Genetic control ofHylemya antiqua III. Differences in pupation ability between two strains

Some relevant traits of a wild (L) and a laboratory (C) strain of Hylemya antiqua (Meigen ), determining differences in their pupation ability under experimental conditions have been investigated in relation to genetic control. The wild strain showed an intrinsic higher pupation ability than the laboratory strain. The minimum feeding period was 0.6 days longer for the C strain. The minimum larval dry weight was different for the two strains. With a normal feeding period C larvae pupated on average 1.63 days later than L larvae. By the shortening of the larval feeding period an acceleration of the larval development of both strains was observed: the acceleration of development was more marked for the wild strain. This result has been contrasted with published works on Drosphila. The consequences of these differences as far as the competitive ability of each strain is concerned, have been discussed in relation to genetic control.     

Intra- and interspecies variations in pupation height in Drosophila.

In order to study the pupation site preference which is an important component of larval behaviour, pupation height was scored in Drosophila ananassae, D. bipectinata and D. malerkotliana by using a large number of strains of these species. The mean pupation height of different strains ranged from 1.1 to 8.7 mm in D. ananassae, from 0.41 to 0.75 mm in D. bipectinata and from 1.0 to 1.3 mm in D. malerkotliana. The analysis of variance and t-test were performed to test intre- and interspecies variations in pupation height. These tests revealed significant variation among three species. Significant variations among different strains of the same species were also found in D. ananassae and D. bipectinata. These observations provide evidence for intra- and interspecies variations in pupation height in Drosophila. Variations among different strains of the same species in pupation height are attributable to genetic heterogeneity among the strains.     

Associations of allozyme variation and behavior in the cowpea weevil (Callosobruchus maculatus)

The cowpea weevil (Callosobruchus maculatus (Fabré)) exhibits several behavioral traits that are stable within, but vary among, strains. These traits are heritable and quantitative. We used cellulose acetate gel electrophoresis to quantify allozyme variation within and among laboratory cultures of four weevil strains and determine whether allozyme variation correlates with behavioral traits. Significant variation exists at 8 of 11 loci assayed and gene frequencies are significantly different among strains. The South Indian strain (SI) is most variable and measures of genetic distance set it apart from the other strains. It is also behaviorally unique. The Brazilian strain (BC) is most different from SI in allozyme diversity and behavioral phenotype, while two African strains (IITA, CAM) are intermediate in allozyme diversity and phenotype. These results are consistent with the known history of these strains and the differences in the allozymes parallel the differences in behavioral traits.     

Ontogenies in mice selected for high voluntary wheel-running activity. I. Mean ontogenies

The evolutionary importance of postnatal ontogenies has long been recognized, but most studies of ontogenetic trajectories have focused exclusively on morphological traits. For animals, this represents a major omission because behavioral traits and their ontogenies often have relatively direct relationships to fitness. Here four replicate lines of house mice artificially selected for high early-age wheel running and their four replicate control lines were used to evaluate the effects of early-age directional selection, genetic drift, and activity environment (presence or absence of a running wheel) on variation in the ontogenies of three traits known to be genetically correlated: voluntary wheel running, body mass, and food consumption. Early-age selection significantly changed both the shape and position of the wheel-running and food-consumption ontogenies while influencing the position, but not the shape, of the body mass ontogeny. Genetic drift (as indicated by variation among replicate lines) produced significant changes in both the position and shape of all three ontogenies; however, its effect differed between the selection and control groups. For wheel running and food consumption, genetic drift only influenced the control ontogenies, whereas for body mass, genetic drift had a significant effect in both selection groups. Both body-mass and food-consumption ontogenies were significantly altered by activity environment, with the environment causing significant changes in the shape and position of both ontogenies. Overall the results demonstrate strong effects of early-age selection, genetic drift, and environmental variation on the evolution and expression of behavioral and morphological ontogenies, with selection changing only the position of the morphological ontogeny but both the position and shape of the behavioral ontogenies.     

Heritability, correlations and in silico mapping of locomotor behavior and neurochemistry in inbred strains of mice

The midbrain dopamine system mediates normal and pathologic behaviors related to motor activity, attention, motivation/reward and cognition. These are complex, quantitative traits whose variation among individuals is modulated by genetic, epigenetic and environmental factors. Conventional genetic methods have identified several genes important to this system, but the majority of factors contributing to the variation remain unknown. To understand these genetic and environmental factors, we initiated a study measuring 21 behavioral and neurochemical traits in 15 common inbred mouse strains. We report trait data, heritabilities and genetic and non-genetic correlations between pheno-types. In general, the behavioral traits were more heritable than neurochemical traits, and both genetic and non-genetic correlations within these trait sets were high. Surprisingly, there were few significant correlations between the behavioral and the individual neurochemical traits. However, striatal serotonin and one measure of dopamine turnover (DOPAC/DA) were highly correlated with most behavioral measures. The variable accounting for the most variation in behavior was mouse strain and not a specific neurochemical measure, suggesting that additional genetic factors remain to be determined to account for these behavioral differences. We also report the prospective use of the in silico method of quantitative trait loci (QTL) analysis and demonstrate difficulties in the use of this method, which failed to detect significant QTLs for the majority of these traits. These data serve as a framework for further studies of correlations between different midbrain dopamine traits and as a guide for experimental cross designs to identify QTLs and genes that contribute to these traits.     

Identification of a genetic network for an ecologically relevant behavioural phenotype in Drosophila melanogaster

Pupation site choice of Drosophila third‐instar larvae is critical for the survival of individuals, as pupae are exposed to various biotic and abiotic dangers while immobilized during the 3–4 days of metamorphosis. This singular behavioural choice is sensitive to both environmental and genetic factors. Here, we developed a high‐throughput phenotyping approach to assay the variation in pupation height in Drosophila melanogaster, while controlling for possibly confounding factors. We find substantial variation of mean pupation height among sampled natural stocks and we show that the Drosophila Genetic Reference Panel (DGRP) reflects this variation. Using the DGRP stocks for genome‐wide association (GWA) mapping, 16 loci involved in determining pupation height could be resolved. The candidate genes in these loci are enriched for high expression in the larval central nervous system. A genetic network could be constructed from the candidate loci, which places scribble (scrib) at the centre, plus other genes known to be involved in nervous system development, such as Epidermal growth factor receptor (Egfr) and p53. Using gene disruption lines, we could functionally validate several of the initially identified loci, as well as additional loci predicted from network analysis. Our study shows that the combination of high‐throughput phenotyping with a genetic analysis of variation captured from the wild can be used to approach the genetic dissection of an environmentally relevant behavioural phenotype.     

Developmental and genetic effects on behavioral and life‐history traits in a field cricket

A fundamental goal of evolutionary ecology is to identify the sources underlying trait variation on which selection can act. Phenotypic variation will be determined by both genetic and environmental factors, and adaptive phenotypic plasticity is expected when organisms can adjust their phenotypes to match environmental cues. Much recent research interest has focused on the relative importance of environmental and genetic factors on the expression of behavioral traits, in particular, and how they compare with morphological and life‐history traits. Little research to date examines the effect of development on the expression of heritable variation in behavioral traits, such as boldness and activity. We tested for genotype, environment, and genotype‐by‐environment differences in body mass, development time, boldness, and activity, using developmental density treatments combined with a quantitative genetic design in the sand field cricket (Gryllus firmus). Similar to results from previous work, animals reared at high densities were generally smaller and took longer to mature, and body mass and development time were moderately heritable. In contrast, neither boldness nor activity responded to density treatments, and they were not heritable. The only trait that showed significant genotype‐by‐environment differences was development time. It is possible that adaptive behavioral plasticity is not evident in this species because of the highly variable social environments it naturally experiences. Our results illustrate the importance of validating the assumption that behavioral phenotype reflects genetic patterns and suggest questions about the role of environmental instability in trait variation and heritability.     

Host plant exodus and larval wandering behaviour in a butterfly: diapause generation larvae wander for longer periods than do non‐diapause generation larvae

1. Prior to pupation, lepidopteran larvae enter a wandering phase lasting up to 30 h before choosing a pupation site. Because stillness is important for concealment, this behaviour calls for an adaptive explanation. 2. The explanation most likely relates to the need to find a suitable pupation substrate, especially in terms of shelter from predation, and given that many predators and parasitoids use host plants as prey‐location cues, mortality probably decreases with distance from the host plant. Hence, remaining on the host includes a long‐term risk, while moving away from the host introduces an increased risk during locomotion. 3. Bivoltine species that overwinter in the pupal stage produce two kinds of pupae; non‐diapausing pupae from which adults emerge after 1–2 weeks, or diapausing pupae that overwinter with adults emerging after 8–10 months. 4. Given the hypothesis of distance‐from‐host‐plant‐related predation, this should select for phenotypic plasticity with larvae in the diapausing generation having a longer wandering phase than larvae under direct development, if there is a trade‐off between mortality during the wandering phase and accumulated mortality during winter. 5. Here this prediction is tested by studying the duration of the wandering period in larvae of the partially bivoltine swallowtail butterfly, Papilio machaon, under both developmental pathways. 6. The results are in agreement with the predictions and show that the larval wandering phase is approximately twice as long under diapause development. The authors suggest that the longer duration of the wandering phase in the diapause generation is a general phenomenon in Lepidoptera.     

Selection on olfactory response to semiochemicals from a plant-host complex in a parasitic wasp

Parasitic wasps (parasitoids) use volatiles from the plants infested by phytophagous insects to locate host herbivores, but their behavioural response to such semiochemicals is highly variable. Bi-directional selection on Cotesia glomerata (Hymenoptera: Braconidae) was conducted to investigate the importance of genetic variation in the olfactory response of parasitoids. Female wasps were assessed for flight orientation and landing success in response to the hexane extract from a plant-host complex in a wind tunnel. After the first generation of selection, two strains significantly differentiated in both flight orientation and landing success, and their divergence continued with further selection. The two selected strains genetically differentiated in olfactory perception rather than upwind flight ability. The realized heritability was estimated as 0.248 for flight orientation and 0.216 for landing success. The selection experiment further demonstrated that a prior exposure to the semiochemicals significantly enhanced the subsequent response of female wasps, independent of genetic differences. These results suggest that both genetic component and environmental conditioning have played an important role in the evolution of host selection and utilization by the parasitoid in a tritrophic system.     

The role of abiotic factors in the pupation of Thaumatotibia leucotreta Meyrick (Lepidoptera: Tortricidae) in the soil

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Experimental evolution of evolutionary potential in fluctuating environments

Variation is the raw material for evolution. Evolutionary potential is determined by the amount of genetic variation, but evolution can also alter the visibility of genetic variation to natural selection. Fluctuating environments are suggested to maintain genetic variation but they can also affect environmental variance, and thus, the visibility of genetic variation to natural selection. However, experimental studies testing these ideas are relatively scarce. In order to determine differences in evolutionary potential we quantified variance attributable to population, genotype and environment for populations of the bacterium Serratia marcescens. These populations had been experimentally evolved in constant and two fluctuating environments. We found that strains that evolved in fluctuating environments exhibited larger environmental variation suggesting that adaptation to fluctuations has decreased the visibility of genetic variation to selection.     

Study of Natural Genetic Variation in Early Fitness Traits Reveals Decoupling Between Larval and Pupal Developmental Time in <Emphasis Type="Italic">Drosophila melanogaster</Emphasis>

Characterizing the relationships between genotype and phenotype for developmental adaptive traits is essential to understand the evolutionary dynamics underlying biodiversity. In holometabolous insects, the time to reach the reproductive stage and pupation site preference are two such traits. Here we characterize aspects of the genetic architecture for Developmental Time (decomposed in Larval and Pupal components) and Pupation Height using lines derived from three natural populations of Drosophila melanogaster raised at two temperatures. For all traits, phenotypic differences and variation in plasticity between populations suggest adaptation to the original thermal regimes. However, high variability within populations shows that selection does not exhaust genetic variance for these traits. This could be partly explained by local adaptation, environmental heterogeneity and modifications in the genetic architecture of traits according to environment and ontogenetic stage. Indeed, our results show that the genetic factors affecting Developmental Time and Pupation Height are temperature-specific. Varying relationships between Larval and Pupal Developmental Time between and within populations also suggest stage-specific modifications of genetic architecture for this trait. This flexibility would allow for a somewhat independent evolution of adaptive traits at different environments and life stages, favoring the maintenance of genetic variability and thus sustaining the traits’ evolvabilities.     

The contribution of ancestry,chance, and past and ongoing selection to adaptive evolution

The relative contributions of ancestry, chance, and past and ongoing election to variation in one adaptive (larval feeding rate) and one seemingly nonadaptive (pupation height) trait were determined in populations ofDrosophila melanogaster adapting to either low or high larval densities in the laboratory. Larval feeding rates increased rapidly in response to high density, and the effects of ancestry, past selection and chance were ameliorated by ongoing selection within 15–20 generations. Similarly, in populations previously kept at high larval density, and then switched to low larval density, the decline of larval feeding rate to ancestral levels was rapid (15-20 generations) and complete, providing support for a previously stated hypothesis regarding the costs of faster feeding inDrosophila larvae. Variation among individuals was the major contributor to variation in pupation height, a trait that would superficially appear to be nonadaptive in the environmental context of the populations used in this study because it did not diverge between sets of populations kept at low versus high larval density for many generations. However, the degree of divergence among populations (FST) for pupation height was significantly less than expected for a selectively neutral trait, and we integrate results from previous studies to suggest that the variation for pupation height among populations is constrained by stabilizing selection, with a flat, plateau-like fitness function that, consequently, allows for substantial phenotypic variation within populations. Our results support the view that the genetic imprints of history (ancestry and past selection) in outbreeding sexual populations are typically likely to be transient in the face of ongoing selection and recombination. The results also illustrate the heuristic point that different forms of selection-for example directional versus stabilizing selection—acting on a trait in different populations may often not be due to differently shaped fitness functions, but rather due to differences in how the fitness function maps onto the actual distribution of phenotypes in a given population. We discuss these results in the light of previous work on reverse evolution, and the role of ancestry, chance, and past and ongoing selection in adaptive evolution.     

Population and behaviour genetics of Drosophila ananassae

Drosophila ananassae is a cosmopolitan and domestic species. It occupies a nuique status among the Drosophila species due to certain peculiarities in its genetic behaviour. The most unusual feature of this species is spontaneous male recombination in appreciable frequency. The present review summarises the work done on population and behaviour genetics of D. ananassae from India. Population dynamics of three cosmopolitan inversions has been studied in Indian population of D. ananassae and it is evident from the results that there is a considerable degree of genetic divergence at the level of inversion polymorphism. In general, the populations from south India show more differentiation than those from the north. These three cosmopolitan inversions, which are coextensive with the species, exhibit heterosis. Interracial hybridization does not lead to beaakdown of heterosis, which suggests that evidence for coadaptation is lacking in geographic populations of D. ananassae. Heterosis appears to be simple luxuriance rather than populational heterosis (coadaptation). Unlinked inversions occur in random associations, indicating no interchromosomal interactions. However, two inversions of the third chromosome often show strong linkage disequilibrium in laboratory populations, which is due to epistatic gene interaction and suppression of crossing-over. Genetic variations for certain allozyme polymorphism and sternoleural bristle phenotypes in Indian populations of D. ananassae have also been observed.A number of investigations have also been carried out on certain aspects of behaviour genetics of Indian D. ananassae. There is evidence for sexual isolation within D. ananassae. Significant variations in mating propensity of several isofemale strains, inversion karyotypes, the diminishing effects of certain mutations on sexual activity of males and positive response to selection for high and low mating propensity provide evidence for genetic control of sexual behaviour in D. ananassae. Males contribute more to variation and thus are more subject to intra-sexual selection than females. Evidence for rare male mating advantage has also been presented. Geographic strains of D. ananassae show variation with respect to oviposition site preference. The results of studies on pupation site preference, which is an important component of larval behaviour, suggest that larval pupation behaviour in D. ananassae is under polygenic control with a substantial amount of additive genetic variation.     

Genetic structure of phenotypic robustness in the collaborative cross mouse diallel panel

Developmental stability and canalization describe the ability of developmental systems to minimize phenotypic variation in the face of stochastic micro‐environmental effects, genetic variation and environmental influences. Canalization is the ability to minimize the effects of genetic or environmental effects, whereas developmental stability is the ability to minimize the effects of micro‐environmental effects within individuals. Despite much attention, the mechanisms that underlie these two components of phenotypic robustness remain unknown. We investigated the genetic structure of phenotypic robustness in the collaborative cross (CC) mouse reference population. We analysed the magnitude of fluctuating asymmetry (FA) and among‐individual variation of cranial shape in reciprocal crosses among the eight parental strains, using geometric morphometrics and a diallel analysis based on a Bayesian approach. Significant differences among genotypes were found for both measures, although they were poorly correlated at the level of individuals. An overall positive effect of inbreeding was found for both components of variation. The strain CAST/EiJ exerted a positive additive effect on FA and, to a lesser extent, among‐individual variance. Sex‐ and other strain‐specific effects were not significant. Neither FA nor among‐individual variation was associated with phenotypic extremeness. Our results support the existence of genetic variation for both developmental stability and canalization. This finding is important because robustness is a key feature of developmental systems. Our finding that robustness is not related to phenotypic extremeness is consistent with theoretical work that suggests that its relationship to stabilizing selection is not straightforward.     

Intrapopulation Genome Size Variation in D. melanogaster Reflects Life History Variation and Plasticity

We determined female genome sizes using flow cytometry for 211 Drosophila melanogaster sequenced inbred strains from the Drosophila Genetic Reference Panel, and found significant conspecific and intrapopulation variation in genome size. We also compared several life history traits for 25 lines with large and 25 lines with small genomes in three thermal environments, and found that genome size as well as genome size by temperature interactions significantly correlated with survival to pupation and adulthood, time to pupation, female pupal mass, and female eclosion rates. Genome size accounted for up to 23% of the variation in developmental phenotypes, but the contribution of genome size to variation in life history traits was plastic and varied according to the thermal environment. Expression data implicate differences in metabolism that correspond to genome size variation. These results indicate that significant genome size variation exists within D. melanogaster and this variation may impact the evolutionary ecology of the species. Genome size variation accounts for a significant portion of life history variation in an environmentally dependent manner, suggesting that potential fitness effects associated with genome size variation also depend on environmental conditions.     

A genetic correlation between age at pupation and melanization immune response of the yellow fever mosquito Aedes aegypti

To investigate the evolutionary cost of an immune response, we selected six lines of the mosquito Aedes aegypti for earlier or later pupation and measured the extent to which this selection procedure changed the mosquito's ability to encapsulate and melanize a negatively charged Sephadex bead. After 10 generations of selection, the age at pupation in the two selection regimes differed by about 0.7 days, accompanied by an increase of wing length of the mosquitoes selected for late pupation. Among the mosquitoes that had been selected for early pupation, only 6% had strongly or completely melanized the bead, while among the individuals that had been selected for late pupation, 32% had melanized the bead. Thus, our results suggest a genetic correlation between age at pupation and immunocompetence. As a consequence, mosquitoes that respond to increased intense parasite pressure with more effective immunity are predicted to pay for the increased defense with slower development.     

Fine-scale abundance and distribution of wild silk moth pupae

Although several factors influence herbivore insect distributions at any particular scale, the most important determinants are likely to differ between species with different life histories. Identifying what these factors are and how they relate to life history forms an important component of understanding the population dynamics of species, and the habitat requirements necessary for their conservation. The pupal stage of two wild silk moth species, Gonometa postica Walker and G. rufobrunnea Aurivillius (Lepidoptera: Lasiocampidae), is the target of harvesting practices that are totally dependent on the availability of pupae from natural populations. Consequently, and partly due to poor knowledge of the species' biology, there is substantial interest in the distribution of pupae among and within trees for both these species. It was investigated whether between- and within-tree pupal distributions in these two species are non-random, and if so, whether there are relationships between pupation site use and tree characteristics such as tree size, available pupation space and branch position. Between-tree patterns in pupal abundance were random in terms of absolute spatial position, but markedly non-random with respect to tree characteristics. The apparent G. postica pupae were aggregated on large larval host plants, whereas the cryptic G. rufobrunnea pupae were aggregated on non-host plants. These patterns reflect the life history differences of the two species. In contrast, at the within-tree scale, branch position, aspect and tree shape influenced pupation site choice similarly for both species. These patterns might be related to microclimate. Documenting between-tree and within-tree patterns in Gonometa pupal distributions is the first step towards explaining pupation site selection, as well as identifying possible evolutionarily selective factors in the species, and generating testable hypotheses from these.     

Lack of evidence for local adaptation to individual plant clones or site by a mobile specialist herbivore

Several studies have documented local adaptation by sedentary insects to individual phenotypes of their host plants. Here, I examined whether a similar phenomenon could be found in a mobile, specialized insect, the sumac flea beetle. Previous work has shown that sumac individuals differ in their suitability as hosts for these beetles and that differences have both an environmental and a genetic basis. Using beetle populations collected as eggs from eight different sumac clones along an east-west transect, a reciprocal transfer experiment was conducted to determine whether there was any evidence for local adaptation by beetles to individual plant clones or to site. Variables examined were larval survivorship past first instar, development time, weight at pupation and patterns of predation by enemies. While no evidence for local adaptation was found, there were significant effects of plant clone on which larvae developed, origin of the larval population and the interaction of these effects on larval performance. For larval weight at pupation, there was also some indication that trade-offs may exist in ability of larvae to use different host plant clones. In addition, there were significant environmental effects on several measures of larval performance. Predation rates differed by plant clone, but not by site or with respect to origin of larvae. While no evidence for local adaptation was found in this study, prerequisites for finding such patterns may exist in this system. Received: 23 May 1996 / Accepted: 26 September 1996     

Selection of Pupation Habitats by Oriental Fruit Fly Larvae in the Laboratory

We performed a series of laboratory experiments to determine the effects of shade, soil moisture, and soil compaction on the selection of pupation habitats by wandering late-instar Oriental fruit flies, Bactrocera dorsalis (Hendel). Larvae showed a strong preference toward pupating in shaded rather than brightly lit areas, in moist rather than dry soil, and in soil with larger particle sizes. These behavioral preferences are likely to lead to clumped distribution of Oriental fruit fly pupae in natural habitats. The implications of this for management of localized populations by chemical and biological methods are discussed.