although artificial selection on body size in insects frequently results in a sizable response, these responses usually bear fitness costs. Further, these experiments tend to select only on size at one landmark age, rather than selecting for patterns of growth over the whole larval life stage. To address whether constraints may be caused by larval growth patterns rather than final size, we implemented a function‐valued (FV) trait method of selection, in which entire larval growth curves from Tribolium were artificially selected. The selection gradient function used was previously predicted to give the maximal response and was implemented using a novel selection index in the FV framework. Results indicated a significant response after one generation of selection, but no response in subsequent generations. Correlated responses included increased mortality, increased critical weight, and decreased development time (DT). The lack of response in size and development time after the first generation was likely caused by increased mortality suffered in selected lines; we demonstrated that the selection criterion caused both increased body size and increased mortality. We conclude that artificial selection on continuous traits using FV methods is very efficient and that the constraint of body size evolution is likely caused by a suite of trade‐offs with other traits.     

The genetics of foraging behaviour: artificial selection for food choice in larvae of the fruitfly,Drosophila melanogaster

Artificial selection for choice between high protein content and high energy content of food was performed for 44 generations of fruitfly larvae. There were significant responses to selection both in a line selected to choose high energy food and in a line selected to choose high protein food. Both lines showed a significantly higher preference for the selected food type than did the control line. The line selected for high energy food also showed a significant increase in response over the generations. Since the selection resulted in lines that differed significantly from controls and from each other, there must be some genetic basis for food choice behaviour. Realized heritability was very low in both selected lines. Viability, as measured by larval weight, pupation time, emergence time and adult size, was significantly higher for flies raised on high protein food than for those raised on high energy food.     

Physiological genetics of the response to a high-sucrose diet byDrosophila melanogaster

A diet medium containing 10% (w/v) sucrose can be inferred to be stressful toDrosophila melanogaster from the increased developmental time and reduced size and fecundity of emerging flies. The metabolic basis for this stress and the genetic response to it are of interest from the point of view of both metabolic regulation and the evolutionary genetics of adaptation to stress. Here the effects of a high-sucrose diet on live weight, total protein, stored lipid and glycogen, and crude activities of 12 enzymes involved in energy metabolism were quantified. Assays were done on a large population ofDrosophila that had been acclimated to the laboratory. A collection of eggs was divided to produce two replicate populations maintained on standard medium and two replicates maintained on high-sucrose medium for 133 generations. At the end of this period, both control and sucrose-selected populations were tested on standard and on high-sucrose medium. Results showed that the immediate effect of the high-sucrose diet (compared to standard medium) for both populations was a reduction in live weight and total protein, and activities of many of the enzymes were also reduced by the sucrose treatment, even after adjusting for the weight effect. Selection resulted in several changes on both the standard and the sucrose medium, but the direction of change was not always the same as the acute effect. In no case was there a significant medium by selection-treatment interaction. The pattern of phenotypic correlations did not resolve the reasons for the direction of the genetic responses. Correlations were generally stable across diets and after selection, but there were notable exceptions.     

Experimental comparison of selection methods to improve a non-linear trait in Tribolium

Summary Five lines of Tribolium castaneum (flour beetle) were selected in each of three, replicates for biomass, using quadratic (Q) and linear (L) selection indices, direct selection for biomass (B) and indirect selection for each component trait, i.e. pupal weight (P) and family size (F), respectively, for five generations. The highest response in biomass was obtained in the Q line using an exact quadratic index in which the genetic value for each trait was based on information from both traits. The expected results were that Q selection would not exceed L selection while the L method would be superior with respect to direct selection for biomass (B line); P selection would be better and F selection worse than B selection. Selection was effective for pupal weight in the Q, L, B and P lines, but was not effective for family size, although the Q line increased slightly. The lines had no linear response for family size. Selection for biomass was influenced more by family size than by pupal weight. It is advantageous to include terms such as squares and products in the selection index, even though the response to selection may have been non-linear because of the low heritability of family size and an intermediate optimum for this component trait.     

Metabolism of pyrenedecanoic acid in Epstein-Barr virus-transformed lymphoid cell lines from normal subjects and from a patient with multisystemic lipid storage myopathy

A lymphoid cell line has been established from a patient with multisystemic lipid storage myopathy and showed a major triacylglycerol storage, whereas the content of other neutral lipids and phospholipids was in the normal range. The metabolism of the triacylglycerols has been investigated in this lymphoid cell line from multisystemic lipid storage myopathy as well as in control cells through pulse-chase experiments using 10-(1-pyrene)decanoic acid (P10), a fluorescent fatty acid derivative, as precursor. After 1 h incubation, the uptake of P10 was not significantly different in multisystemic lipid storage myopathy and control lymphoid cells. The amount of fluorescent lipids synthesized by the lymphoid cells was proportional to the concentration of P10 in the culture medium. After 24 h incubation, at any extracellular concentration of P10, the content of P10-labelled triacylglycerols was much higher in multisystemic lipid storage myopathy cells than in controls. Chase experiments showed an impairment in the rate of degradation of biosynthesized triacylglycerols in multisystemic lipid storage myopathy lymphoblasts compared to controls with time of chase (the ratio P10-triacylglycerols/P10-phospholipids increased in mutant cells while it decreased in normal cells). Elsewhere, no enzyme deficiency of the neutral triacylglycerol lipase activity, has been found in multisystemic lipid storage myopathy lymphoid cells.     

Condition factor and hepatosomatic index as estimates of energy status in male three-spined stickleback

Seasonal variation in direct and indirect measures of energy status was examined using estimates of glycogen, lipid and protein levels in a single cohort of male three-spined sticklebacks from an annual population collected each month over one complete year. Condition factor, somatic condition factor and hepatosomatic index (HSI) were calculated as indirect indices of energy status and the accuracy of these indirect measures as predictors of energy status was investigated. Results indicate that both condition factors were significant predictors of energy reserves (lipid, protein, glycogen and total energy), but that the proportion of variance accounted for was small. Both condition factors perform better as predictors of energy content per unit body weight. The HSI was a significant, but a weak predictor of total glycogen levels over the whole year. On a seasonal basis the relationship between HSI and energy reserves was highly variable. These indices are therefore poor predictors of energy reserves in male three-spined sticklebacks.     

Do random fluctuations in daily ration affect the growth rate of juvenile three-spined sticklebacks?

The effect of random fluctuations in daily ration on the growth performance of individual juvenile three-spined sticklebacks Gasterosteus aculeatus was studied in experiments lasting 21 days at 14°C and a photoperiod of 10L:14D. Two mean ration levels were used: a maintenance ration of 2%, and a high ration of 6% of initial body weight per day. For experimental fish, the daily ration varied randomly about the required mean value with a coefficient of variation of 33%. The controls received a constant daily ration. The experiment was replicated in winter (Dec.-Jan.) and spring (Mar.-Apr.). At a given ration, there was no significant difference between the specific growth rates of fish receiving constant or varying ration. Neither the final dry weight, final body water content nor final lipid content (% dry weight) differed significantly. As expected, ration had a significant effect on growth rate, final dry weight and body lipid and water content. There was a significant difference in mean growth rate between the winter and spring replicates. The growth rates observed in these experiments were compared with those predicted from a previously described empirical growth model developed for sticklebacks fed constant rations. The model underestimated mean growth rates.     

An empirical comparison of selection methods for the improvement of biomass

Summary A single generation of upward truncation selection on families with 20% selected was carried out in each of five replicates using Tribolium castaneum as the test organism. The experiment involved eight lines: N — selected for offspring number; W — selected for pupal weight; B — selected for biomass; Q — quadratic index selected; L₂₁ — linear index selected with relative economic weights of 21 offspring number to pupal weight; L₁₁ — linear index selected with relative economic weights of 11 offspring number to pupal weight; L₁₂ — linear index selected with relative economic weights of 12 offspring number to pupal weight; C — an unselected control.Biomass (weight of offspring per family), offspring number, and pupal weight were measured. No differences in response to selection were found among the linear index lines and the pupal weight line with regard to any trait analysed. Generally, response to selection in the linear index lines and pupal weight line was small for offspring number and high for pupal weight. Selection pressure on offspring number in these lines seemed to be dependent on the correlation between offspring number and pupal weight. As a result, response to selection for biomass was poor in the linear index and pupal weight selected lines. In the case of the linear indices, poor response to selection for biomass appeared to be due to the violation of the assumption of additivity of the traits included in the definition of aggregate genotype.The responses in the quadratic index, biomass, and offspring number selected lines were equal with respect to selection for biomass. The response of the quadratic index selected line was less than the responses of the biomass and offspring number selected lines for offspring number, but the response in the quadratic index line was as large as that of any other line included in the experiment and greater than the biomass and offspring number selected lines where pupal weight was the criterion.Highly significant amounts of variation were found for all traits indicating that more replicates are needed for precise evaluation of selection systems.