Individual and group selection for productivity in Tribolium castaneum

Summary The usefulness of a non-random group selection method in the improvement of a character influenced by interactions among individuals has been experimentally tested in a population of Tribolium castaneum. The selected trait —number of adults produced in a fixed period of time —showed a clear increase after 11 generations of selection. This increase is related to a reduction in developmental time, and it was specific for the population structure in which selection was applied, vanishing when this structure was altered.     

Artificial selection for 18-day pupa weight and opposing simulated naturel selection in Tribolium castaneum

Summary The effect of simulated opposing natural selection on the response to mass selection for 18-day pupa weight of Tribolium castaneum was studied for 10 generations of selection. Natural selection was simulated in replicated treatment lines by imposing a negative relationship between mid-parent genetic value for pupa weight and fertility. Responses to selection and realized heritabilities were smaller (P < 0.05 and P < 0.10, respectively) for the treatment lines than for control lines under selection for pupa weight only. One treatment, line E3G1, reached an intermediate selection plateau by generation 10, and responded linearly to 4 generations of artificial selection after natural selection had been discontinued. Possible explanations for the different behaviors of the replicate lines E3G1 and E3G2 were also discussed.     

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.     

Reproductive fitness and artificial selection in animal breeding: culling on fitness prevents a decline in reproductive fitness in lines of Drosophila melanogaster selected for increased inebriation time

Summary The maintenance of reproductive fitness in lines subjected to artificial selection is one of the major problems in animal breeding. The decline in reproductive performance has neither been predictable from heritabilities and genetic correlations, nor have conventional selection indices been adequate to avoid the problem. Gowe (1983) has suggested that the heritabilities of reproductive traits are non-linear, with heritabilities being higher on the lower fitness side. Consequently, he has predicted that culling on reproductive fitness in artificial selection lines will be effective in preventing the usual declines in fitness. An experimental evaluation of Gowe's prediction has been carried out by comparing fitnesses of replicated lines of three treatments: selection for increased inebriation time without culling on fitness (HO), selection for inebriation time with culling of 20% (4/20) of selected females on reproductive fitness (HS), and unselected controls (C). Response to selection for inebriation time in the two selection treatments was similar. After 25 generations, the competitive index, a measure of reproductive fitness, was significantly lower in the HO treatment than the HS treatment, while the HS treatment did not differ from the control lines or the base population. These results demonstrate for the first time that culling on reproductive fitness in selection lines can be used to prevent the usual decline in reproductive performance.     

Epistatic contributions to quantitative traits in Tribolium castaneum

Summary Triple-testcross experiments were used to analyze epistatic contributions to % hatchability of eggs, age of pupation, number of eggs laid in 24-hour period, and survival from hatching to day 35. Seven diverse inbred lines and the F₁ produced by crossing the two tester lines were examined for the presence of epistasis. There was evidence of epistasis for each of the 4 traits in at least one of the 8 lines tested. Epistasis was a major source of variation in survival in all of the lines tested.     

Epistatic contributions to quantitative traits in Tribolium castaneum

Summary Triple-testcross experiments were used to analyze epistatic contributions to larva weight, pupa weight, pupa width and adult weight in Tribolium castaneum. Seven diverse inbred lines and the F₁. produced by crossing the two tester lines were examined for indications of epistasis. Larva weight was the only trait for which no significant epistasis was detected. There was significant epistasis for pupa weight in three of the inbred lines; for pupa width in four of the inbred lines; for adult weight in five of the inbred lines. Only one inbred line and the F₁ line failed to exhibit significant epistasis for any trait. Each inbred line had a unique pattern of epistasis, suggesting that a number of different loci were contributing to the detected epistasis.This paper (No. 76-5-158) is published with the approval of the Director of the Kentucky Agricultural Experiment Station.     

Experimental comparison of methods for simultaneous selection of two correlated traits in Tribolium

Summary Two lines of Tribolium castaneum were selected in each of three replicates for egg laying between 7 and 11 days after adult emergency and for adult weight at 12 days, using theoretical (IT) and empirical (IP) index selection methods. Index coefficients were given empirically in the IP line and they were adjusted in the successive generations of selection according to the results obtained in the previous ones. Highly repeatable selection responses in all replicates occurred in both lines for the aggregate genotype (egg laying plus adult weight) and for each individual trait. The IP line tended to increase slightly more than the IT line for aggregate genotype and egg laying, while the highest response in adult weight was obtained with the IT method. The two methods gave consistently different responses in each replicate. The expected results were that IT selection should not exceed IP selection for the aggregate genotype and egg laying while theoretically the IT method should have been superior for increase adult weight. Theoretical expectations for adult weight have been fulfilled in practice. The IP method would be preferred in a practical sense because of its simplicity and freedom from need of parameter estimation.     

Effects on heterozygosity and reproductive fitness of inbreeding with and without selection on fitness in Drosophila melanogaster

The effects of inbreeding, with (IS) and without selection (IO) for reproductive fitness, on inbreeding depression and heterozygosity were evaluated in 20 lines of each treatment inbred over seven generations using full-sib mating. The survival of lines was significantly greater in IS (20/20) than in IO (15/20). The competitive index measure of reproductive fitness was significantly lower in the inbred lines than in the outbred base population, but not significantly different in surviving IS and IO lines. There was a trend for higher fitness in the IS treatment as relative fitnesses were 19% higher in IS than IO for surviving lines and 59% higher for all lines. Heterozygosities were lower in the inbred lines than in the base population, and significantly higher in the IS than the IO lines. Consequently, the reduction of inbreeding depression in IS has been achieved, at least in part, by slowing the rate of fixation.     

Sites of Fgf signalling and perception during embryogenesis of the beetle Tribolium castaneum

The development of multicellular embryos depends on coordinated cell-to-cell signalling events. Among the numerous cell-signalling pathways, fibroblast growth factors (FGFs) are involved in important processes during embryogenesis, such as mesoderm formation during gastrulation and growth. In vertebrates, the Fgf superfamily consists of 22 family members, whereas only few FGFs are contained in the less complex genomes of insects and worms. In the recently sequenced genome of the beetle Tribolium, we identified four Fgf family members representing three subfamilies. Tribolium has Fgf1 genes that are absent in Drosophila but known from vertebrates. By phylogenetic analysis and microsynteny to Drosophila, we further classify Tc-fgf 8 as an ancestor of pyramus and thisbe, the fly Fgf8 genes. Tc-fgf8 expression in the growth zone suggests an involvement in mesoderm formation. In the embryonic head, expression of Tc-fgf8 subdivides the brain into a larger anterior and a smaller posterior region. The Fgf Tc-branchless is expressed in the embryonic tracheal placodes and in various gland-like structures. The expression patterns of the only Tribolium Fgf receptor and the adaptor molecule Downstream-of-Fgfr are largely congruent with Tc-Fgf8 and Tc-bnl. Thus, in contrast to Drosophila, only one Fgf receptor canalises Fgf signalling in different tissues in Tribolium. Our findings significantly advance our understanding of the evolution of Fgf signalling in insects.     

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.     

The genetic basis of egg lay response to conditioned medium in the flour beetle,Tribolium castaneum

Summary The rate of egg laying in Tribolium castaneum is affected by the quality of the environment and can thus serve not only as a component of fitness of the individual, but also as an adaptation to the long-term survival of the population. In an attempt to determine whether it is an independent genetic character, selection for high and low rates of response was carried jut in beetles from three wild populations. When tests were done on virgin females, and responsiveness, corrected for scale effects, was used as the criterion for selection, separation between the lines was achieved in one generation. It is suggested that in natural populations of Tribolium, the responsiveness of the rate of egg laying to the quality of the environment is controlled by a small number of genes.     

Genetic structure,allozyme-habitat associations and reproductive fitness in Gypsophila fastigiata (Caryophyllaceae)

Relationships between allozyme differentiation, habitat variation and individual reproductive success were examined in local populations of a perennial herb, Gypsophila fastigiata, on the Baltic island of Öland (Sweden). Relatively little (c. 2%) of the total allozyme diversity in this largely outcrossing species is explained by differentiation between sites tens of kilometres apart. The low level of geographic differentiation suggests that gene flow between sites is, or has recently been, extensive. Yet the component of allozyme diversity due to differentiation between plots (only tens of meters apart) within sites is 3 times larger than the between-site component of diversity. Allozyme variation, especially at the Pgi-2 locus, is significantly associated with habitat variation within sites. Different allele x habitat combinations for the Pgi-2 locus are associated with differences in individual reproductive fitness. Differential selection in different local habitats may thus contribute to the fine-scale structuring of genetic diversity within sites.     

Bidirectional selection for body mass and correlated response of pyrethroid resistance and fitness in Sitophilus zeamais

Responses to artificial selection on body mass in the maize weevil Sitophilus zeamais (Coleoptera: Curculionidae) were investigated to determine whether changes in body mass are associated with insecticide susceptibility, rate of population growth, and metabolic rate. Two strains of the maize weevil differing in susceptibility to pyrethroid insecticides were subjected to bidirectional selection on body mass. The susceptible strain responded to selection resulting in individuals with lower or higher body mass, but the resistant strain responded significantly only to selection for lower body mass. The resistant strain selected for low body mass increased its level of deltamethrin resistance in 44 × . In contrast, selection for low body mass in the susceptible parental strain led to increased deltamethrin susceptibility (50 × ) and selection for high body mass increased deltamethrin resistance (4 × ). Thus, the correlated response of insecticide resistance to selection for body mass differed between strains, a likely consequence of their distinct genetic background. Regardless, body mass was positively correlated with fitness (reproductive output) (r = 0.79; P < 0.001), while such correlation with respiration rate was significant only at P = 0.07 (r = 0.44). Therefore, the association between body mass and deltamethrin resistance is population‐dependent in the maize weevil, and the confluence of deltamethrin resistance and high body mass in a given strain will likely favour its energy metabolism and lead to the mitigation of fitness costs usually associated with insecticide resistance. The genetic background and selection history of insecticide resistant populations should not be neglected since they may favour the confluence of insecticide resistance with mitigation mechanisms of its associated fitness costs limiting the tactics available to their management.     

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.     

Directional selection in lines founded from different parts of the phenotypic distribution of sternopleural chaetae number in Drosophila melanogaster

Summary Divergent directional selection lines were initiated from base populations founded from parents taken from different parts of the sternopleural chaetae distribution in a cage population of Drosophila melanogaster. Lines founded from parents taken from the central part of the distribution showed greater response and higher realised heritability than lines derived from parents with extreme high or extreme low chaetae number. The results suggest that centrally derived phenotypes have higher heterozygosity for chaetae factors than extreme phenotypes and that these factors have a large effect on the character.     

Direct and correlated responses to artificial selection on sexual size dimorphism in the flour beetle, Tribolium castaneum

Sexual size dimorphism (SSD) is a conspicuous yet poorly understood pattern across many organisms. Although artificial selection is an important tool for studying the evolution of SSD, previous studies have applied selection to only a single sex or to both sexes in the same direction. In nature, however, SSD likely arises through sex-specific selection on body size. Here, we use Tribolium castaneum flour beetles to investigate the evolution of SSD by subjecting males and females to sexually antagonistic selection on body size (sexes selected in opposite directions). Additionally, we examined correlated responses to body size selection in larval growth rates and development time. After seven generations, SSD remained unchanged in all selected lines; this observed lack of response to short-term selection may be attributed to evolutionary constraints arising from between-sex body size correlations. Developmental traits showed complex correlated responses under different selection treatments. These results suggest that sex-specific larval development patterns may facilitate the evolution of SSD.     

Effect of delayed mating and prolonged engorgement on the reproductive fitness of female Amblyomma limbatum (Acari: Ixodidae) in marginal population areas

Reptile ticks mate while females are attached to their host. Following mating, females engorge, detach and then lay their eggs. This study examines whether the time that females of the reptile tick Amblyomma limbatum spend on hosts prior to mating and the time they take to engorge on hosts after mating influence their reproductive fitness, as measured by the number of viable eggs they produce. When compared with females that experienced no temperature-induced delay in mating, females attached for an initial period to hosts with body temperatures too low to induce mating had no decline in their reproductive fitness, once provided with suitable temperatures for mating. However, on hosts with suitable body temperatures for mating, female ticks that took longer to mate, took longer to engorge and had reduced reproductive fitness. In addition, females that spent more than 20 days feeding on hosts after mating had reduced reproductive fitness, irrespective of the time they spent on hosts prior to mating compared with females with shorter feeding periods. Thus, the time that A. limbatum females spent on hosts prior to mating and/or after mating has a significant bearing on the number viable progeny produced. The ecological implications of these results are discussed in relation to females colonizing marginal population areas at the edge of their distributional range.