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Cyclically parthenogenetic animals such as aphids are able alternating sexual and asexual reproduction during its life cycle, and represent good models for studying short-term evolutionary consequences of sex. In aphids, different morphs, whether sexual or asexual, winged or wingless, are produced in response to specific environmental cues. The production of these morphs could imply a differential energy investment between the two reproductive phases (i.e., sexual and asexual), which can also be interpreted in terms of changes in genetic variation and/or trade-offs between the associated traits. In this study we compared the G-matrices of energy metabolism, life-history traits and morph production in 10 clonal lineages (genotypes) of the pea aphid, Acyrthosiphon pisum, during both sexual and asexual phases. The heritabilities (broad-sense) were significant for almost all traits in both phases; however the only significant genetic correlation we found was a positive correlation between resting metabolic rate and production of winged parthenogenetic females during the asexual phase. These results suggest the pea aphid shows some lineage specialization in terms of energy costs, but a higher specialization in the production of the different morphs (e.g., winged parthenogenetic females). Moreover, the production of winged females during the asexual phase appears to be more costly than wingless females. Finally, the structures of genetic variance-covariance matrices differed between both phases. These differences were mainly due to the correlation between resting metabolic rate and winged parthenogenetic females in the asexual phase. This structural difference would be indicating that energy allocation rules changes between phases, emphasizing the dispersion role of asexual morphs., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Wing dimorphism has been proposed as a strategy to face trade-offs between flight capability and fecundity. In aphids, individuals with functional wings have slower development and lower fecundity compared with wingless individuals. However, differential maintenance costs between winged and wingless aphids have not been deeply investigated. In the current study, we studied the combined effect of wing dimorphism with the effects of aphid genotypes and of wheat hosts having different levels of chemical defences (hydroxamic acids, Hx) on adult body mass and standard metabolic rates (SMR) of winged and wingless morphs of the grain aphid, Sitobion avenae. We found that wingless aphids had higher body mass than winged aphids and that body mass also increased towards host with high Hx levels. Furthermore, winged aphids showed a plastic SMR in terms of Hx levels, whereas wingless aphids displayed a rigid reaction norm (significant interaction between morph condition and wheat host). These findings suggest that winged aphids have reduced adult size compared to wingless aphids, likely due to costs associated to the development of flight structure in early-life stages. These costs contrast with the absence of detectable metabolic costs related to fuelling and maintenance of the flight apparatus in adults., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

Induced defences are a typical case of phenotypic plasticity, involving benefits for 'plastic' phenotypes under environments with variable degree of stress. Defence induction, in turn, could be energetically expensive incurring costs on growth and reproduction. In this study, we investigated the genetic variation and induction of detoxification enzymes mediated by wheat chemical defences (hydroxamic acids; Hx), and their metabolic and fitness costs using five multilocus genotypes of the grain aphid (Sitobion avenae). Cytochrome P450 monooxygenases and glutathione S-transferases activities were seen to increase with Hx levels, whereas esterases activity and standard metabolic rate increased in wheat hosts with low Hx levels. Additionally, the intrinsic rate of increase (a fitness proxy) increased in highly defended hosts. However, we did not find significant genetic variation or genotype-host interaction for any studied trait. Therefore, aphids feeding on host plants with elevated chemical defences appeared to reduce their detoxification costs and to increase their reproductive performance, which we interpret as a novel adaptation to defended plants. In brief, this study supports the notion that aphids perform better on highly defended host plants, probably related to the selective pressures during the colonization of New World agroecosystems, characterized by highly defended host plants., (© 2010 The Authors. Journal Compilation © 2010 European Society For Evolutionary Biology.)

'Superclones' are predominant and time-persistent genotypes, exhibiting constant fitness across different environments. However, causes of this ecological success are still unknown. Therefore, we studied the physiological mechanisms that could explain this success, evaluating the effects of wheat chemical defences on detoxification enzymes [cytochrome P450 monooxygenases (P450), glutathione S-transferases (GST), esterases (EST)], standard metabolic rate (SMR), and fitness-related traits [adult body mass and intrinsic rate of increase (r(m))] of two 'superclones' (Sa1 and Sa2) of the grain aphid, Sitobion avenae. Additionally, we compared 'superclones' with a less-frequent genotype (Sa46). Genotypes were reared on three wheat cultivars with different levels of hydroxamic acids (Hx; wheat chemical defences). Detoxification enzymes and SMR did not differ between wheat hosts. However, GST and EST were different between 'superclones' and Sa46, while Sa1 showed a higher SMR than Sa2 or Sa46 (p=0.03). Differences between genotypes were found for r(m), which was higher for Sa1 than for Sa2 or Sa46. For all cases, genotype-host interactions were non-significant, except for aphid body mass. In conclusion, 'superclones' exhibit a broad host range, flat energetic costs for non-induced detoxification enzymes, and low variation in their reproductive performance on different defended hosts. However, physiological specialization of 'superclones' that could explain their ecological success was not evident in this study.

Life-history theory postulates that evolution is constrained by trade-offs (i.e., negative genetic correlations) among traits that contribute to fitness. However, in organisms with complex life cycles, trade-offs may drastically differ between phases, putatively leading to different evolutionary trajectories. Here, we tested this possibility by examining changes in life-history traits in an aphid species that alternates asexual and sexual reproduction in its life cycle. The quantitative genetics of reproductive and dispersal traits was studied in 23 lineages (genotypes) of the bird cherry-oat aphid Rhopalosiphum padi, during both the sexual and asexual phases, which were induced experimentally under specific environmental conditions. We found large and significant heritabilities (broad-sense) for all traits and several negative genetic correlations between traits (trade-offs), which are related to reproduction (i.e., numbers of the various sexual or asexual morphs) or dispersal (i.e., numbers of winged or wingless morphs). These results suggest that R. padi exhibits lineage specialization both in reproductive and dispersal strategies. In addition, we found important differences in the structure of genetic variance-covariance matrices (G) between phases. These differences were due to two large, negative genetic correlations detected during the asexual phase only: (1) between fecundity and age at maturity and (2) between the production of wingless and winged parthenogenetic females. We propose that this differential expression in genetic architecture results from a reallocation scheme during the asexual phase, when sexual morphs are not produced. We also found significant G x E interaction and nonsignificant genetic correlations across phases, indicating that genotypes could respond independently to selection in each phase. Our results reveal a rather unique situation in which the same population and even the same genotypes express different genetic (co)variation under different environmental conditions, driven by optimal resource allocation criteria.

Herbivorous insects have developed mechanisms to cope with plant barriers, including enzymatic systems to detoxify plant allelochemicals. Detoxification systems may be induced when insects are feeding on plants with increasing levels of allelochemicals. Increases in enzymatic activity have been related to energetic costs, and therefore less energy may be allocated to fitness-related traits. In this study, we explored the induction and energetic costs of detoxifying hydroxamic acids (Hx; a wheat allelochemical) in the grain aphid, Sitobion avenae. Aphids were reared on three wheat cultivars with different levels of Hx (0.26+/-0.08, 2.09+/-0.6 and 5.91+/-1.18 mmol kg(-1) fresh mass). We performed a nested ANOVA to test the effect of Hx (main factor) and intrahost variation (nested factor) on body mass, standard metabolic rate (SMR) and the enzymatic activity of cytochrome P450s monooxygenases (P450s), glutathione S-transferases (GSTs) and esterases (ESTs). We found non-significant effects of Hx levels (P>0.5 for all tests), but there was significant intrahost variation (P<0.05 for all tests). In addition, we found a negative correlation between SMR and ESTs (P=0.003) and no correlation between SMR and GSTs or P450s (P=n.s after a Bonferroni correction). Multiple regression between SMR (dependent variable) and enzymatic activities (predictor variables) was significant (P=0.007), but detoxification enzymes only explained about 5% of the variation of SMR. Finally, we found a non-significant path coefficient between ;metabolism' and ;detoxifying capacity' (P>0.05). These results suggest that increased enzymatic activities do not entail increased metabolic rate. Therefore, low energetic costs in aphids would facilitate the use of different hosts and promote a wider ecological niche.

One of the most important factors that determine the evolutionary trajectory of a suite of traits in a population is the structure of the genetic variance-covariance matrix (G). We studied the cyclically parthenogenetic aphid Rhopalosiphum padi, whose populations exhibit two types of reproductive lineages respectively specialized in sexuality (that is, cyclically parthenogenetic lineages) and in asexuality (that is, obligate parthenogenetic lineages). We compared the quantitative genetics of life histories in these two lineage types. Our results suggest that both, the elements and the whole structure of the resulting G matrices differ in the very short term, between lineage types. This would involve the evolution toward different evolutionary optima in the same population, depending on whether sexual or asexual lineages predominate. Since sexual and asexual lineages vary seasonally in their abundance, a fluctuating selective regime has been proposed for this species, which would contribute to the maintenance of the reproductive polymorphism that these populations exhibit.