Influences of elevated CO2 and pest damage on the allocation of plant defense compounds in Bt‐transgenic cotton and enzymatic activity of cotton aphid

Abstract Plant allocation to defensive compounds by elevated CO₂‐grown non‐transgenic and transgenic Bt cotton in response to infestation by cotton aphid, Aphis gossypii (Glover) in open‐top chambers under elevated CO₂ were studied. The results showed that significantly lower foliar nitrogen concentration and Bt toxin protein occurred in transgenic Bt cotton with and without cotton aphid infestation under elevated CO_2. However, significantly higher carbon/nitrogen ratio, condensed tannin and gossypol were observed in transgenic Bt cotton “GK‐12” and non‐transgenic Bt cotton ‘Simian‐3’ under elevated CO_2. The CO₂ level and cotton variety significantly influenced the foliar nitrogen, condensed tannin and gossypol concentrations in the plant leaves after feeding by A. gossypii. The interaction between CO₂ level × infestation time (24 h, 48 h and 72 h) showed a significant increase in cotton condensed tannin concentrations, while the interaction between CO₂ level × cotton variety significantly decreased the true choline esterase (TChE) concentration in the body of A. gossypi. This study exemplified the complexities of predicting how transgenic and non‐transgenic plants will allocate defensive compounds in response to herbivorous insects under differing climatic conditions. Plant defensive compound allocation patterns and aphid enzyme changes observed in this study appear to be broadly applicable across a range of plant and herbivorous insect interactions as CO₂ atmosphere rises.     

Effects of elevated CO2 on the interspecific competition between two sympatric species of Aphis gossypii and Bemisia tabaci fed on transgenic Bt cotton

Abstract Effects of elevated CO₂ (twice ambient vs. ambient) and Bt Cry1Ac transgene (Bt cotton cv. 33^B vs. its nontransgenic parental line cv. DP5415) on the interspecific competition between two ecologically similar species of cotton aphid Aphis gossypii and whitefly biotype‐Q Bemisia tabaci were studied in open‐top chambers. The results indicated that elevated CO₂ and Bt cotton both affected the population abundances of A. gossypii and biotype‐Q B. tabaci when introduced solely (i.e., without interspecific competition) or two species coexisted (i.e., with interspecific competition). Compared with ambient CO₂, elevated CO₂ increased the population abundances of A. gossypii and biotype‐Q B. tabaci as fed on Bt and nontransgenic cotton on 45 (i.e., seedling stage) and 60 (i.e., flowering stage) days after planting (DAP), but only significantly enhanced aphid abundance without interspecific competition on the 45‐DAP nontransgenic cotton and 60‐DAP Bt cotton, and significantly increased whitefly abundance with interspecific competition on the 45‐DAP Bt cotton and 60‐DAP nontransgenic cotton. In addition, compared with nontransgenic cotton at elevated CO₂, Bt cotton significantly reduced biotype‐Q B. tabaci abundances without and with interspecific competition during seedling and flowering stage, while only significantly decreasing A. gossypii abundances without interspecific competition during the seedling stage. When the two insect species coexisted, the proportions of biotype‐Q B. tabaci were significantly higher than those of A. gossypii on Bt and nontransgenic cotton at the same CO₂ levels, and elevated CO₂ only significantly increased the percentages of biotype‐Q B. tabaci and significantly reduced the proportions of A. gossypii on seedling and flowering nontransgenic cotton. Therefore, the effects of elevated CO₂ were favorable for biotype‐Q B. tabaci to out‐compete A. gossypii under the predicted global climate change.     

Response of amino acid changes in Aphis gossypii (Glover) to elevated CO2 levels

Effects of elevated CO₂ levels on the amino acid constituents of cotton aphid, Aphis gossypii (Glover), fed on transgenic Bacillus thuringiensis (Berliner) (Bt) cotton [Cryl A(c)], grown in ambient and double‐ambient CO₂ levels in closed‐dynamics CO₂ chambers, were investigated. Lower amounts of amino acids were found in cotton phloem under elevated CO₂ than under ambient CO₂ levels. However, higher amounts of free amino acids were found in A. gossypii fed on elevated CO₂‐grown cotton than those fed ambient CO₂‐grown cotton, and the contents of amino acids in honeydew were not significantly affected by elevated CO₂ levels. A larger amount of honeydew was produced by cotton aphids feeding on leaves under elevated CO₂ treatment than those feeding on leaves under ambient CO₂ treatment, which indicates that A. gossypii ingests more cotton phloem because of the higher C:N ratio of cotton phloem under elevated CO₂ levels. Moreover, the amino acid composition was similar in bodies of aphids ingesting leaves under both CO₂ treatments, except for two alkaline amino acids, lysine and arginine. This suggests that the nutritional constitution of the phloem sap was important for A. gossypii. Our data suggest that more phloem sap will be ingested by A. gossypii to satisfy its nutritional requirement and balance the break‐even point of amino acid in elevated CO₂. Larger amounts of honeydew produced by A. gossypii under elevated CO₂ will reduce the photosynthesis and result in the occurrence of some Entomophthora spp.     

How does atmospheric elevated CO2 affect crop pests and their natural enemies? Case histories from China

Abstract Global atmospheric CO₂ concentrations have risen rapidly since the Industrial Revolution and are considered as a primary factor in climate change. The effects of elevated CO₂ on herbivore insects were found to be primarily through the CO₂‐induced changes occurring in their host plants, which then possibly affect the intensity and frequency of pest outbreaks on crops. This paper reviews several ongoing research models using primary pests of crops (cotton bollworm, whitefly, aphids) and their natural enemies (ladybeetles, parasitoids) in China to examine insect responses to elevated CO₂. It is generally indicated that elevated CO₂ prolonged the development of cotton bollworm, Helicoverpa armigera, a chewing insect, by decreasing the foliar nitrogen of host plants. In contrast, the phloem‐sucking aphid and whitefly insects had species‐specific responses to elevated CO₂ because of complex interactions that occur in the phloem sieve elements of plants. Some aphid species, such as cotton aphid, Aphis gossypii and wheat aphid, Sitobion avenae, were considered to represent the only feeding guild to respond positively to elevated CO₂ conditions. Although whitefly, Bemisia tabaci, a major vector of Tomato yellow leaf curl virus, had neutral response to elevated CO₂, the plants became less vulnerable to the virus infection under elevated CO₂. The predator and parasitoid response to elevated CO₂ were frequently idiosyncratic. These documents from Chinese scientists suggested that elevated CO₂ initially affects the crop plant and then cascades to a higher trophic level through the food chain to encompass herbivores (pests), their natural enemies, pathogens and underground nematodes, which disrupt the natural balance observed previously in agricultural ecosystems.     

Aphid and host‐plant genotype × genotype interactions under elevated CO2

1. Elevated CO₂ can alter plant physiology and morphology, and these changes are expected to impact diet quality for insect herbivores. While the plastic responses of insect herbivores have been well studied, less is known about the propensity of insects to adapt to such changes. Genetic variation in insect responses to elevated CO₂ and genetic interactions between insects and their host plants may exist and provide the necessary raw material for adaptation. 2. We used clonal lines of Rhopalosiphum padi (L.) aphids to examine genotype‐specific responses to elevated CO₂. We used the host plant Schedonorus arundinaceus (tall fescue; Schreb), which is capable of asexual reproduction, to investigate host plant genotype‐specific effects and possible host plant‐by‐insect genotype interactions. The abundance and density of three R. padi genotypes on three tall fescue genotypes under three concentrations of CO₂ (ambient, 700, and 1000 ppm) in a controlled greenhouse environment were examined. 3. Aphid abundance decreased in the 700 ppm CO₂ concentration, but increased in the 1000 ppm concentration relative to ambient. The effect of CO₂ on aphid density was dependent on host plant genotype; the density of aphids in high CO₂ decreased for two plant genotypes but was unchanged in one. No interaction between aphid genotype and elevated CO₂ was found, nor did we find significant genotype‐by‐genotype interactions. 4. This study suggests that the density of R. padi aphids feeding on tall fescue may decrease under elevated CO₂ for some plant genotypes. The likely impact of genotype‐specific responses on future changes in the genetic structure of plant and insect populations is discussed.     

Aphid Honeydew Quality as a Food Source for Parasitoids Is Maintained in Bt Cotton

Bt-transgenic cotton has proven to be highly efficient in controlling key lepidopteran pests. One concern with the deployment of Bt cotton varieties is the potential proliferation of non-target pests. We previously showed that Bt cotton contained lower concentrations of insecticidal terpenoids as a result of reduced caterpillar damage, which benefited the aphid Aphis gossypii. It is thus important that non-target herbivores are under biological control in Bt cotton fields. The induction or lack of induction of terpenoids could also influence the quality of aphid honeydew, an important food source for beneficial insects. We therefore screened A. gossypii honeydew for cotton terpenoids, that are induced by caterpillars but not the aphids. We then tested the influence of induced insect-resistance of cotton on honeydew nutritional quality for the aphid parasitoid Lysiphlebus testaceipes and the whitefly parasitoid Eretmocerus eremicus. We detected the cotton terpenoids gossypol and hemigossypolone in A. gossypii honeydew. Although a feeding assay demonstrated that gossypol reduced the longevity of both parasitoid species in a non-linear, dose-dependent manner, the honeydew was capable of sustaining parasitoid longevity and reproduction. The level of caterpillar damage to Bt and non-Bt cotton had no impact on the quality of honeydew for the parasitoids.These results indicate that the nutritional quality of honeydew is maintained in Bt cotton and is not influenced by induced insect resistance.     

Transgenic Bacillus thuringiensis (Bt) cotton (Gossypium hirsutum) allomone response to cotton aphid, Aphis gossypii, in a closed-dynamics CO(2) chamber (CDCC)

Allocation of allomones of transgenic Bacillus thuringiensis Gossypium hirsutum (Bt cotton) (cv. GK-12) and non-Bt-transgenic cotton (cv. Simian-3) grown in elevated CO₂ in response to infestation by cotton aphid, Aphis gossypii Glover, was studied in a closed-dynamics CO₂ chamber. Significant increases in foliar condensed tannin and carbon/nitrogen ratio for GK-12 and Simian-3 were observed in elevated CO₂ relative to ambient CO_2, as partially supported by the carbon nutrient balance hypothesis, owing to limiting nitrogen and excess carbon in cotton plants in response to elevated CO₂. The CO₂ level significantly influenced the foliar nutrients and allomones in the cotton plants. Aphid infestation significantly affected foliar nitrogen and allomone compounds in the cotton plants. Allomone allocation patterns in transgenic Bt cotton infested by A. gossypii may have broader implications across a range of plant and herbivorous insects as CO₂ continues to rise. Gang Wu and Fa Jun Chen contributed equally to this work.     

Impacts of elevated CO2 on expression of plant defensive compounds in Bt‐transgenic cotton in response to infestation by cotton bollworm

• 1 The allocation of defensive compounds of transgenic Bt (cv. GK‐12) and nontransgenic cotton (cv. Simian‐3) grown in elevated CO₂ in response to infestation by cotton bollworm Helicoverpa armigera (Hübner) was studied in closed‐dynamics CO₂ chambers.
• 2 A significant reduction in foliar nitrogen content and Bt toxin protein occurred when transgenic Bt cotton grew under elevated CO₂. A significantly higher carbon/nitrogen ratio as well as condensed tannin and gossypol contents was observed for transgenic Bt (cv. GK‐12) and nontransgenic cotton in elevated CO₂, in partial support of the carbon nutrient balance hypothesis as a result of limiting nitrogen and excess carbon in cotton plants in response to elevated CO₂.
• 3 The CO₂ level and infestation time significantly affected the foliar nitrogen, condensed tannin, gossypol and Bt toxin protein contents of cotton plants after feeding by H. armigera. The interaction between CO₂ levels × cotton variety had a significant effect on foliar nitrogen content after injury by H. armigera.

     

Plant phenolics mediated bottom-up effects of elevated CO2 on Acyrthosiphon pisum and its parasitoid Aphidius avenae

Elevated concentrations of atmospheric CO2 can alter plant secondary metabolites,which play important roles in the interactions among plants,herbivorous insects and natural enemies.However,few studies have examined the cascading effects of host plant secondary metabolites on tri-trophic interactions under elevated CO2(eCO2).In this study,we determined the effects of eCO2 on the growth and foliar phenolics of Medicago truncatula and the cascading effects on two color genotypes oiAcyrthosiphon pisum(pink vs.green)and their parasitoid Aphidius avenae in the field open-top chambers.Our results showed that eCO2 increased photosynthetic rate,nodule number,yield and the total phenolic content of M.truncatula.eCO2 had contrasting effects on two genotypes of A.pisum;the green genotype demonstrated increased population abundance,fecundity,growth and feeding efficiency,while the pink genotype showed decreased fitness and these were closely associated with the foliar genstein content.Furthermore,eCO2 decreased the parasitic rate of A.avenae independent of aphid genotypes.eCO2 prolonged the emergence time and reduced the emergence rate and percentage of females when associated with the green genotype,but little difference,except for increased percentage of females,was observed in A.avenae under eCO2 when associated with the pink genotype,indicating that parasitoids can perceive and discriminate the qualities of aphid hosts.We concluded that eCO2 altered plant phenolics and thus the performance of aphids and parasitoids.Our results indicate that plant phenolics vary by different abiotic and biotic stimuli and could potentially deliver the cascading effects of eCO2 to the higher trophic levels.Our results also suggest that the green genotype is expected to perform better in future eCO2 because of decreased plant resistance after its infestation and decreased parasitic rate.     

Elevated CO2 lessens predation of Chrysopa sinica on Aphis gossypii

Most studies on the effects of elevated atmospheric CO₂ on organisms have focused on the performance of plants or herbivores. Few reports have examined the impact on the ability of predators at the third trophic level. In this experiment, we made use of open‐top chambers to quantify the effects of elevated CO₂ on growth, development, and predatory ability of two successive generations of Chinese lacewing, Chrysopa sinica (Tjeder) (Neuroptera: Chrysopidae), feeding on cotton aphids, Aphis gossypii (Glover) (Hemiptera: Aphididae), which were reared on cotton, Gossypium hirsutum L. (Malvaceae), grown under elevated CO₂ (double ambient vs. ambient). Higher atmospheric CO₂ concentrations reduced the duration of larval development and the survival rate of pupae, and caused decreased weight in adult female C. sinica, but had no significant effects on survival rate of each larval stage, female adult fecundity, egg hatch rate, or adult life span. The predatory ability of larvae in the third instar and the total larval stage of C. sinica that fed on A. gossypii were significantly lower in elevated CO₂ environments. The number of aphids consumed by first‐generation lacewing population did not change significantly with different CO₂ treatments; however, significantly fewer aphids were consumed by the second generation of the lacewing population with elevated CO₂. We speculate that A. gossypii may become a more serious pest under an environment with elevated CO₂ concentrations because of the reduced predatory ability of C. sinica on A. gossypii.     

Individual growth rates do not predict aphid population densities under altered atmospheric conditions

• 1 Altered atmospheric composition, associated with climate change, can modify herbivore population dynamics through CO₂ and/or O₃‐mediated changes in plant quality.
• 2 Although pea aphid Acyrthosiphon pisum genotypes exhibit intraspecific variation in population growth in response to atmospheric composition, the proximate mechanisms underlying this variation are largely unknown.
• 3 By rearing single (green, pink) and mixed (green + pink) pea aphid genotypes on red clover Trifolium pratense at the Aspen Free Air CO₂ and O₃ Enrichment (Aspen FACE) site, we assessed whether: (i) elevated CO₂ and/or O₃ concentrations alter aphid growth and development and (ii) individual aphid growth rates predict aphid population densities.
• 4 We showed that growth and development of individual green and pink aphids were not influenced by CO₂ and/or O₃ concentrations when reared as individual or mixed genotypes. Individual growth rates, however, did not predict population densities.
• 5 Reared as a single genotype, green pea aphid populations decreased in response to elevated CO₂ concentrations, but not in response to elevated CO₂ + O₃ concentrations. Pink pea aphid populations reared as a single genotype were unaffected by augmented CO₂ or O₃. Populations of mixed genotypes, however, were reduced under elevated CO₂ concentrations, irrespective of O₃ concentrations.
• 6 Herbivore population sizes may not readily be predicted from growth rates of individual organisms under atmospheric conditions associated with global climate change.

     

Effects of elevated CO2 and transgenic Bt cotton on plant chemistry, performance, and feeding of an insect herbivore, the cotton bollworm

Effects of elevated atmospheric CO₂ (double‐ambient CO₂) on the growth and metabolism of cotton bollworm, Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae), fed on transgenic Bacillus thuringiensis (Berliner) (Bt) cotton [Cry1A(c)], grown in open‐top chambers, were studied. Two levels of CO₂ (ambient and double‐ambient) and two cotton cultivars (non‐transgenic Simian‐3 and transgenic GK‐12) were deployed in a completely randomized design with four treatment combinations, and the cotton bollworm was reared on each treatment simultaneously. Plants of both cotton cultivars had lower nitrogen and higher total non‐structural carbohydrates (TNC), TNC:Nitrogen ratio, condensed tannin, and gossypol under elevated CO₂. Elevated CO₂ further resulted in a significant decrease in Bt toxin level in GK‐12. The changes in chemical components in the host plants due to increased CO₂ significantly affected the growth parameters of H. armigera. Both transgenic Bt cotton and elevated CO₂ resulted in a reduced body mass, lower fecundity, decreased relative growth rate (RGR), and decreased mean relative growth rate in the bollworms. Larval life‐span was significantly longer for H. armigera fed transgenic Bt cotton. Significantly reduced larval, pupal, and adult moth weights were observed in the bollworms fed elevated CO₂‐grown transgenic Bt cotton compared with those of bollworms reared on non‐transgenic cotton, regardless of the CO₂ level. The efficiency of conversion of ingested food and of digested food of the bollworm were significantly reduced when fed transgenic Bt cotton, but there was no significant CO₂ or CO₂× cotton cultivar interaction. Approximate digestibility of larvae reared on transgenic cotton grown in elevated CO₂ was higher compared to that of larvae fed non‐transgenic cotton grown at ambient CO₂. The damage inflicted by cotton bollworm on cotton, regardless of the presence or absence of insecticidal genes, is predicted to be more serious under elevated CO₂ conditions because of individual compensatory feeding on host plants caused by nitrogen deficiency.     

Impacts of elevated CO2 on Bemisia tabaci infesting Bt cotton and its parasitoid Encarsia formosa

Atmospheric carbon dioxide concentration is expected to rise in the coming decades. Rising atmospheric CO₂ levels may alter plant‐insect‐parasitoid associations due to the indirect effects of CO₂ enrichment on phytochemicals important for herbivore and parasitoid nutrition. Tritrophic effects of elevated CO₂ on Bt cotton (GK‐12) and non‐transgenic (Simian‐3, or S3) cotton [Gossypium hirsutum L. (Malvaceae)], Bemisia tabaci (Gennadius) biotype B (Hemiptera: Aleyrodidae), and its parasitoid Encarsia formosa Gahan (Hymenoptera: Aphelinidae), were examined in open‐top chambers. Significantly, longer egg‐adult developmental duration and higher mortality of nymphs were observed under elevated CO₂ concentrations on both cotton cultivars during three successive generations. However, no significant differences were found in adult longevity, offspring sex ratio, and the number of eggs laid per female adult of B. tabaci fed on transgenic (GK‐12) or non‐transgenic cotton (S3) grown under elevated CO₂. Abundance of B. tabaci adults increased from 10 to 120 per plant and then decreased to 40 per plant through the growing season, but no significant differences in density occurred between CO₂ treatments and between cultivar treatments. Similarly, no significant differences were found in the developmental duration, parasitization rate, and adult emergence rate of E. formosa after parasitizing B. tabaci for three successive generations. Our results showed that the effects of transgenic Bt cotton did not significantly affect the development, survivorship, life span, or fecundity of B. tabaci and its parasitoids. Moreover, interactions between B. tabaci and E. formosa were not significantly affected by elevated CO₂. These results suggest that the biological control of B. tabaci by E. formosa would not be influenced by transgenic Bt cotton and/or elevated CO₂, indicating that the current risk management strategy regarding B. tabaci outbreaks and biocontrol by E. formosa will remain effective if the atmospheric CO₂ level continues to rise.     

Pea aphid promotes amino acid metabolism both in Medicago truncatula and bacteriocytes to favor aphid population growth under elevated CO2

Rising atmospheric CO₂ levels can dilute the nitrogen (N) resource in plant tissue, which is disadvantageous to many herbivorous insects. Aphids appear to be an exception that warrants further study. The effects of elevated CO₂ (750 ppm vs. 390 ppm) were evaluated on N assimilation and transamination by two Medicago truncatula genotypes, a N‐fixing‐deficient mutant (dnf1) and its wild‐type control (Jemalong), with and without pea aphid (Acyrthosiphon pisum) infestation. Elevated CO₂ increased population abundance and feeding efficiency of aphids fed on Jemalong, but reduced those on dnf1. Without aphid infestation, elevated CO₂ increased photosynthetic rate, chlorophyll content, nodule number, biomass, and pod number for Jemalong, but only increased pod number and chlorophyll content for dnf1. Furthermore, aphid infested Jemalong plants had enhanced activities of N assimilation‐related enzymes (glutamine synthetase, Glutamate synthase) and transamination‐related enzymes (glutamate oxalate transaminase, glutamine phenylpyruvate transaminase), which presumably increased amino acid concentration in leaves and phloem sap under elevated CO₂. In contrast, aphid infested dnf1 plants had decreased activities of N assimilation‐related enzymes and transmination‐related enzymes and amino acid concentrations under elevated CO₂. Furthermore, elevated CO₂ up‐regulated expression of genes relevant to amino acid metabolism in bacteriocytes of aphids associated with Jemalong, but down‐regulated those associated with dnf1. Our results suggest that pea aphids actively elicit host responses that promote amino acid metabolism in both the host plant and in its bacteriocytes to favor the population growth of the aphid under elevated CO₂.     

Presence of two host races of Aphis gossypii Glover (Hemiptera: Aphididae) collected in Turkey

The aphid, Aphis gossypii, is a primary pest of citrus, cotton, cucurbits and greenhouse‐grown vegetables in Turkey and throughout Europe. There is some previous empirical data suggesting that host‐adapted genotypes of this aphid exist which may in fact be host‐races. To determine if host races of A. gossypii are indeed present in the eastern Mediterranean region of Turkey, reciprocal host transfer experiments and life table analyses were performed with multiple asexual lineages (= clones) of the aphid collected from different hosts. The collection hosts included citrus, cucumber, eggplant, okra, sweet pepper and cotton. Aphid developmental times on the host from which the aphid was originally collected (= collection or natal host) were shorter (5.2–6.0 days) and had a higher intrinsic rate of population growth (r_m = 0.25–0.44) than the 6.6–7.3 days required when the aphid was reared on a non‐original collection host (= non‐collection host or non‐natal host) and had r_m = 0.03–0.30. Total immature mortality of the cotton clone, especially in the first nymphal stage, was high (51–100%) with low r_m (0–0.03) on cucumber, citrus and sweet pepper. Aphid populations transferred from citrus, eggplant and okra to cotton (r_m = 0.29–0.30) did not differ significantly in their performance from that of the cotton population on cotton (r_m = 0.34), whereas that from sweet pepper and cucumber populations (r_m = 0.22–0.24) were significantly lower. These data have allowed us to separate A. gossypii into two distinct biological groups: (a) a ‘generalist’ population obtained from cucumber, sweet pepper, citrus, eggplant and okra which exhibited statistically better development on cotton; versus (b) a population from cotton which, by comparison on reciprocal hosts, developed poorly on non‐natal hosts except on eggplant. Development of the cotton clone on cucumber and okra was not improved after four successive generations on the non‐natal host. The good development of A. gossypii from eggplant and cotton on these reciprocal hosts suggests that these particular clones were similar, if not identical, host races.     

Plant stomatal closure improves aphid feeding under elevated CO2

Stomata help plants regulate CO₂ absorption and water vapor release in response to various environmental changes, and plants decrease their stomatal apertures and enhance their water status under elevated CO₂. Although the bottom‐up effect of elevated CO₂ on insect performance has been extensively studied, few reports have considered how insect fitness is altered by elevated CO₂‐induced changes in host plant water status. We tested the hypothesis that aphids induce stomatal closure and increase host water potential, which facilitates their passive feeding, and that this induction can be enhanced by elevated CO₂. Our results showed that aphid infestation triggered the abscisic acid (ABA) signaling pathway to decrease the stomatal apertures of Medicago truncatula, which consequently decreased leaf transpiration and helped maintain leaf water potential. These effects increased xylem‐feeding time and decreased hemolymph osmolarity, which thereby enhanced phloem‐feeding time and increased aphid abundance. Furthermore, elevated CO₂ up‐regulated an ABA‐independent enzyme, carbonic anhydrase, which led to further decrease in stomatal aperture for aphid‐infested plants. Thus, the effects of elevated CO₂ and aphid infestation on stomatal closure synergistically improved the water status of the host plant. The results indicate that aphid infestation enhances aphid feeding under ambient CO₂ and that this enhancement is increased under elevated CO₂.     

Parasitoid spectrum (Hymenoptera: Braconidae; Aphelinidae) of the soybean aphid Aphis glycines (Homoptera: Aphididae) in Japan and Indonesia (Java and Bali)

Aphidiine and aphelinid parasitoids collected from the soybean aphid, Aphis glycines, on Glycine max in Japan and Indonesia (Java and Bali) were identified to clarify the parasitoid spectrum of the aphid there. Nine parasitoid species from Japan (Aphidiinae: Aphidius gifuensis, Aphidius sp., Binodoxys communis, Diaeretiella rapae, Lipolexis gracilis, Lysiphlebia japonica; Aphelinidae: Aphelinus asychis, A. gossypii, A. varipes) and two parasitoid species from Indonesia (B. communis, A. gossypii) were found to be associated with A. glycines.     

Top‐down control by Harmonia axyridis mitigates the impact of elevated atmospheric CO2 on a plant–aphid interaction

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Foliar quality influences tree-herbivore-parasitoid interactions: effects of elevated CO<Subscript>2</Subscript>, O<Subscript>3</Subscript>, and plant genotype

This study examined the effects of carbon dioxide (CO₂)-, ozone (O₃)-, and genotype-mediated changes in quaking aspen (Populus tremuloides) chemistry on performance of the forest tent caterpillar (Malacosoma disstria) and its dipteran parasitoid (Compsilura concinnata) at the Aspen Free-Air CO₂ Enrichment (FACE) site. Parasitized and non-parasitized forest tent caterpillars were reared on two aspen genotypes under elevated levels of CO₂ and O₃, alone and in combination. Foliage was collected for determination of the chemical composition of leaves fed upon by forest tent caterpillars during the period of endoparasitoid larval development. Elevated CO₂ decreased nitrogen levels but had no effect on concentrations of carbon-based compounds. In contrast, elevated O₃ decreased nitrogen and phenolic glycoside levels, but increased concentrations of starch and condensed tannins. Foliar chemistry also differed between aspen genotypes. CO₂, O₃, genotype, and their interactions altered forest tent caterpillar performance, and differentially so between sexes. In general, enriched CO₂ had little effect on forest tent caterpillar performance under ambient O₃, but reduced performance (for insects on one aspen genotype) under elevated O₃. Conversely, elevated O₃ improved forest tent caterpillar performance under ambient, but not elevated, CO₂. Parasitoid larval survivorship decreased under elevated O₃, depending upon levels of CO₂ and aspen genotype. Additionally, larval performance and masses of mature female parasitoids differed between aspen genotypes. These results suggest that host-parasitoid interactions in forest systems may be altered by atmospheric conditions anticipated for the future, and that the degree of change may be influenced by plant genotype.     

Co‐occurrence of defensive traits in the potato aphid Macrosiphum euphorbiae

1. Insecticide usage selects strongly for resistance in aphid populations, but this could entail fitness costs in other resistance traits. The potato aphid Macrosiphum euphorbiae Thomas exhibits intraspecific variation in susceptibility to parasitism by braconid wasps and provides a suitable species to study the relation between the defensive traits of parasitism and insecticide resistance. 2. Clonal lines (23 in total) of M. euphorbiae were established from aphids collected in 2013 from geographically separate populations in the U.K. Clonal lines belonged to five aphid genotypes, but one genotype predominated (78% of samples), and the facultative endosymbiont Hamiltonella defensa was detected in c. 40% of lines. 3. Total esterase activity in aphid tissues varied significantly between aphid genotypes and collection areas, but there was no clear pattern in relation to H. defensa infection or between collection sites likely to differ in insecticide pressures. 4. Five clonal lines representing low to moderate levels of enzyme activity, which included different aphid genotypes and presence/absence of H. defensa infection, were assayed for their susceptibility to the parasitoid wasp Aphidius ervi Haliday. Aphid mummification varied significantly between aphid genotypes, with low values in one genotype of aphids irrespective of H. defensa presence. 5. The results revealed that aphid lines belonging to the parasitism‐resistant genotype exhibited moderate levels of total esterase activity, indicating a competitve advantage for this genotype of M. euphorbiae when exposed to chemical and biological control factors in agroecosystems.