Significant Additive Genetic Variation Research Articles

Genetic variation in male-induced harm in Drosophila melanogaster.

In Drosophila melanogaster, prolonged exposure to males reduces the longevity and fecundity of females. This harm arises from the effects of male courtship behaviours and the toxic side effects of the accessory gland proteins (Acps) in their seminal fluids. Here, we examine the relationship between male exposure and its harmful effect on the lifetime fitness of his mates, and quantify the genetic basis for this variation. We found significant additive genetic variation in the magnitude of harm that males impose on females by exposing females to males from a variety of hemiclonal backgrounds for either a brief or prolonged period of time and measuring their fecundity, a meaningful fitness index. Furthermore, we discovered a strong negative correlation between the magnitude of harm and the short-term effects of male exposure on female fitness. We discuss the evolutionary significance of these results with regards to potential life-history trade-offs in females, and its relationship to male body size.

Genetic control of Eucalyptus globulus harvest traits

The cost of harvesting short-rotation plantation eucalypts can be in excess of AU$2500·ha−1. Despite this high cost, the extent to which harvesting productivity is affected by tree genetics is not well understood. We address this issue in a study of two 10-year-old genetic field trials of Eucalyptus globulus Labill. in Australia. Standing-tree traits analysed were survival, diameter at breast height, basal area, and stem straightness and forking. Harvest traits analysed were volume, time, and productivity. Genetic group and within-group genetic variation (additive and dominance), stand-level family variation, phenotypic and genetic correlations, and the effects of inbreeding were estimated for these traits. The different scenarios studied showed that plantation harvest productivity was affected by tree genetics to some degree but was mainly affected through positive covariation with stem diameter. Harvest productivity is thus unlikely to have been adversely affected by past selection. Although no significant additive or dominance genetic variation in stem forking or straightness was detected, weak phenotypic correlations were consistent with harvest productivity being higher in straighter trees with no forking. High inbreeding depression was evident for growth and survival; however, in open-pollinated progeny, this resulted in only a slight reduction in harvest productivity (5.5%) compared with out-crossed progeny.

Additive genetic variation for tolerance to estrogen pollution in natural populations of Alpine whitefish (Coregonus sp., Salmonidae).

The evolutionary potential of natural populations to adapt to anthropogenic threats critically depends on whether there exists additive genetic variation for tolerance to the threat. A major problem for water-dwelling organisms is chemical pollution, and among the most common pollutants is 17α-ethinylestradiol (EE2), the synthetic estrogen that is used in oral contraceptives and that can affect fish at various developmental stages, including embryogenesis. We tested whether there is variation in the tolerance to EE2 within Alpine whitefish. We sampled spawners from two species of different lakes, bred them in vitro in a full-factorial design each, and studied growth and mortality of embryos. Exposure to EE2 turned out to be toxic in all concentrations we tested (≥1 ng/L). It reduced embryo viability and slowed down embryogenesis. We found significant additive genetic variation in EE2-induced mortality in both species, that is, genotypes differed in their tolerance to estrogen pollution. We also found maternal effects on embryo development to be influenced by EE2, that is, some maternal sib groups were more susceptible to EE2 than others. In conclusion, the toxic effects of EE2 were strong, but both species demonstrated the kind of additive genetic variation that is necessary for an evolutionary response to this type of pollution.

Hemiclonal analysis of interacting phenotypes in male and female Drosophila melanogaster.

BackgroundIdentifying the sources of variation in mating interactions between males and females is important because this variation influences the strength and/or the direction of sexual selection that populations experience. While the origins and effects of variation in male attractiveness and ornamentation have received much scrutiny, the causes and consequences of intraspecific variation in females have been relatively overlooked. We used cytogenetic cloning techniques developed for Drosophila melanogaster to create “hemiclonal” males and females with whom we directly observed sexual interaction between individuals of different known genetic backgrounds and measured subsequent reproductive outcomes. Using this approach, we were able to quantify the genetic contribution of each mate to the observed phenotypic variation in biologically important traits including mating speed, copulation duration, and subsequent offspring production, as well as measure the magnitude and direction of intersexual genetic correlation between female choosiness and male attractiveness.ResultsWe found significant additive genetic variation contributing to mating speed that can be attributed to male genetic identity, female genetic identity, but not their interaction. Furthermore we found that phenotypic variation in copulation duration had a significant male-associated genetic component. Female genetic identity and the interaction between male and female genetic identity accounted for a substantial amount of the observed phenotypic variation in egg size. Although previous research predicts a trade-off between egg size and fecundity, this was not evident in our results. We found a strong negative genetic correlation between female choosiness and male attractiveness, a result that suggests a potentially important role for sexually antagonistic alleles in sexual selection processes in our population.ConclusionThese results further our understanding of sexual selection because they identify that genetic identity plays a significant role in phenotypic variation in female behaviour and fecundity. This variation may be potentially due to ongoing sexual conflict found between the sexes for interacting phenotypes. Our unexpected observation of a negative correlation between female choosiness and male attractiveness highlights the need for more explicit theoretical models of genetic covariance to investigate the coevolution of female choosiness and male attractiveness.

No Evidence for Heritability of Male Mating Latency or Copulation Duration across Social Environments in Drosophila melanogaster

A key assumption underpinning major models of sexual selection is the expectation that male sexual attractiveness is heritable. Surprisingly, however, empirical tests of this assumption are relatively scarce. Here we use a paternal full-sib/half-sib breeding design to examine genetic and environmental variation in male mating latency (a proxy for sexual attractiveness) and copulation duration in a natural population of Drosophila melanogaster. As our experimental design also involved the manipulation of the social environment within each full-sibling family, we were able to further test for the presence of genotype-by-environment interactions (GEIs) in these traits, which have the potential to compromise mate choice for genetic benefits. Our experimental manipulation of the social environment revealed plastic expression of both traits; males exposed to a rival male during the sensitive period of adult sexual maturation exhibited shorter mating latencies and longer copulation durations than those who matured in isolation. However, we found no evidence for GEIs, and no significant additive genetic variation underlying these traits in either environment. These results undermine the notion that the evolution of female choice rests on covariance between female preference and male displays, an expectation that underpins indirect benefit models such as the good genes and sexy sons hypotheses. However, our results may also indicate depletion of genetic variance in these traits in the natural population studied, thus supporting the expectation that traits closely aligned with reproductive fitness can exhibit low levels of additive genetic variance.

Assessing genetic variation to improve stem straightness in Eucalyptus globulus

Stem straightness is an important trait for growers and processors of Eucalyptus globulus logs for solid-wood products. The aims of the study were to determine the extent of genetic variation in stem deviation from straightness in E. globulus and assess the utility of a six-point subjective scoring method as a selection criterion for stem straightness. Two E. globulus progeny trials, grown under solid-wood product regimes, were studied. At age 9 years (post-thinning), stem straightness was measured using both image analysis and a six-point subjective scale. Diameter at breast height (DBH; 1.3 m) was measured at both age 5 (pre-thinning) and age 9 years. Significant additive genetic variation was observed. Strong, positive and significant additive genetic correlations were observed between the stem straightness assessment methods and between DBH at ages 5 and 9 years. Significant positive genetic correlations were shown between subjectively scored stem straightness and DBH at both ages 5 and 9 years. The six-point subjective scoring method is a cost-effective selection criterion for stem straightness in E. globulus. The image measurement technique may be applied where objective estimates of stem straightness are required, for training purposes and to verify subjective scores.

Pathogen-induced hatching and population-specific life-history response to waterborne cues in brown trout (Salmo trutta)

Hatching is an important niche shift, and embryos in a wide range of taxa can either accelerate or delay this life-history switch in order to avoid stage-specific risks. Such behavior can occur in response to stress itself and to chemical cues that allow anticipation of stress. We studied the genetic organization of this phenotypic plasticity and tested whether there are differences among populations and across environments in order to learn more about the evolutionary potential of stress-induced hatching. As a study species, we chose the brown trout (Salmo trutta; Salmonidae). Gametes were collected from five natural populations (within one river network) and used for full-factorial in vitro fertilizations. The resulting embryos were either directly infected with Pseudomonas fluorescens or were exposed to waterborne cues from P. fluorescens-infected conspecifics. We found that direct inoculation with P. fluorescens increased embryonic mortality and induced hatching in all host populations. Exposure to waterborne cues revealed population-specific responses. We found significant additive genetic variation for hatching time, and genetic variation in trait plasticity. In conclusion, hatching is induced in response to infection and can be affected by waterborne cues of infection, but populations and families differ in their reaction to the latter.

Personality pace-of-life hypothesis: testing genetic associations among personality and life history

Over the last few years, animal personality researchers have called for integrative approaches to study behavioral, immunological, and life-historical traits. This is because life history and personality have become implied as part of integrative “pace-of-life” syndromes. Immune defense is one of the traits that have been suggested to associate with personality traits, such as boldness, mainly because behavioral types may differ in parasite encounter rates. Here, we quantified the narrow-sense heritabilities (h 2 = VA/(VA + VR)) and genetic (rA) and phenotypic (rP) correlations between 2 measures of behavior (overall boldness and defreezing, i.e., recovery from disturbance-induced immobility), 1 measure of immune function (encapsulation response), and 2 life-history traits (body mass and maturation time) using the western stutter-trilling cricket, Gryllus integer. All nonbehavioral traits showed strong heritabilities (range: encapsulation response and life-history: h 2 = 0.42–0.84), whereas behaviors were only marginally heritable (h 2 = 0.06–0.11). Boldness and encapsulation were positively associated, and defreezing and body mass were negatively associated phenotypically, whereas only defreezing was positively genetically correlated with encapsulation. However, the lack of significant additive genetic variation in defreezing suggests that the genetic correlation may be an artifact and therefore, that there were only environmentally induced phenotypic correlations between behaviors and other measured traits. Life-history traits and encapsulation were positively phenotypically associated and these associations were mostly genetically underpinned. Overall, our results support the hypothesis that life history is structured in an evolutionarily significant “pace-of-life” syndrome, but suggest that behavior does not intrinsically integrate with such life-history variation.

Cuticular hydrocarbons as a basis for chemosensory self‐referencing in crickets: a potentially universal mechanism facilitating polyandry in insects

Females of many species obtain benefits by mating polyandrously, and often prefer novel males over previous mates. However, how do females recognise previous mates, particularly in the face of cognitive constraints? Female crickets appear to have evolved a simple but effective solution: females imbue males with their own cuticular hydrocarbons (CHCs) at mating and utilise chemosensory self-referencing to recognise recent mates. Female CHC profiles exhibited significant additive genetic variation, demonstrating that genetically unique chemical cues are available to support chemosensory self-referencing. CHC profiles of males became more similar to those of females after mating, indicating physical transfer of CHCs between individuals during copulation. Experimental perfuming of males with female CHCs resulted in a female aversion to males bearing chemical cues similar to their own. Chemosensory self-referencing, therefore, could be a widespread mechanism by which females increase the diversity of their mating partners.

Genetic variation in the susceptibility of Eucalyptus globulus to drought damage

Genetic variation in drought damage in Eucalyptus globulus was studied in a sublined trial series across four neighbouring sites in Western Australia linked by ten common families. The trials included approximately 400 open-pollinated families, encompassing 51 native stand collection localities and 19 subraces from throughout the geographic range of the species. Data were analysed using mixed models, with spatial analysis used to better identify genetic effects. Significant subrace differences in drought damage were detected, with both broad-scale, regional and localised clines evident. The quantitative genetic differentiation between subraces as measured by QST (0.39 ± 0.091) was significantly greater than the FST for neutral marker expectations and consistent with diversifying selection shaping the patterns of subrace divergence in drought susceptibility. This conclusion is supported by the significant association of subrace drought susceptibility with bioclimatic parameters, particularly those associated with temperature seasonality. Less drought damage was observed in subraces originating from areas with more temperature seasonality, but also less radiation and rainfall seasonality, less winter rainfall, higher radiation and higher temperatures in the warmest month. Significant additive genetic variation in drought damage was detected within subraces, with narrow-sense heritabilities ranging from 0.14 to 0.20. We argue that spatial genetic variation in drought susceptibility of E. globulus has been shaped by natural selection acting at multiple scales and discuss opportunities for exploiting this genetic variation in breeding and deployment programs.

Genetic and nutritional effects on male traits and reproductive performance in Tribolium flour beetles

In Tribolium flour beetles and other organisms, individuals migrate between heterogeneous environments where they often encounter markedly different nutritional conditions. Under these circumstances, theory suggests that genotype-by-environment interactions (GEI) may be important in facilitating adaptation to new environments and maintaining genetic variation for male traits subject to directional selection. Here, we used a nested half-sib breeding design with Tribolium castaneum to partition the separate and joint effects of male genotype and nutritional environment on phenotypic variation in a comprehensive suite of life-history traits, reproductive performance measures across three sequential sexual selection episodes, and fitness. When male genotypes were tested across three nutritional environments, considerable phenotypic plasticity was found for male mating and insemination success, longevity and traits related to larval development. Our results also revealed significant additive genetic variation for male mating rate, sperm offence ability (P(2)), longevity and total fitness and for several traits reflecting both larval and adult resource use. In addition, we found evidence supporting GEI for sperm defence ability (P(1)), adult longevity and larval development; thus, no single male genotype outperforms others in every nutritional environment. These results provide insight into the potential roles of phenotypic plasticity and GEI in facilitating Tribolium adaptation to new environments in ecological and evolutionary time.

Genetic variation in resistance to infectious pancreatic necrosis in rainbow trout (Oncorhynchus mykiss) after a challenge test

In this study, we wanted to evaluate genetic variation in resistance to infectious pancreatic necrosis (IPN) in a breeding population of rainbow trout. Two hundred families were challenged at a commercial test station, using methods that were previously developed for testing resistance to IPN in Atlantic salmon. Thirty-five days after the challenge the accumulated mortality was 26%. The results show that resistance to IPN is moderately heritable in the tested population (h2=0.30). The genetic correlation between IPN resistance and body weight was found to be low and non-significant. The significant additive genetic variation found in IPN resistance after a controlled challenge test gives promise for successful breeding for increased resistance to IPN in rainbow trout.

Heritable victimization and the benefits of agonistic relationships.

Here, we present estimates of heritability and selection on network traits in a single population, allowing us to address the evolutionary potential of social behavior and the poorly understood link between sociality and fitness. To evolve, sociality must have some heritable basis, yet the heritability of social relationships is largely unknown. Recent advances in both social network analyses and quantitative genetics allow us to quantify attributes of social relationships and estimate their heritability in free-living populations. Our analyses addressed a variety of measures (in-degree, out-degree, attractiveness, expansiveness, embeddedness, and betweenness), and we hypothesized that traits reflecting relationships controlled by an individual (i.e., those that the individual initiated or were directly involved in) would be more heritable than those based largely on the behavior of conspecifics. Identifying patterns of heritability and selection among related traits may provide insight into which types of relationships are important in animal societies. As expected, we found that variation in indirect measures was largely explained by nongenetic variation. Yet, surprisingly, traits capturing initiated interactions do not possess significant additive genetic variation, whereas measures of received interactions are heritable. Measures describing initiated aggression and position in an agonistic network are under selection (0.3 < |S| < 0.4), although advantageous trait values are not inherited by offspring. It appears that agonistic relationships positively influence fitness and seemingly costly or harmful ties may, in fact, be beneficial. Our study highlights the importance of studying agonistic as well as affiliative relationships to understand fully the connections between sociality and fitness.

VARIANCES AND COVARIANCES OF PHENOLOGICAL TRAITS IN A WILD MAMMAL POPULATION

In a seasonal environment, there are multiple aspects of timing, or phenology, that contribute to an individual's fitness. Several studies have shown a genetic basis to variation between individuals in breeding time, but we know little about the heritability of other phenological traits in wild populations. Furthermore, the presence of genetic correlations between phenological variables could act to constrain or promote any response to selection, but less is known of the multivariate genetic relationships underlying phenological traits in the wild. Here, we use data from a wild population of red deer on the Isle of Rum, Scotland, to investigate covariances between eight phenological traits. Variation was characterized at the level of the phenotype, genotype, and year, and traits measured in different sexes enabled us to test for cross-sex genetic correlations. Phenotypic correlations were broadly strong and positive, as were correlations between traits expressed in the same year. We found evidence of significant additive genetic variation in five of the eight phenological traits studied. However there was little evidence of genetic correlations between traits, implying that much of the observed phenotypic correlation was environmentally induced. Our results suggest that different phenological traits may be free to move along independent evolutionary trajectories.

Heritability of morphology in brook trout with variable life histories.

Distinct morphological variation is often associated with variation in life histories within and among populations of both plants and animals. In this study, we examined the heritability of morphology in three hatchery strains of brook trout (Salvelinus fontinalis), which were historically or are currently used for stocking and supplementation of both migratory and resident ecotypes in the upper Great Lakes region. In a common garden experiment, significant variation in body morphology was observed within and across populations sampled at three time periods. The most notable differences among strains were differences in dorso-ventral body depth and the shape of the caudal peduncle, with some differences in the anterior-posterior placement of the dorsal and ventral fins. Variation with and among 70 half-sib families indicates that heritabilities of morphology and body size were significant at most developmental time points both within and across strains. Heritabilities for morphological characters within strains ranged from 0 to 0.95 across time points. Significant within-strain heritabilities for length ranged from 0 to 0.93 across time points and for weight ranged from 0 to 0.88. Significant additive genetic variation exists within and across hatchery brook trout strains for morphology and size, indicating that these traits are capable of responding to natural or artificial selection.

Stiffness and checking of Eucalyptus nitens sawn boards: genetic variation and potential for genetic improvement

A trial was undertaken to assess the extent to which variation in sawn-board quality traits of plantation-grown Eucalyptus nitens is under genetic control and amenable to genetic improvement. Five hundred and sixty trees from 129 families and three central Victorian races were sampled from an open-pollinated progeny trial in Tasmania, Australia. Acoustic wave velocity (AWV) was assessed on standing-trees and sawlogs. Wedges from disks extracted from sawlogs were assessed for basic density and checking. Processed boards from 496 of the trees were assessed for board stiffness (static modulus of elasticity, MOE), and internal and surface checking. Genetic differences among races were significant for AWV and MOE traits. The Southern race had the highest mean values for these traits. Significant additive genetic variation within races was observed in all traits, demonstrating that the quality of plantation-grown E. nitens boards could be improved through breeding. Estimated narrow-sense heritabilities were 0.85 for standing-tree AWV, 0.71 for log AWV, 0.37 for board MOE, and ranged from 0.20 to 0.52 for checking traits. A strongly positive genetic correlation (rg = 1.05) was observed between standing-tree AWV and board MOE, indicating that AWV could be used as a selection trait to improve E. nitens board stiffness. The genetic correlation between basic density and board MOE was also positive (rg = 0.62). However, a significant and adverse genetic correlation (rg = 0.61) was identified between basic density and surface check length. Wood stiffness and checking traits were more-or-less genetically independent, and genetic correlations between surface and internal checking were positive but only moderate (rg = 0.48-0.52).

Genetic determination of fatty acid composition in Spanish Churra sheep milk

The objective of this study was to estimate the genetic variation of ovine milk fatty acid (FA) composition. We collected 4,100 milk samples in 14 herds from 976 Churra ewes sired mostly by 15 AI rams and analyzed them by gas–liquid chromatography for milk fatty acid composition. The studied traits were 12 individual FA contents (proportion in relation to the total amount of FA), 3 groups of fatty acids [saturated fatty acids (SFA), monounsaturated FA (MUFA), and polyunsaturated FA (PUFA)], and 2 FA ratios (n-6:n-3 and C18:2 cis-9,trans-11:C18:1 trans-11). In addition, percentages of fat and protein and daily milk yield were studied. For the analysis, repeatability animal models were implemented using Bayesian methods. In an initial step, univariate methods were conducted to test the hypothesis of the traits showing additive genetic determination. Deviance information criterion and Bayes factor were employed as model choice criteria. All the studied SFA showed additive genetic variance, but the estimated heritabilities were low. Among unsaturated FA (UFA), only C18:1 trans-11 and C18:2 cis-9,cis-12 showed additive genetic variation, their estimated heritabilities being [marginal posterior mean (marginal posterior SD)] 0.02(0.01) and 0.11(0.04), respectively. For the FA groups, only PUFA showed significant additive genetic variation. None of the studied ratios of FA showed additive genetic variation. In second multitrait analyses, genetic correlations between individual FA and production traits, and between groups of FA and ratios of FA and production traits, were investigated. Positive genetic correlations were estimated among medium-chain SFA, ranging from 0 to 0.85, but this parameter was close to zero between long-chain SFA (C16:0 and C18:0). Between long- and medium-chain SFA, estimated genetic correlations were negative, around −0.6. Among those UFA showing significant additive genetic variance, genetic correlations were close to zero. The estimated genetic correlations among all the investigated FA, milk yield, and fat and protein percentages were not different from zero. Our results suggest that low additive genetic variation is involved in the determination of the FA composition of milk fat in Churra sheep under current production conditions, which results in low values of heritabilities.

The Effects of Drying Temperature and Method of Assessment on the Expression of Genetic Variation in Gross Shrinkage of Eucalyptus globulus Wood Samples

Abstract Genetic variation in wood-sample gross shrinkage and basic density was examined in a Eucalyptus globulus base population trial growing in Tasmania, Australia. Gross shrinkage, which includes all components of shrinkage including collapse, was assessed in four ways (calliper- and visually-assessed tangential shrinkage, volumetric shrinkage and radial shrinkage) on samples dried at three temperatures (22°C, 60°C and 105°C). Significant differences between subraces were observed using all measures of gross shrinkage for two or more of the three drying treatments. Furthermore, significant additive genetic variation within subraces was observed in calliper- and visually-assessed gross shrinkage under two or more of the drying treatments, with narrowsense heritabilities greater than or equal to 0.35. There was no obvious trend in heritabilities or coefficients of additive genetic variation with drying temperature. Under the 105°C drying treatment, subrace correlations among calliper-, visually- and volume-assessed gross shrinkage were positive and very strong (≥ 0.97), while these measures were less strongly correlated with radial gross shrinkage at the subrace level (≤ 0.77). Withinsubrace genetic correlations among the first three measures were also strongly positive (≥ 0.95). These high genetic correlations suggest that different drying regimes and the calliper, visual and volume methods of assessment could be used interchangeably to select for reduced tangential gross shrinkage. Estimated subrace and genetic correlations between basic density and measures of gross shrinkage were universally negative (i.e. favourable), although not all were significantly different to zero.

Genetic stability of wood density and diameter in Pinus radiata D. Don plantation estate across Australia

Genetic variation for wood quality traits and diameter growth for radiata pine (Pinus radiata D. Don) at age 20/21 years was estimated from eight trials in Australia. The traits studied were wood density, acoustic time-of-flight (an indirect measure of stiffness) and diameter at breast height (DBH). Wood density and DBH exhibited significant additive genetic variation whereas non-additive effects were not significantly different from zero. Time of flight was also not significantly different from zero for both additive and non-additive effects, respectively. Average single-site heritability estimates (±SE) for wood density and DBH were 0.38 ± 0.10 and 0.16 ± 0.08, respectively. Pooled-site heritability estimates for wood density and DBH were 0.38 ± 0.10 and 0.08 ± 0.10, respectively. For density, there was little evidence of genotype-by-environment interaction (GEI) across the eight trials at the additive level (type B additive genetic correlation; rBADD = 0.73 ± 0.08) and type B genetic correlation for full-sib families (rBFS = 0.64 ± 0.08). In contrast, the type B additive genetic correlation for DBH was lower, (rBADD = 0.51 ± 0.14), suggesting evidence of GEI. However, type B genetic correlation for full-sib families was moderate (0.63 ± 0.11) for DBH, suggesting that there may be some stable full-sib families. On the basis of the results of this study, GEI should be considered in order to optimise deployment of improved germplasm in Australia.

Genetic variation and constraints on the evolution of defense against spittlebug (Philaenus spumarius) herbivory in Mimulus guttatus

Plants mediate carbon into most ecosystems and are thus under persistent attack by diverse enemies. The evolution of defense against such assaults will depend on the availability of genetic variation, as well as the costs and constraints on defense. We estimated the magnitude of genetic variation for defense against spittlebug (Philaenus spumarius) herbivory in Mimulus guttatus using a diallel cross-grown in a greenhouse. Except for flowering time, additive genetic variation for the plant traits we measured was negligible, regardless of herbivory environment. In contrast, nonadditive genetic variation contributed significantly to all plant traits measured. We found significant additive genetic variation among plants for biomass of adult spittlebugs, suggesting heritability for resistance to herbivory. The other putative resistance trait measured, spittlebug maturation time, was not significantly heritable. We found no evidence for significant genetic variation for tolerance to herbivory except for a small non-nuclear paternal contribution to tolerance for flower number. Additive genetic correlations indicated that more resistant plant genotypes (in terms of adult spittlebug biomass) were also smaller in the absence of spittlebugs, suggesting a potential cost of resistance to herbivory. We found no other significant genetic correlations indicating a cost of defense, nor did we find evidence for a tradeoff between resistance and tolerance to herbivory. Overall, these results suggest the future adaptive evolution of tolerance to spittlebugs in this population will be limited primarily by available genetic variation, whereas the future evolution of antibiosis resistance may be constrained by allocation costs of resistance.