Genetic Variation In Body Size Research Articles

Genetic Variation in Sexual Size Dimorphism Is Associated with Variation in Sex-Specific Plasticity in Drosophila.

AbstractThe difference in body size between females and males, or sexual size dimorphism (SSD), is ubiquitous, yet we have a poor understanding of the developmental genetic mechanisms that generate it and how these mechanisms may vary within and among species. Such an understanding of the genetic architecture of SSD is important if we are to evaluate alternative models of SSD evolution, but the genetic architecture is difficult to describe because SSD is a characteristic of populations, not individuals. Here, we overcome this challenge by using isogenic lineages of Drosophila to measure SSD for 196 genotypes. We demonstrate extensive genetic variation for SSD, primarily driven by higher levels of genetic variation for body size among females than among males. While we observe a general increase in SSD with sex-averaged body size (pooling for sex) among lineages, most of the variation in SSD is independent of sex-averaged body size and shows a strong genetic correlation with sex-specific plasticity, such that increased female-biased SSD is associated with increased body size plasticity in females. Our data are consistent with the condition dependence hypothesis of sexual dimorphism and suggest that SSD in Drosophila is a consequence of selection on the developmental genetic mechanisms that regulate the plasticity of body size.

Transgenerational plasticity and the capacity to adapt to low salinity in the eastern oyster, Crassostrea virginica.

Salinity conditions in oyster breeding grounds in the Gulf of Mexico are expected to drastically change due to increased precipitation from climate change and anthropogenic changes to local hydrology. We determined the capacity of the eastern oyster, Crassostrea virginica, to adapt via standing genetic variation or acclimate through transgenerational plasticity (TGP). We outplanted oysters to either a low- or medium-salinity site in Louisiana for 2 years. We then crossed adult parents using a North Carolina II breeding design, and measured body size and survival of larvae 5 dpf raised under low or ambient salinity. We found that TGP is unlikely to significantly contribute to low-salinity tolerance since we did not observe increased growth or survival in offspring reared in low salinity when their parents were also acclimated at a low-salinity site. However, we detected genetic variation for body size, with an estimated heritability of 0.68 ± 0.25 (95% CI). This suggests there is ample genetic variation for this trait to evolve, and that evolutionary adaptation is a possible mechanism through which oysters will persist with future declines in salinity. The results of this experiment provide valuable insights into successfully breeding low-salinity tolerance in this commercially important species.

Environment-specific heritabilities and maternal effects for body size, morphology and survival in juvenile Atlantic salmon (Salmo salar): evidence from a field experiment

Environmental heterogeneity may strongly influence the amount of heritable variation in phenotypic traits and thus affect evolutionary responses to natural selection. However, the question of whether heritabilities change across environmental gradients has received little empirical attention, particularly for wild vertebrates. We tested whether levels of heritable variation in body size, morphology and survival of juvenile Atlantic salmon (Salmo salar) differed between water flow regimes. We exposed individuals of known genetic relationships to rearing habitats characterized by slow and rapid water flows in a field experiment. We found that the additive genetic variation in body size tended to be higher for individuals reared under rapid water flows. By contrast, the heritabilities of other morphological traits were not consistently higher in either water flow. We also found that salmon grew faster under rapid water flows but also suffered high mortality rates with little heritable variation explaining the variation in survival. However, part of the variation in survival in the rapid water flow was explained by maternal effects. Our results suggest a strong tendency for heritable variation, particularly in body size to be revealed only under specific environmental conditions, such as those that allow for rapid growth. We provide support for the hypothesis that genotype by environment interactions have important effects on the adaptive potential of phenotypes in nature.

Natural variation and the capacity to adapt to ocean acidification in the keystone sea urchin Strongylocentrotus purpuratus

A rapidly growing body of literature documents the potential negative effects of CO2 -driven ocean acidification (OA) on marine organisms. However, nearly all this work has focused on the effects of future conditions on modern populations, neglecting the role of adaptation. Rapid evolution can alter demographic responses to environmental change, ultimately affecting the likelihood of population persistence, but the capacity for adaptation will differ among populations and species. Here, we measure the capacity of the ecologically important purple sea urchin Strongylocentrotus purpuratus to adapt to OA, using a breeding experiment to estimate additive genetic variance for larval size (an important component of fitness) under future high-pCO2 /low-pH conditions. Although larvae reared under future conditions were smaller than those reared under present-day conditions, we show that there is also abundant genetic variation for body size under elevated pCO2 , indicating that this trait can evolve. The observed heritability of size was 0.40±0.32 (95% CI) under low pCO2 , and 0.50±0.30 under high-pCO2 conditions. Accounting for the observed genetic variation in models of future larval size and demographic rates substantially alters projections of performance for this species in the future ocean. Importantly, our model shows that after incorporating the effects of adaptation, the OA-driven decrease in population growth rate is up to 50% smaller, than that predicted by the 'no-adaptation' scenario. Adults used in the experiment were collected from two sites on the coast of the Northeast Pacific that are characterized by different pH regimes, as measured by autonomous sensors. Comparing results between sites, we also found subtle differences in larval size under high-pCO2 rearing conditions, consistent with local adaptation to carbonate chemistry in the field. These results suggest that spatially varying selection may help to maintain genetic variation necessary for adaptation to future OA.

Narrow-sense heritability of body size and its response to different developmental temperatures in Culex quinquefasciatus (Say 1923)

Body size is an important trait involved in overall fitness through its effects on mating success, fecundity, resource acquisition and mortality, and desiccation resistance. In this study, we raised inbred Culex quinquefasciatus mosquito cohorts at different developmental temperatures of 20°, 23°, and 27° C. As an indicator of the amount of genetic variation in body size, we estimated the narrow-sense heritability of body sizes defined as wing aspect ratios. Our results show that narrow-sense heritability of the body size increased as the developmental temperature increased. We also detected the presence of strong genotype-by-environment (G × E) interaction from low cross-environmental correlations. The body size of each temperature regime followed the general rule that higher temperatures produce smaller individuals. We suggest that the increase in genetic variation with increasing temperature might be due to an unleashing of the cryptic genetic variation of the putative genes affecting body size. We conclude that this increase in genetic variation tracking the environmental (developmental temperature) change could have considerable implications for the distribution and range expansion of Cx. quinquefasciatus, especially in warmer environments.

CRYPTIC GENETIC VARIATION AND BODY SIZE EVOLUTION IN THREESPINE STICKLEBACK

The role of environment as a selective agent is well-established. Environment might also influence evolution by altering the expression of genetic variation associated with phenotypes under selection. Far less is known about this phenomenon, particularly its contribution to evolution in novel environments. We investigated how environment affected the evolvability of body size in the threespine stickleback (Gasterosteus aculeatus). Gasterosteus aculeatus is well suited to addressing this question due to the rapid evolution of smaller size in the numerous freshwater populations established following the colonization of new freshwater habitats by an oceanic ancestor. The repeated, rapid evolution of size following colonization contrasts with the general observation of low phenotypic variation in oceanic stickleback. We reared an oceanic population of stickleback under high and low salinity conditions, mimicking a key component of the ancestral environment, and freshwater colonization, respectively. There was low genetic variation for body size under high salinity, but this variance increased significantly when fish were reared under low salinity. We therefore conclude that oceanic populations harbor the standing genetic variation necessary for the evolution of body size, but that this variation only becomes available to selection upon colonization of a new habitat.

Quantifying the effects of genetic selection and genetic variation for body size, carcass composition, and meat quality in the domestic fowl (Gallus domesticus)

A multistrain experiment was conducted to quantify the extent of genetic differences in carcass and muscle yields, muscle quality, support organs, and taste panel assessments of cooked breast muscle of 296 birds from 37 lines of commercial broiler, layer, and traditional chickens. The birds were reared as broilers and 4 males from each line were slaughtered at 6 and 10 wk of age. The extent of genetic variation was measured as the intraclass correlation. The intraclass correlation for live weight; carcass yields; breast, drum, and wing portions; and associated muscle yields were high, whereas those for the thigh portion and yield were low. Broilers had more breast and thigh muscle but similar drum muscle as a proportion of carcass weight compared with layer and traditional lines. Genetic variation for muscle quality (plasma creatine kinase activity) was high; that for muscle color (L, a, and b) and hemorrhage score were moderate in size and were greater at 10 than at 6 wk of age. Broiler lines had greater creatine kinase activity indicative of greater muscle pathology; breast muscle was lighter, less red and yellow in color, and had a greater hemorrhage score than muscle from layer and traditional lines, which were similar. Intraclass correlations for taste panel scores were low and generally not significant except for texture, chicken flavor intensity, flavor liking, and overall liking at 6 wk of age. Significantly greater scores from broiler compared with layer and traditional lines for texture, chicken flavor intensity, and overall liking were observed. At 10 wk of age, chicken flavor intensity did not differ between broiler or layer birds but was significantly greater in both groups than traditional birds. Genetic variation for relative weight of abdominal fat, spleen, and heart was moderately high and greater at 10 than at 6 wk of age. Broiler carcasses had a relatively high proportion of abdominal fat and smaller spleen and heart weights.

Ecological correlates of body size in relation to cell size and cell number: patterns in flies, fish, fruits and foliage

Body size is important to most aspects of biology and is also one of the most labile traits. Despite its importance we know remarkably little about the proximate (developmental) factors that determine body size under different circumstances. Here, I review what is known about how cell size and number contribute to phenetic and genetic variation in body size in Drosophila melanogaster, several fish, and fruits and leaves of some angiosperms. Variation in resources influences size primarily through changes in cell number while temperature acts through cell size. The difference in cellular mechanism may also explain the differences in growth trajectories resulting from food and temperature manipulations. There is, however, a poorly recognized interaction between food and temperature effects that needs further study. In addition, flies show a sexual dimorphism in temperature effects with the larger sex responding by changes in cell size and the smaller sex showing changes in both cell size and number. Leaf size is more variable than other organs, but there appears to be a consistent difference between how shade-tolerant and shade-intolerant species respond to light level. The former have larger leaves via cell size under shade, the latter via cell number in light conditions. Genetic differences, primarily from comparisons of D. melanogaster, show similar variation. Direct selection on body size alters cell number only, while temperature selection results in increased cell size and decreased cell number. Population comparisons along latitudinal clines show that larger flies have both larger cells and more cells. Use of these proximate patterns can give clues as to how selection acts in the wild. For example, the latitudinal pattern in D. melanogaster is usually assumed to be due to temperature, but the cellular pattern does not match that seen in laboratory selection at different temperatures.

Studies in quantitative inheritance. XI. Genetic and environmental correlation between body size and egg production in Drosophila melanogaster. 1957.

1. The variation of body size and egg production between individuals of a laboratory population ofDrosophila melanogaster has been examined. Estimates of genetic and environmental contributions to the phenotypic variation have been derivedby comparing the variability of the flies of the wild population with that of genetically uniform flies from crosses between inbred lines reared under similar favourable conditions. Apparently about 50 % of the variance of body size and about 60 % of the variance of egg production is attributable to genetic segregation. 2. From the results of two-way selection for body size and also egg production, the heritability of the two characters is estimated as respectively 0.43 and 0.18, suggesting that most of the genetic variation in body size behaves as additive in the statistical sense, while non-additive effects predominate in the variability of egg production. 3. There is a clear-cut phenotypic correlation between size and egg production among the individuals of the wild population and this is mostly genetic in origin. Selection for large and small size leads to comparatively little change in egg production. A series of different genotypes, each represented by a number of genetically identical individuals; were prepared by combining haploid sets of chromosomes from the wild population with a haploid set from an inbred line—and these provided no evidence of genetic correlation between size and egg production. Hence the genetic correlation in the wild stock is essentially non-additive in behaviour. 4. Gene-environment interaction has been demonstrated for both egg production and body size by comparing estimates of variance and heritability on the live medium with those found on a synthetic medium which permits normal growth. It is suggested that gene-environment interaction will be particularly important under the suboptimal condition in which the population usually lives. 5. Reduction of the body size of genetically uniform flies by varying the nutrition of the larvae leads to a proportional reduction in the rate at which eggs are laid. 6. The variability of development time has also been examined in preliminary experiments which indicate that about half the variance is genetic in origin. Under favourable conditions there is a high correlation between length of development and body size, while egg production tends to be negatively correlated, suggesting that egg production and growth rate are genetically associated. 7. The results are discussed in relation to the stability of average body size in. the population and the ease with which selection either way leads to striking change. There are strong indications that the apparently continuous variation of attributes such as size or egg production are heterogeneous with respect to the effects on physiology and development which contribute to the final variation, while the relative contribution of different sorts of effect may vary according to environmental conditions.

Spontaneous Mutational Variation for Body Size in Caenorhabditis elegans

We measured the impact of new mutations on genetic variation for body size in two independent sets of C. elegans spontaneous mutation-accumulation (MA) lines, derived from the N2 strain, that had been maintained by selfing for 60 or 152 generations. The two sets of lines gave broadly consistent results. The change of among-line genetic variation between cryopreserved controls and the MA lines implied that broad sense heritability increased by 0.4% per generation. Overall, MA reduced mean body size by approximately 0.1% per generation. The genome-wide rate for mutations with detectable effects on size was estimated to be approximately 0.0025 per haploid genome per generation, and their mean effects were approximately 20%. The proportion of mutations that increase body size was estimated by maximum likelihood to be no more than 20%, suggesting that the amount of mutational variation available for selection for increased size could be quite small. This hypothesis was supported by an artificial selection experiment on adult body size, started from a single highly inbred N2 individual. We observed a strongly asymmetrical response to selection of a magnitude consistent with the input of mutational variance observed in the MA experiment.

Quantitative genetic analysis of natural variation in body size in Drosophila melanogaster.

Latitudinal, genetic variation in body size is a commonly observed phenomenon in many invertebrate species and is shaped by natural selection. In this study, we use a chromosome substitution and a quantitative trait locus (QTL) mapping approach to identify chromosomes and genomic regions associated with adaptive variation in body size in natural populations of Drosophila melanogaster from the extreme ends of clines in South America and Australia. Chromosome substitution revealed the largest effects on chromosome three in both continents, and minor effects on the X and second chromosome. Similarly, QTL analysis of the Australian cline identified QTL with largest effects on the third chromosome, with smaller effects on the second. However, no QTL were found on the X chromosome. We also compared the coincidence of locations of QTL with the locations of five microsatellite loci previously shown to vary clinally in Australia. Permutation tests using both the sum of the LOD scores and the sum distance to nearest QTL peak revealed there were no significant associations between locations of clinal markers and QTL's. The lack of significance may, in part, be due to broad QTL peaks identified in this study. Future studies using higher resolution QTL maps should reveal whether the degree of clinality in microsatellite allele frequencies can be used to identify QTL in traits that vary along an environmental gradient.

Seasonality and genetic architecture of development time and body size of the birch feeding sawfly Priophorus pallipes.

We tested, using the sawfly Priophorus pallipes feeding on leaves of mountain birch, whether the expression of genetic (co)variation of larval development time and body size can be altered by exposing larvae to diets with differential seasonal changes in quality. In nature, larvae feed mainly on mature leaves, but occasionally they are forced to consume senescing leaves. Sixty families were assayed on three experimentally simulated diets: mature leaves of high quality, senescing leaves of rapidly declining quality, and senesced leaves of low quality. The intuitively obvious positive phenotypic and genetic correlations between development time and final mass were observed when the larvae consumed leaves of stable high quality, but low and declining food quality prevented long-growing individuals and families from achieving high final mass, switching the correlations to close to zero or negative in these treatments. The amount of genetic variation for body size showed a non-linear change across the diet quality gradient, whereas genetic variation for development time increased with decreasing diet quality. The among-trait difference in the degree reaction norms crossed along the diet gradient caused the changes in the expression of genetic (co)variation within the environments. Our results show that seasonally varying diet quality induces dramatic changes in the genetic (co)variation of development time and body size, and that simultaneous analysis of reaction norms and environment-specific expression of genetic (co)variation is necessary for the understanding of the genetic characteristics underlying the construction of phenotypes in heterogeneous environments.

Genetic and environmental components of growth in nestling blue tits (Parus caeruleus)

Abstract We investigated the effect of brood-size mediated food availability on the genetic and environmental components of nestling growth in the blue tit (Parus caeruleus), using a cross-fostering technique. We found genetic variation for body size at most nestling ages, and for duration of mass increase, but not of tarsus growth. Hence, nestling growth in our study population seems to have the potential to evolve further. Furthermore, significant genotype–environment interactions indicated heritable variation in reaction norms of growth rates and growth periods, i.e. that our study population had a heritable plasticity in the growth response to environmental conditions. The decreasing phenotypic variance with nestling age indicated compensatory growth in all body traits. Furthermore, the period of weight increase was longer for nestlings growing up in enlarged broods, while there was no difference to reduced broods in the period of tarsus growth. At fledging, birds in enlarged broods had shorter tarsi and lower weights than birds in reduced broods, but there was no difference in wing length or body condition between the two experimental groups. The observed flexibility in nestling growth suggests that growing nestlings are able to respond adaptively to food constraint by protecting the growth of ecologically important traits.

Genetic variance in temperature dependent adult size deriving from physiological genetic variation at temperature boundaries.

An increase in genetic variation in body size has often been observed under stress; an increase in dominance variance and interaction variance as well as in additive genetic variance has been reported. The increase in genetic variation must be caused by physiological mechanisms that are specific to adverse environments. A model is proposed to explain the occurrence of an increase in genetic variation in body size in Drosophila at extreme temperatures. The model has parameters specific to the low- and high-temperature regions of the viable range. Additive genetic variation in the boundary temperatures leads to a marked increase in additive genetic variation in development rate and body size at extreme temperatures. Additive genetic variation in the temperature sensitivity in the low- and high-temperature regions adds non-additive genetic variation. Development rate shows patterns in additive genetic variation that differ from the patterns of genetic variation in body size; therefore, the genetic correlation between development rate and body size changes sign repeatedly as a function of temperature. The existence of dominance in the genetic variation in the boundary temperatures or in the low- and high-temperature sensitivities leads to a higher total genetic variance due to higher dominance and interaction variance, for both development rate and body size.

Genetic variation and causes of genotype-environment interaction in the body size of blue tit (Parus caeruleus).

In several studies of natural populations of birds, the heritability of body size estimated by parent-offspring regression has been lower when offspring have developed in poor feeding regimens than when they developed in good feeding regimens. This has led to the suggestion that adaptation under poor regimens may be constrained by lack of genetic variation. We examined the influence of environmental conditions on expression of genetic variation in body size of nestling blue tits (Parus caeruleus) by raising full sibs in artificially reduced and enlarged broods, corresponding to good and poor feeding regimens, respectively. Individuals grown in the poor regimen attained smaller body size than their sibs grown in the good regimen. However, there was among-family variation in response to the treatments--i.e., genotype-environment interactions (GEIs). Partitioning the GEI variance into contributions attributable to (1) differences in the among-family genetic variance between the treatments and (2) imperfect correlation of genotypic values across treatments identified the latter as the main cause of the GEI. Parent-offspring regressions were not significantly different when offspring were reared in the good environment (h2 = 0.75) vs. when they were reared in the poor environment (h2 = 0.63). Thus, there was little evidence that genetic variance in body size was lower under the poor conditions than under the good conditions. These results do not support the view that the genetic potential for adaptation to poor feeding conditions is less than that for adaptation to good conditions, but they do suggest that different genotypes may be favored under the different conditions.

An Interaction Between Environmental Temperature and Genetic Variation for Body Size for the Fitness of Adult Female Drosophila melanogaster

An Interaction Between Environmental Temperature and Genetic Variation for Body Size for the Fitness of Adult Female Drosophila melanogaster

AN INTERACTION BETWEEN ENVIRONMENTAL TEMPERATURE AND GENETIC VARIATION FOR BODY SIZE FOR THE FITNESS OF ADULT FEMALE DROSOPHILA MELANOGASTER.

- Drosophila and other ectotherms show geographic genetic variation in body size, with larger individuals at higher latitudes and altitudes. Temperature is implicated as an important selective agent because long-term laboratory culture of Drosophila leads to the evolution of larger body size at lower temperatures. In this paper, we tested the hypothesis that, in Drosophila melanogaster, larger size is favored at lower temperatures in part because of selection on adult females. We used replicated lines of D. melanogaster artificially selected for increased and decreased wing area with constant cell area. The resulting size differences between the selected lines were due solely to differences in cell number, and thereby were similar to the cellular basis of clinal variation in body size in nature. We examined life-history traits of adult females at 18 and 25°C. Rearing for two generations at the two temperatures did not affect the extent of the size differences between lines from the different selection regimes. There was a strong interaction between temperature and size selection for both survival and lifetime reproductive success, with larger females living significantly longer and producing more offspring over their lifetime only when reared and tested in the colder environment. There was also an increase in average daily progeny production in large-line females relative to the control and small lines again, only in the colder environment. Thus, the females from the large selection lines were relatively fitter at the colder temperature. At both experimental temperatures, especially the lower one, the small- line females rescheduled their progeny production to later ages. Larger body size may have evolved at higher latitudes and altitudes because of the advantages to the adult female of being larger at lower temperatures.

Expression of Genetic Variation in Body Size of the Collared Flycatcher Under Different Environmental Conditions

Expression of Genetic Variation in Body Size of the Collared Flycatcher Under Different Environmental Conditions

Quantitative genetic variation in body size of mice from new mutations.

To measure the amount of new genetic variation in 6-week weight of mice arising each generation from mutation, selection lines derived from an initially inbred strain were maintained for 25 generations. An analysis using an animal model with restricted maximum likelihood was applied to estimate a mutational genetic component of variance for the infinitesimal model of many genes of small effect. Assuming that the inbred base population was at a mutation-drift equilibrium, it is estimated that the heritability for body size has increased by 1.0% per generation, with lower and upper confidence limits of 0.6% and 1.6%, respectively. A model which includes a mutational genetic component of variance fits the data much better than one involving only base population genetic variance. A model with no genetic component fits the data very poorly. An environmental covariance of body size of mother and offspring was included in the model and accounts for 10% of the variance. By using information only from the observed response to selection, the estimated increase in heritability from mutation is 0.3% per generation. These values are higher than published estimates for the increase in variance from spontaneous mutations in bristle traits of Drosophila, for which there are extensive data, but similar to estimates for various skeletal traits in mice.

Population dynamics and body‐size selection in Daphnia

We examined the population dynamics and seasonal change in body size of a population of Daphnia galeata mendotae experiencing strong vertebrate predation. The intensity of food limitation was quantified seasonally by in situ food addition. We also collected live animals early and late in the summer for laboratory studies designed to estimate genetic variation and seasonal change in life‐history traits for the population.The population was characterized by nearly equal short‐term birth and death rates that were strongly correlated with population density, suggesting density dependence. The population generally exhibited very high birth and death rates and no significant response to experimental food addition. The one exception was June 1988, when the population achieved high densities and exhibited strong food limitation. Throughout both summers, minimal and average body sizes of adults decreased.Laboratory studies document substantial genetic variation for body size and other life‐history traits among animals collected early in summer. By late summer, however, there was little genetic variation for these same traits. Furthermore, clones collected early in summer were genetically larger in body size at birth and maturity than clones collected in late summer. We interpret these results as indicating natural selection for body size.