Effects of elevated CO2 on the foraging behavior of cotton bollworm, Helicoverpa armigera

Effects of elevated CO2 on the foraging behavior of cotton bollworm Helicoverpa arrnigera Hübner reared on milky grains of spring wheat grown in ambient, 550μL/L and 750μL/L CO2 concentration atmospheres in open-top chambers （OTC） were studied. The results indicated that： （i） elevated CO2 significantly affected both the type and amount of food eaten by H.arrnigera reared on milky grains of ambient CO2-grown wheat were significant higher than those for bollworm larvae reared on wheat grains grown in 550 and 750μL/L CO2 atmospheres; （ii） when bollworm larvae were reared on mixed milky grains from different CO2-grown wheat （food-choice condition）, larval duration increased significantly-pupal weight, adult longevity, and fecundity decreased significantly, comparing with those reared on milky grains of ambient CO2-grown wheat, 550μL/L CO2-grown wheat and 750μL/L CO2-grown wheat respectively; （iii） significant decreases in the contents of fructose and gross protein （GP） and significant increases in the contents of glucose, amylose, total saccharides （TSC）, TSC： GP ratio, free amino acids and soluble protein in the wheat grains with CO2 rising; （iv） and selected-foraging amount/food-choice index of cotton bollworm H.armigera were significantly positive correlated with the contents of fructose and GP of wheat grains, but they had significantly negative relationships with the contents of glucose, amylose, TSC and TSC： GP ratio of wheat grains.     

The effect of fertilizer‐N on the inter‐specific competition among three wheat aphids under elevated CO2

Increasing atmospheric carbon dioxide (ab. CO₂) and fertilizer‐nitrogen (ab. N) applications may have marked direct effects on the plant growth of agricultural crops, and in turn affect the higher trophic level of insect herbivores. In this study, the effects of elevated CO₂ (i.e., 650 µl/L vs. ambient 400 µl/L) and fertilizer‐N (0, 50, 100, 200 kg/ha) on the population abundances and the inter‐specific competition among three co‐occurring species of wheat aphids, Sitobion avenae, Rhopalosiphum padi and Schizaphis graminum, were studied. The grain weight per ear and the 1,000‐grain weight were generally increased when grown under elevated CO₂ and showed a significant effect at the 100 kg/ha (grain weight per ear) and 0, 50 and 100 kg/ha (1,000‐grain weight) N. These two yield indexes increased with increasing fertilizer‐N levels within reasonable limits and reached a maximum at 100 kg/ha. Elevated CO₂ combined with fertilizer‐N levels formed complex indirect effects on the three wheat aphids through the wheat crops they fed on. Elevated CO₂ significantly decreased the niche overlap index (ab. NOI) between S. avenae and R. padi under 0 and 100 kg/ha and that between R. padi and S. graminum under 0 kg/ha, while significantly increased the three NOIs under 50 kg/ha and that between R. padi and S. graminum under 100 and 200 kg/ha. S. avenae and R. padi had the larger population and stronger competition in low‐N condition (0 and 50 kg/ha), which was harmful to wheat yield and quality when combined with its own poor nutrition. Overall, the 100 kg/ha N level was the best option based on the aphid population, competition and wheat yields. Therefore, the balanced relationship formed among fertilizers, plants and insects under 100 kg/ha N was vital for the interactive ecosystem.     

Nitrogen fertiliser affects the functional response and prey consumption of Harmonia axyridis (Coleoptera: Coccinellidae) feeding on cereal aphids

Predator–prey interactions are influenced by nitrogen availability. Wheat (Triticum aestivum cv. Solstice) plants were provided with four levels of nitrogen and examined the responses of coccinellid predator, Harmonia axyridis to cereal aphids, Rhopalosiphum padi and Sitobion avenae. Increased nitrogen application improved nitrogen contents of the plants and also the body weight of cereal aphids feeding on them. In no‐choice feeding trials, H. axyridis consumed more aphids on low fertilised plants, suggesting a compensatory consumption to overcome reduced biomass (lower aphid size). Total biomass devoured by H. axyridis on all nitrogen fertiliser treatments was not statistically different. Logistic regression analysis of the proportion of prey consumed demonstrated that all developmental stages (larval and adult) of H. axyridis exhibited the type II functional response on all nitrogen fertiliser treatments. The rate of successful search (a′) of third and fourth instars and adults were the same across all fertiliser treatments suggesting that nitrogen fertilisation did not affect a′. Maximum handling time for the first instars of H. axyridis on R. padi (3.81 h^?1) and S. avenae (4.59 h^?1) was on the highest nitrogen treatment while minimum handling time was for the adults of H. axyridis on R. padi (0.20 h^?1) and S. avenae (0.20 h^?1) on the lowest nitrogen treatment. Handling time varied at varying fertiliser treatments within all instars and affected the predator's efficiency. The functional response curve, rate of successful search and handling time provide the information needed to understand the predator–prey interaction between H. axyridis and these cereals aphids. This could lead to the development of a better strategy for the biological control of R. padi and S. avenae at any particular level of nitrogen fertiliser regime in the field crops.     

Responses of Photosynthetic and Defence Systems to High Temperature Stress in Quercus suber L Seedlings Grown under Elevated CO2

Abstract: Growth in elevated CO₂ led to an increase in biomass production per plant as a result of enhanced carbon uptake and lower rates of respiration, compared to ambient CO₂-grown plants. No down-regulation of photosynthesis was found after six months of growth under elevated CO₂. Photosynthetic rates at 15°C or 35 °C were also higher in elevated than in ambient CO₂-grown plants, when measured at their respective CO₂ growth condition. Stomata of elevated CO₂-grown plants were less responsive to temperature as compared to ambient CO₂ plants. The after effect of a heat-shock treatment (4 h at 45 °C in a chamber with 80% of relative humidity and 800–1000 tmol m^-2 s^-1 photon flux density) on A_max was less in elevated than in ambient CO₂-grown plants. At the photochemical level, the negative effect of the heat-shock treatment was slightly more pronounced in ambient than in elevated CO₂-grown plants. A greater tolerance to oxidative stress caused by high temperatures in elevated CO₂-grown plants, in comparison to ambient CO₂ plants, is suggested by the increase in superoxide dismutase activity, after 1 h at 45 °C, as well as its relatively high activity after 2 and 4 h of the heat shock in the elevated CO₂-grown plants in contrast with the decrease to residual levels of superoxide dismutase activity in ambient CO₂-grown plants immediately after 1 h at 45 °C. The observed increase in catalase after 1 h at 45 °C in both ambient and elevated CO₂-grown plants, can be ascribed to the higher rates of photorespiration and respiration under this high temperature.     

Effects of elevated CO2 and plant genotype on interactions among cotton,aphids and parasitoids

Abstract Effects of CO₂ level (ambient vs. elevated) on the interactions among three cotton (Gossypium hirsutum) genotypes, the cotton aphid (Aphis gossypii Glover), and its hymenoptera parasitoid (Lysiphlebia japonica Ashrnead) were quantified. It was hypothesized that aphid‐parasitoid interactions in crop systems may be altered by elevated CO₂, and that the degree of change is influenced by plant genotype. The cotton genotypes had high (M9101), medium (HZ401) and low (ZMS13) gossypol contents, and the response to elevated CO₂ was genotype‐specific. Elevated CO₂ increased the ratio of total non‐structural carbohydrates to nitrogen (TNC: N) in the high‐gossypol genotype and the medium‐gossypol genotype. For all three genotypes, elevated CO₂ had no effect on concentrations of gossypol and condensed tannins. A. gossypii fitness declined when aphids were reared on the high‐gossypol genotype versus the low‐gossypol genotype under elevated CO₂. Furthermore, elevated CO₂ decreased the developmental time of L. japonica associated with the high‐gossypol genotype and the low‐gossypol genotype, but did not affect parasitism or emergence rates. Our study suggests that the abundance of A. gossypii on cotton will not be directly affected by increases in atmospheric CO₂. We speculate that A. gossypii may diminish in pest status in elevated CO₂ and high‐gossypol genotype environments because of reduced fitness to the high‐gossypol genotype and shorter developmental time of L. japonica.     

Photosynthetic acclimation and photosynthate partitioning in soybean leaves in response to carbon dioxide enrichment

Photosynthetic rates and photosynthate partitioning were studied in three-week-old soybean [Glycine max (L.) Merr. cv. Williams] plants exposed to either ambient (35 Pa) or elevated (70 Pa) CO₂ in controlled environment chambers. Ambient CO₂-grown plants also were given a single 24 h treatment with 70 Pa CO₂ 1 d prior to sampling. Photosynthetic rates of ambient CO₂-grown plants initially increased 36% when the measurement CO₂ was doubled from 35 to 70 Pa. Photosynthetic rates of the third trifoliolate leaf, both after 1 and 21 d of elevated CO₂ treatment, were 30 to 45% below those of ambient CO₂-grown plants when measured at 35 Pa CO₂. These reduced photosynthetic rates were not due to increased stomatal resistance and were observed for 2 to 8 h after plants given 1 d of CO₂ enrichment were returned to ambient CO₂. Initial and total ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) activities, percent activation, Rubisco protein, soluble protein and leaf chlorophyll content were similar in all CO₂ treatments. Quantum yields of photosynthesis, determined at limiting irradiances and at 35 Pa CO₂, were 0.049±0.003 and 0.038±0.005 mol CO₂ fixed per mol quanta for ambient and elevated CO₂-grown plants, respectively (p<0.05). Leaf starch and sucrose levels were greater in plants grown at 70 than at 35 Pa CO₂. Starch accumulation rates during the day were greater in ambient CO₂-grown plants than in plants exposed to elevated CO₂ for either 1 or 21 d. However, the percentage of C partitioned to starch relative to total C fixed was unaffected by 1 d of CO₂ enrichment. The above results showed that both photosynthetic and starch accumulation rates of soybean leaflets measured at 35 Pa CO₂ were temporarily reduced after 1 and 21 d of CO₂ enrichment. The biochemical mechanism affecting these responses was not identified.Abbreviations SLW- specific leaf weight (g m^–2) - Rubisco- ribulose 1,5-bisphosphate carboxylase/oxygenase - Rul- 5bisP, ribulose 1,5 bisphosphate - DAP- days after planting - SAR- starch accumulation rate - C_i- intercellular CO₂ concentration     

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

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CO2 enrichment predisposes foliage of a eucalypt to freezing injury and reduces spring growth

Seedlings of Eucalyptus pauciflora, were grown in open-top chambers fumigated with ambient and elevated [CO₂], and were divided into two populations using 10% light transmittance screens. The aim was to separate the effects of timing of light interception, temperature and [CO₂] on plant growth. The orientation of the screens exposed plants to a similar total irradiance, but incident during either cold mornings (east-facing) or warm afternoons (west-facing). Following the first autumn freezing event elevated CO₂-grown plants had 10 times more necrotic leaf area than ambient CO₂ plants. West-facing plants had significantly greater (25% more) leaf damage and lower photochemical efficiency (F_v/F_m) in comparison with east-facing plants. Following a late spring freezing event east-facing elevated CO₂ plants suffered a greater sustained loss in F_v/F_m than west-facing elevated CO₂- and ambient CO₂-grown plants. Stomatal conductance was lower under elevated CO₂ than ambient CO₂ except during late spring, with the highest leaf temperatures occurring in west-facing plants under elevated CO₂. These higher leaf temperatures apparently interfered with cold acclimation thereby enhancing frost damage and reducing the ability to take advantage of optimal growing conditions under elevated CO₂.     

Yellowing and photosynthetic decline of barley primary leaves in response to atmospheric CO2 enrichment

The photosynthetic response of barley (Hordeum vulgare L. cv. Brant) primary leaves was studied as a function of chlorosis induced by CO₂ enrichment. Leaf yellowing, measured as changes of chlorophyll a and b, was more extensive in controlled environments at elevated (680 ± 17 µl l^?1) than at ambient (380 ± 21 µl l^?1) CO₂. Stomatal conductance of primary leaves was decreased by growth in elevated CO₂ between 11 and 18 days after sowing (DAS) when measured at both 380 and 680 µl l^?1 CO₂. Internal leaf CO₂ concentration (C_i) was also lower for elevated- compared to ambient-CO₂-grown primary leaves between 11 and 14 DAS. Results suggest that non-stomatal factors were responsible for the decreased photosynthetic rates of elevated- compared to ambient-CO₂-grown primary leaves 18 DAS. Various photochemical measurements, including quantum absorptance (α), minimal (F₀), maximal (F_m), and variable (F_v) chlorophyll fluorescence, as well as the F_v/F_m ratio, were significantly decreased 18 DAS in the elevated- compared to ambient-CO₂ treatment. Photochemical (qP) and nonphotochemical (qN) chlorophyll fluorescence quenching coefficients of 18-day-old primary leaves did not differ between CO₂ treatments. Photosynthetic electron transport rates of photosystem II were slightly lower for elevated- compared to ambient-CO₂-grown primary leaves 18 DAS. Concentrations of α-amino N (i.e. free amino acids) in barley primary leaves were increased by CO₂ enrichment 10 DAS, but subsequently, α-amino N decreased in association with photosynthetic decline. Total acid protease activity was greater in elevated- than in ambient-CO₂-grown leaves 18 DAS. The above findings suggest that photoinhibition and premature senescence were factors in the CO₂-dependent yellowing of barley primary leaves.     

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.     

Leaf temperature of soybean grown under elevated CO2 increases Aphis glycines (Hemiptera: Aphididae) population growth

Abstract Plants grown under elevated carbon dioxide (CO₂) experience physiological changes that influence their suitability as food for insects. To determine the effects of living on soybean (Glycine max Linnaeus) grown under elevated CO₂, population growth of the soybean aphid (Aphis glycines Matsumura) was determined at the SoyFACE research site at the University of Illinois, Urbana‐Champaign, Illinois, USA, grown under elevated (550 μL/L) and ambient (370 μL/L) levels of CO₂. Growth of aphid populations under elevated CO₂ was significantly greater after 1 week, with populations attaining twice the size of those on plants grown under ambient levels of CO₂. Soybean leaves grown under elevated levels of CO₂ were previously demonstrated at SoyFACE to have increased leaf temperature caused by reduced stomatal conductance. To separate the increased leaf temperature from other effects of elevated CO₂, air temperature was lowered while the CO₂ level was increased, which lowered overall leaf temperatures to those measured for leaves grown under ambient levels of CO₂. Aphid population growth on plants grown under elevated CO₂ and reduced air temperature was not significantly greater than on plants grown under ambient levels of CO₂. By increasing Glycine max leaf temperature, elevated CO₂ may increase populations of Aphis glycines and their impact on crop productivity.     

Leaf and canopy responses to elevated CO2 in a pine forest under free-air CO2 enrichment

Physiological responses to elevated CO₂ at the leaf and canopy-level were studied in an intact pine (Pinus taeda) forest ecosystem exposed to elevated CO₂ using a free-air CO₂ enrichment (FACE) technique. Normalized canopy water-use of trees exposed to elevated CO₂ over an 8-day exposure period was similar to that of trees exposed to current ambient CO₂ under sunny conditions. During a portion of the exposure period when sky conditions were cloudy, CO₂-exposed trees showed minor (7%) but significant reductions in relative sap flux density compared to trees under ambient CO₂ conditions. Short-term (minutes) direct stomatal responses to elevated CO₂ were also relatively weak (5% reduction in stomatal aperture in response to high CO₂ concentrations). We observed no evidence of adjustment in stomatal conductance in foliage grown under elevated CO₂ for nearly 80 days compared to foliage grown under current ambient CO₂, so intrinsic leaf water-use efficiency at elevated CO₂ was enhanced primarily by direct responses of photosynthesis to CO₂. We did not detect statistical differences in parameters from photosynthetic responses to intercellular CO₂ (A _net-C _i curves) for Pinus taeda foliage grown under elevated CO₂ (550 mol mol^–1) for 50–80 days compared to those for foliage grown under current ambient CO₂ from similar-sized reference trees nearby. In both cases, leaf net photosynthetic rate at 550 mol mol^–1 CO₂ was enhanced by approximately 65% compared to the rate at ambient CO₂ (350 mol mol^–1). A similar level of enhancement under elevated CO₂ was observed for daily photosynthesis under field conditions on a sunny day. While enhancement of photosynthesis by elevated CO₂ during the study period appears to be primarily attributable to direct photosynthetic responses to CO₂ in the pine forest, longer-term CO₂ responses and feedbacks remain to be evaluated.     

Effects of elevated pCO2 on the metabolism of a temperate rhodolith Lithothamnion corallioides grown under different temperatures

Coralline algae are considered among the most sensitive species to near future ocean acidification. We tested the effects of elevated pCO₂ on the metabolism of the free‐living coralline alga Lithothamnion corallioides (“maerl”) and the interactions with changes in temperature. Specimens were collected in North Brittany (France) and grown for 3 months at pCO₂ of 380 (ambient pCO₂), 550, 750, and 1000 μatm (elevated pCO₂) and at successive temperatures of 10°C (ambient temperature in winter), 16°C (ambient temperature in summer), and 19°C (ambient temperature in summer +3°C). At each temperature, gross primary production, respiration (oxygen flux), and calcification (alkalinity flux) rates were assessed in the light and dark. Pigments were determined by HPLC. Chl a, carotene, and zeaxanthin were the three major pigments found in L. corallioides thalli. Elevated pCO₂ did not affect pigment content while temperature slightly decreased zeaxanthin and carotene content at 10°C. Gross production was not affected by temperature but was significantly affected by pCO₂ with an increase between 380 and 550 μatm. Light, dark, and diel (24 h) calcification rates strongly decreased with increasing pCO₂ regardless of the temperature. Although elevated pCO₂ only slightly affected gross production in L. corallioides, diel net calcification was reduced by up to 80% under the 1,000 μatm treatment. Our findings suggested that near future levels of CO₂ will have profound consequences for carbon and carbonate budgets in rhodolith beds and for the sustainability of these habitats.     

Impact of elevated CO2 concentration on ultrastructure of pericarp and composition of grain in three Triticum species of different ploidy levels

Triticum species of three ploidies were grown under ambient (375 μl/l) or elevated (FACE, 550 μl/l) CO₂ concentration [CO₂] to evaluate their response to CO₂ enrichment. The consistent effect of elevated CO₂ was an increase in concentration of starch and decrease in concentration of protein in the grain. Transmission electron micrographs revealed an increase in width and area of chloroplasts, and change in shape from elliptical in ambient to round in elevated [CO₂]. There was a corresponding increase in starch grain size and number in chloroplasts. The large starch grains distributed among thylakoids resulted in separation and distortion of internal membrane system in chloroplasts. The level of response was different in species of different ploidy levels. Maximum increase in starch concentration, and least decrease in protein concentration, was observed in Triticum dicoccoides, which also proved the most suitable species in terms of C:N ratio.     

Does carbon partitioning in ectomycorrhizal pine seedlings under elevated CO2 vary with fungal species?

Enhanced soil respiration in response to elevated atmospheric CO₂ has been demonstrated, and ectomycorrhizal (ECM) fungi are of particular interest since they partition host-derived photoassimilates belowground. Although a strong response of ECM fungi to elevated CO₂ has been shown, little is still known about the functional diversity among species. We studied carbon (C) partitioning in mycorrhizal Scots pine seedlings in response to short-term CO₂ enrichment, using seven ECM species with different ecological strategies. Mycorrhizal associations were synthesised and seedlings grown in large Petri dishes containing peat:vermiculite and nutrient solution for 10–15 weeks, after which half of the microcosms were exposed to elevated CO₂ treatment (710 ppm) for 15 days and the other half were kept in ambient CO₂ treatment. Partitioning of C was quantified by pulse labelling the seedlings with ¹⁴CO₂ and examining the distribution of labelled assimilates in shoot, root and extraradical mycelial compartments by destructive harvest and liquid scintillation counting. Fungal biomass was determined with PLFA analysis. The respiratory loss of ¹⁴CO₂ was on average greater in the elevated CO₂ treatment for most species compared to the ambient CO₂ treatment. More label was retrieved in the shoots in the ambient CO₂ treatment compared to elevated CO₂ (significant for P. involutus and P. fallax). Greater amounts of label were found in the extraradical mycelial compartment in all species (except P. involutus) in elevated CO₂ compared to ambient CO₂ (significant for L. bicolor, P. byssinum, P. fallax and R. roseolus). Fungal biomass production increased significantly with elevated CO₂ for two species (H. velutipes and A. muscaria); three species (P. fallax, P. involutus and R. roseolus) showed a similar but non-significant trend, whereas L. bicolor and P. byssinum produced less biomass in elevated CO₂ compared to ambient CO₂. When ¹⁴C in the mycelial compartment and respiration was expressed per unit fungal PLFA the difference between CO₂ treatments disappeared. We demonstrated that different ECM fungal isolates respond differently in C partitioning in response to CO₂ enrichment. These results suggest that under certain growth conditions, when nutrients are not limiting, ECM fungi respond rapidly to increasing C-availability through changed biomass production and respiration.     

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.     

Leaf age effects of elevated CO2-grown white oak leaves on spring-feeding lepidopterans

Folivorous insect responses to elevated CO₂-grown tree species may be complicated by phytochemical changes as leaves age. For example, young expanding leaves in tree species may be less affected by enriched CO₂-alterations in leaf phytochemistry than older mature leaves due to shorter exposure times to elevated CO₂ atmospheres. This, in turn, could result in different effects on early vs. late instar larvae of herbivorous insects. To address this, seedlings of white oak (Quercus alba L.), grown in open-top chambers under ambient and elevated CO₂, were fed to two important early spring feeding herbivores; gypsy moth (Lymantria dispar L.), and forest tent caterpillar (Malacosoma disstria Hübner). Young, expanding leaves were presented to early instar larvae, and older fully expanded or mature leaves to late instar larvae. Young leaves had significantly lower leaf nitrogen content and significantly higher total nonstructural carbohydrate:nitrogen ratio as plant CO₂ concentration rose, while nonstructural carbohydrates and total carbon-based phenolics were unaffected by plant CO₂ treatment. These phytochemical changes contributed to a significant reduction in the growth rate of early instar gypsy moth larvae, while growth rates of forest tent caterpillar were unaffected. The differences in insect responses were attributed to an increase in the nitrogen utilization efficiency (NUE) of early instar forest tent caterpillar larvae feeding on elevated CO₂-grown leaves, while early instar gypsy moth larval NUE remained unchanged among the treatments. Later instar larvae of both insect species experienced larger reductions in foliage quality on elevated CO₂-grown leaves than earlier instars, as the carbohydrate:nitrogen ratio of leaves substantially increased. Despite this, neither insect species exhibited changes in growth or consumption rates between CO₂ treatments in the later instar. An increase in NUE was apparently responsible for offsetting reduced foliar nitrogen for the late instar larvae of both species.     

Elevated CO2 and plant nitrogen-use: is reduced tissue nitrogen concentration size-dependent?

Plants often respond to elevated atmospheric CO₂ levels with reduced tissue nitrogen concentrations relative to ambient CO₂-grown plants when comparisons are made at a common time. Another common response to enriched CO₂ atmospheres is an acceleration in plant growth rates. Because plant nitrogen concentrations are often highest in seedlings and subsequently decrease during growth, comparisons between ambient and elevated CO₂-grown plants made at a common time may not demonstrate CO₂-induced reductions in plant nitrogen concentration per se. Rather, this comparison may be highlighting differences in nitrogen concentration between bigger, more developed plants and smaller, less developed plants. In this study, we directly examined whether elevated CO₂ environments reduce plant nitrogen concentrations independent of changes in plant growth rates. We grew two annual plant species. Abutilon theophrasti (C₃ photosynthetic pathway) and Amaranthus retroflexus (C₄ photosynthetic pathway), from seed in glass-sided growth chambers with atmospheric CO₂ levels of 350 mol·mol^–1 or 700 mol·mol^–1 and with high or low fertilizer applications. Individual plants were harvested every 2 days starting 3 days after germination to determine plant biomass and nitrogen concentration. We found: 1. High CO₂-grown plants had reduced nitrogen concentrations and increased biomass relative to ambient CO₂-grown plants when compared at a common time; 2. Tissue nitrogen concentrations did not vary as a function of CO₂ level when plants were compared at a common size; and 3. The rate of biomass accumulation per rate of increase in plant nitrogen was unaffected by CO₂ availability, but was altered by nutrient availability. These results indicate that a CO₂-induced reduction in plant nitrogen concentration may not be due to physiological changes in plant nitrogen use efficiency, but is probably a size-dependent phenomenon resulting from accelerated plant growth.     

Possible coexistence of Harmonia axyridis Pallas (Coleoptera: Coccinellidae) and Diaeretiella rapae M'Intosh (Hymenoptera: Braconidae) in the biological control of Lipaphis erysimi (Homoptera: Aphididae)

To study the interactions between the aphidophagous predator Harmonia axyridis Pallas (Coleoptera: Coccinellidae) and the specialist aphid parasitoid Diaeretiella rapae M'Intosh (Hymenoptera: Braconidae) in the biological control of mustard aphid, Lipaphis erysimi (Homoptera: Aphididae), the prey discrimination by H. axyridis among unparasitized, non-mummified parasitized and mummified aphids was examined under laboratory conditions. Prey/host selections were also tested by offering L. erysimi at various developmental stages to assess the possibility of coexistence between the two species, so the prey preference of H. axyridis when D. rapae parasitize aphids, and the host preference of D. rapae when H. axyridis interfered with the parasitization were detected. We found that H. axyridis could discriminate against mummies rather than non-mummified parasitized aphids. The ladybug showed a significantly positive preference for adult prey when D. rapae turned aphids into mummies, while D. rapae tended to parasitize younger nymphal aphids when H. axyridis was introduced. The present study suggests the prey discrimination against mummies by H. axyridis, and indicates that H. axyridis and D. rapae can avoid resource competition by attacking different and non-overlapping developmental stages of aphid. Thus, H. axyridis and D. rapae can potentially coexist and establish a stable ecosystem in the biological control of L. erysimi.     

Decline in gypsy moth (Lymantria dispar) performance in an elevated CO2 atmosphere depends upon host plant species

Plant species differ broadly in their responses to an elevated CO₂ atmosphere, particularly in the extent of nitrogen dilution of leaf tissue. Insect herbivores are often limited by the availability of nutrients, such as nitrogen, in their host plant tissue and may therefore respond differentially on different plant species grown in CO₂-enriched environments. We reared gyspy moth larvae (Lymantria dispar) in situ on seedlings of yellow birch (Betula allegheniensis) and gray birch (B. populifolia) grown in an ambient (350 ppm) or elevated (700 ppm) CO₂ atmosphere to test whether larval responses in the elevated CO₂ atmosphere were species-dependent. We report that female gypsy moths (Lymantria dispar) reared on gray birch (Betula populifolia) achieved similar pupal masses on plants grown at an ambient or an elevated CO₂ concentration. However, on yellow birch (B. allegheniensis), female pupal mass was 38% smaller on plants in the elevated-CO₂ atmosphere. Larval mortality was significantly higher on yellow birch than gray birch, but did not differ between the CO₂ treatments. Relative growth rate declined more in the elevated CO₂ atmosphere for larvae on yellow birch than for those on gray birch. In preference tests, larvae preferred ambient over elevated CO₂-grown leaves of yellow birch, but showed no preference between gray birch leaves from the two CO₂ atmospheres. This differential response of gypsy moths to their host species corresponded to a greater decline in leaf nutritional quality in the elevated CO₂ atmosphere in yellow birch than in gray birch. Leaf nitrogen content of yellow birch dropped from 2.68% to 1.99% while that of gray birch leaves only declined from 3.23% to 2.63%. Meanwhile, leaf condensed tannin concentration increased from 8.92% to 11.45% in yellow birch leaves while gray birch leaves only increased from 10.72% to 12.34%. Thus the declines in larval performance in a future atmosphere may be substantial and host-species-specific.