Attractiveness of CO2 released by root respiration fades on the background of root exudates

Plants are endangered at their roots by soil-dwelling rhizophagous insects. These below-ground living herbivores may orient to the source of carbon dioxide (CO₂), an ubiquitous volatile released by respiring plant roots. Here, we studied the interaction of CO₂ and other plant root-derived chemical stimuli with regard to the chemical orientation of the polyphagous larvae of Melolontha melolontha L. (Scarabaeidae). A soil arena was developed that enabled both determination of the actual soil CO₂ concentration and the behavioural response of an insect to (a) CO₂ gradients per se, (b) chemical stimuli released from respiring, undamaged roots of plants potted into vermiculite in this arena and (c) combinations of CO₂ gradients and root-derived stimuli. In a root-free arena, larvae of M. melolontha oriented to the source of synthetic CO₂. However, similar CO₂ gradients generated by host plant roots did not attract the larvae. Neither did a synthetic CO₂ gradient combined with aqueous extracts from rhizospheres with undamaged plant roots elicit an attractive effect. Our data suggest that orientation of cockchafer larvae within CO₂ gradients generated by respiring roots is ‘masked’ by an aqueous extract from a rhizosphere with undamaged roots. The results emphasise that effects of behaviour modifying plant-derived compounds need to be investigated against the background of naturally co-occurring chemicals. The significance of our results for orientation of soil living insects is discussed with respect to abiotic conditions in natural soil and the role of soil microorganisms for the attractiveness of plant roots.     

Laboratory and field evaluations of chemical and plant‐derived potential repellents against Culicoides biting midges in northern Spain

The efficacy of 23 compounds in repelling Culicoides biting midges (Diptera: Ceratopogonidae), particularly Culicoides obsoletus (Meigen) females, was determined by means of a Y‐tube olfactometer. The 10 most effective compounds were further evaluated in landing bioassays. The six most promising compounds (including chemical and plant‐derived repellents) were evaluated at 10% and 25% concentrations in field assays using Centers for Disease Control (CDC) light traps. At least three compounds showed promising results against Culicoides biting midges with the methodologies used. Whereas olfactometer assays indicated DEET at 1 µg/µL to be the most effective repellent, filter paper landing bioassays showed plant‐derived oils to be better. Light traps fitted with polyester mesh impregnated with a mixture of octanoic, decanoic and nonanoic fatty acids at 10% and 25% concentrations collected 2.2 and 3.6 times fewer midges than control traps and were as effective as DEET, which is presently considered the reference standard insect repellent. The best plant‐derived product was lemon eucalyptus oil. Although these have been reported as safe potential repellents, the present results indicate DEET and the mixture of organic fatty acids to be superior and longer lasting.     

Assessment of the repellent effect of Lippia alba essential oil and major monoterpenes on the cattle tick Rhipicephalus microplus

The control of Rhipicephalus microplus (Ixodida: Ixodidae) is achieved using synthetic acaricides. However, resistant tick populations are widespread around the world. Plant essential oils can act as repellents, keeping ticks away from hosts and decreasing the selection pressure on synthetic acaricides. The aim of this study was to evaluate the in vitro repellent effect of Lippia alba essential oil on R. microplus larvae. Leaves from two L. alba genotypes maintained under the same agronomic and environmental conditions were collected. Essential oil was extracted by hydrodistillation and analysed by gas chromatography–mass spectrometry (GC‐MS). The major monoterpenes detected in the chemical analysis were commercially acquired and tested. For the repellency test, a glass rod was vertically fixed to measure active climbing of approximately 30 R. microplus larvae aged 14–21 days in response to essential oils and monoterpenes. Repellency was evaluated at 1 h, 3 h and 5 h after treatment. Variation in repellent action was detected between the genotypes. The major monoterpenes identified in the essential oils (limonene and carvone) showed low repellent effects in comparison with intact essential oils. Thus, the present results showed that L. alba essential oil contains bioactive compounds with great repellent activity against ticks that varies according to the plant genotype.     

Host-odour recognition in two tick species is coded in a blend of vertebrate volatiles

The questing behaviour of ixodid ticks serves for identification and localisation of approaching hosts and is evoked by carbon dioxide, vibrations, visual and odour stimuli. In an olfactometer, we examined the specificity of the questing response of larvae of Boophilus microplus, a one-host tick which develops mainly on cattle, and Ixodes ricinus, a three-host tick with a broader host spectrum. While all mammalian odours tested were equally stimulatory for I. ricinus, B. microplus was clearly more activated by bovine odours. A phenolic fraction of bovine odour stimulated B. microplus only. Attractive components of the host odours were identified by exposing the ticks to single chemicals and mixtures. Single chemicals stimulated questing responses only at levels higher than the levels detected in the bovine odour. However, an artificial odour blend of 37 pure chemicals, diluted to concentrations at which the individual components were inactive, proved to be as effective as natural host odour for both tick species. Further fractionation of the blend revealed that the combinatory effect was achieved by only 7 compounds in both species. Although B. microplus responded to the same synergistic mixture of volatiles as I. ricinus, it showed significant higher sensitivity to the cattle-associated compounds 1-octen-3-ol and 2-nitrophenol and this might contribute to its host-specificity. Accepted: 20 March 1999     

Will krill fare well under Southern Ocean acidification?

Antarctic krill embryos and larvae were experimentally exposed to 380 (control), 1000 and 2000 µatm pCO₂ in order to assess the possible impact of ocean acidification on early development of krill. No significant effects were detected on embryonic development or larval behaviour at 1000 µatm pCO₂; however, at 2000 µatm pCO₂ development was disrupted before gastrulation in 90 per cent of embryos, and no larvae hatched successfully. Our model projections demonstrated that Southern Ocean sea water pCO₂ could rise up to 1400 µatm in krill''s depth range under the IPCC IS92a scenario by the year 2100 (atmospheric pCO₂ 788 µatm). These results point out the urgent need for understanding the pCO₂-response relationship for krill developmental and later stages, in order to predict the possible fate of this key species in the Southern Ocean.     

Rising CO2 concentrations affect settlement behaviour of larval damselfishes

Reef fish larvae actively select preferred benthic habitat, relying on olfactory, visual and acoustic cues to discriminate between microhabitats at settlement. Recent studies show exposure to elevated carbon dioxide (CO₂) impairs olfactory cue recognition in larval reef fishes. However, whether this alters the behaviour of settling fish or disrupts habitat selection is unknown. Here, the effect of elevated CO₂ on larval behaviour and habitat selection at settlement was tested in three species of damselfishes (family Pomacentridae) that differ in their pattern of habitat use: Pomacentrus amboinensis (a habitat generalist), Pomacentrus chrysurus (a rubble specialist) and Pomacentrus moluccensis (a live coral specialist). Settlement-stage larvae were exposed to current-day CO₂ levels or CO₂ concentrations that could occur by 2100 (700 and 850 ppm) based on IPCC emission scenarios. First, pair-wise choice tests were performed using a two-channel flume chamber to test olfactory discrimination between hard coral, soft coral and coral rubble habitats. The habitat selected by settling fish was then compared among treatments using a multi-choice settlement experiment conducted overnight. Finally, settlement timing between treatments was compared across two lunar cycles for one of the species, P. chrysurus. Exposure to elevated CO₂ disrupted the ability of larvae to discriminate between habitat odours in olfactory trials. However, this had no effect on the habitats selected at settlement when all sensory cues were available. The timing of settlement was dramatically altered by CO₂ exposure, with control fish exhibiting peak settlement around the new moon, whereas fish exposed to 850 ppm CO₂ displaying highest settlement rates around the full moon. These results suggest larvae can rely on other sensory information, such as visual cues, to compensate for impaired olfactory ability when selecting settlement habitat at small spatial scales. However, rising CO₂ could cause larvae to settle at unfavourable times, with potential consequences for larval survival and population replenishment.     

Behavioural responses of Ixodes ricinus nymphs to carbon dioxide and rodent odour

Many haematophagous ectoparasites use carbon dioxide (CO₂) and host odour to detect and locate their hosts. The tick Ixodes ricinus (Linnaeus) (Ixodida: Ixodidae) walks only small distances and quests in vegetation until it encounters a host. The differential effects of CO₂ and host odour on the host‐finding behaviour of I. ricinus have, however, never been clarified and hence represent the subject of this study. The effects of CO₂ and odour from bank voles on the activation and attraction of I. ricinus nymphs were analysed in a Y‐tube olfactometer. Carbon dioxide evoked a response in the absence and presence of host odour, but did not attract nymphs. Host odour, however, did not evoke a response but did attract nymphs in the absence and presence of CO₂. The current results show that CO₂ is an activator, but not an attractant, and that host odour is an attractant, but not an activator, of I. ricinus nymphs, and provide ecological insights into the host‐finding behaviour of I. ricinus.     

Interactive direct and plant‐mediated effects of elevated atmospheric [CO2] and temperature on a eucalypt‐feeding insect herbivore

Understanding the direct and indirect effects of elevated [CO₂] and temperature on insect herbivores and how these factors interact are essential to predict ecosystem‐level responses to climate change scenarios. In three concurrent glasshouse experiments, we measured both the individual and interactive effects of elevated [CO₂] and temperature on foliar quality. We also assessed the interactions between their direct and plant‐mediated effects on the development of an insect herbivore of eucalypts. Eucalyptus tereticornis saplings were grown at ambient or elevated [CO₂] (400 and 650 μmol mol^?1 respectively) and ambient or elevated ( + 4 °C) temperature for 10 months. Doratifera quadriguttata (Lepidoptera: Limacodidae) larvae were feeding directly on these trees, on their excised leaves in a separate glasshouse, or on excised field‐grown leaves within the temperature and [CO₂] controlled glasshouse. To allow insect gender to be determined and to ensure that any sex‐specific developmental differences could be distinguished from treatment effects, insect development time and consumption were measured from egg hatch to pupation. No direct [CO₂] effects on insects were observed. Elevated temperature accelerated larval development, but did not affect leaf consumption. Elevated [CO₂] and temperature independently reduced foliar quality, slowing larval development and increasing consumption. Simultaneously increasing both [CO₂] and temperature reduced these shifts in foliar quality, and negative effects on larval performance were subsequently ameliorated. Negative nutritional effects of elevated [CO₂] and temperature were also independently outweighed by the direct positive effect of elevated temperature on larvae. Rising [CO₂] and temperature are thus predicted to have interactive effects on foliar quality that affect eucalypt‐feeding insects. However, the ecological consequences of these interactions will depend on the magnitude of concurrent temperature rise and its direct effects on insect physiology and feeding behaviour.     

Early Exposure of Bay Scallops (Argopecten irradians) to High CO2 Causes a Decrease in Larval Shell Growth

Ocean acidification, characterized by elevated pCO₂ and the associated decreases in seawater pH and calcium carbonate saturation state (Ω), has a variable impact on the growth and survival of marine invertebrates. Larval stages are thought to be particularly vulnerable to environmental stressors, and negative impacts of ocean acidification have been seen on fertilization as well as on embryonic, larval, and juvenile development and growth of bivalve molluscs. We investigated the effects of high CO₂ exposure (resulting in pH = 7.39, Ω_ar = 0.74) on the larvae of the bay scallop Argopecten irradians from 12 h to 7 d old, including a switch from high CO₂ to ambient CO₂ conditions (pH = 7.93, Ω_ar = 2.26) after 3 d, to assess the possibility of persistent effects of early exposure. The survival of larvae in the high CO₂ treatment was consistently lower than the survival of larvae in ambient conditions, and was already significantly lower at 1 d. Likewise, the shell length of larvae in the high CO₂ treatment was significantly smaller than larvae in the ambient conditions throughout the experiment and by 7 d, was reduced by 11.5%. This study also demonstrates that the size effects of short-term exposure to high CO₂ are still detectable after 7 d of larval development; the shells of larvae exposed to high CO₂ for the first 3 d of development and subsequently exposed to ambient CO₂ were not significantly different in size at 3 and 7 d than the shells of larvae exposed to high CO₂ throughout the experiment.     

Host location and behavioural response patterns of the parasitoid,Tachinaephagus zealandicus Ashmead (Hymenoptera: Encyrtidae), to host and host‐habitat odours

Abstract 1. To overcome the challenge of host location in patchy and complex environments, many parasitoids exploit host‐habitat derived odour cues. This study investigated the role of odour cues used during host location by the generalist parasitoid, Tachinaephagus zealandicus, a common parasitoid of Dipteran larvae found in association with decomposing carrion. 2. A Y‐tube olfactometer was used to test aspects of host searching behaviour of T. zealandicus females under different choice scenarios: combinations of host–liver complex, eaten liver, unwashed host larvae, washed host larvae, liver in six stages of decay, and control (no odour). 3. T. zealandicus females demonstrated an innate response to, and preference for, odours arising from liver substrates following interaction with the host. However, this was dependent on the stage of meat decay. Females were not attracted to liver that had never been in contact with a host, regardless of the stage of decay. There were discernible differences in parasitoid movement and behaviour under different choice scenarios. The absence of odour stimulus elicited limited movement about the olfactometer. In the presence of any odour stimuli, females were more active within the olfactometer, and when presented with two odour options T. zealandicus females demonstrate a heightened investigation response, moving across all regions of the olfactometer. 4. Knowledge of the cues that mediate host location for T. zealandicus can contribute to fields such as forensic entomology, where the presence of parasitoids on decomposing remains can be used to estimate time since death.     

Aedes aegypti Mosquitoes Exhibit Decreased Repellency by DEET following Previous Exposure

DEET (N,N-Diethyl-m-toluamide) is one of the most widely used mosquito repellents. Although DEET has been shown to be extremely effective, recent studies have revealed that certain individual insects are unaffected by its presence. A genetic basis for this has been shown in Aedes aegypti mosquitoes and the fruit fly Drosophila melanogaster, but, for the triatomine bug, Rhodnius prolixus, a decrease in response to DEET occurred shortly after previous exposure, indicating that non-genetic factors may also be involved in DEET “insensitivity”. In this study, we examined host-seeking behaviour and electrophysiological responses of A. aegypti after pre-exposure to DEET. We found that three hours after pre-exposure the mosquitoes showed behavioural insensitivity, and electroantennography revealed this correlated with the olfactory receptor neurons responding less to DEET. The change in behaviour as a result of pre-exposure to DEET has implications for the use of repellents and the ability of mosquitoes to overcome them.     

Pre‐treatment of Stegomyia aegypti mosquitoes with a sublethal dose of imidacloprid impairs behavioural avoidance induced by lemon oil and DEET

The present study was conducted to determine whether imidacloprid can impair the avoidance behaviour of the mosquito Stegomyia aegypti. Laboratory investigations using a T‐maze apparatus showed that St. aegypti mosquitoes present long term avoidance behaviour when they are exposed to repetitive trials with lemon oil and DEET. The present study tested the effect of a sublethal dose of imidacloprid on the avoidance behaviour of St. aegypti mosquitoes over a 48 h period. Data suggest that 0.5 ng of imidacloprid/mosquito reduces the avoidance behaviour of mosquitoes exposed to lemon oil, on the first day of exposure, after the second trial; whereas imidacloprid affected DEET repellency only the first day of exposure, after the second trial. Imidacloprid was toxic against St. aegypti mosquitoes, and at sublethal doses was able to impair the repellency induced by lemon oil and DEET. The present data were consistent with the finding that St. aegypti mosquitoes exhibit long term avoidance behaviour, and treatment of mosquitoes with a sublethal dose of imidacloprid under DEET application can affect the repellency of DEET against St. aegypti.     

Effects of elevated CO2 on development and larval food-plant preference in the butterfly Coenonympha pamphilus (Lepidoptera, Satyridae)

The objective of this study was to determine how increasing atmospheric CO₂ change plant tissue quality in four native grassland grass species (Agrostis stolonifera, Anthoxanthum odoratum, Festuca rubra, Poa pratensis) which are all larval food‐plants of Coenonympha pamphilus (Lepidoptera, Satyridae). We assessed the effect of these changes on the performance and larval food‐plant preference of C. pamphilus in a greenhouse experiment. Furthermore, we tested the interactive effects of elevated CO₂ and soil nutritional availability in F. rubra and its effect an larval development of C. pamphilus. In general, elevated CO₂ decreased leaf water concentration, nitrogen concentration and specific leaf area (SLA), while leaf starch concentration was increased in all grass species. A species‐specific reaction to elevated CO₂ was only found for foliar starch concentration. P. pratensis did not increase its starch concentration under elevated CO₂ conditions, whereas the other three species did. Fertilisation, investigated only for F. rubra, increased leaf nitrogen concentration and amplified the CO₂‐induced decrease in leaf nitrogen. Development time of C. pamphilus was on the average prolonged by two days under elevated CO₂ and the prolongation differed from 0.7 to 5.3 days among food‐plant species. Pupal fresh weight differed marginally between CO₂ treatments. Fertilisation of the larval food‐plant F. rubra shortened development time by one day and significantly increased pupal and adult fresh weights. C. pamphilus larvae showed a clear food‐plant preference among grass species at the age of 36 h or older. Additionally, a change of food‐plant preference under elevated CO₂ was found. Larvae at ambient CO₂ preferred Agrostis stolonifera and F. rubra, while under elevated CO₂Anthoxanthum odoratum and P. pratensis were preferred. The present study demonstrates that larval development of C. pamphilus is affected by food‐plant species and CO₂ induced changes in foliar chemistry. Although we found some species‐specific reactions to elevated CO₂ for foliar chemistry, no such CO₂ by species interaction was found for insect development. The change in food‐plant preference of larvae under elevated CO₂ implies potential changes in selection pressure for grass species and might therefore affect evolutionary processes.     

Repellencies and toxicities of five ant-derived defensive compounds against the lone star tick, <Emphasis Type="Italic">Amblyomma americanum</Emphasis> (Acari: Ixodidae)

The lone star tick, Amblyomma americanum, is a vector of several important human and animal diseases. This tick species has rapidly expanded in its geographic distribution, and its aggressive behavior has increased the risk of tick-borne diseases in these new areas. Repellents are recommended by the Centers for Disease Control and Prevention (CDC) for protection against tick bites. DEET is the most common repellent, but public concerns over its safety have increased the need for alternative safe and efficacious tick repellents. Several naturally derived animal compounds have been tested against other species of ticks or other arthropod pests, but not against A. americanum. Based on EC₅₀ values obtained using a vertical paper bioassay, decylamine and MT-710 (a 2-tridecanone formulation) were found to be as repellent as DEET. Those two substances along with 2-tridecanone were also found to be as repellent as DEET when their EC₉₅ values were compared. Lone star ticks were more susceptible to the toxic effects of DEET in glass vial assays than all of the ant-derived defensive compounds/formulations. These results suggest that the ant-derived defensive compounds are likely more effective lone star tick repellents than DEET, but they are not as toxic as DEET towards the ticks. The suitability of these compounds for use as personal repellents, as well as at the landscape scale, should be explored.     

CO2 and O3 effects on host plant preferences of the forest tent caterpillar (Malacosoma disstria)

Elevated levels of CO₂ and O₃ affect plant growth and phytochemistry, which in turn can alter physiological performance of associated herbivores. Little is known, however, about how generalist insect herbivores respond behaviorally to CO₂‐ and O₃‐mediated changes in their host plants. This research examined the effects of elevated CO₂ and O₃ levels on host plant preferences and consumption of forest tent caterpillar (FTC, Malacosoma disstria Hbn.) larvae. Dual choice feeding assays were performed with foliage from birch (Betula papyrifera Marsh.) and aspen (Populus tremuloides Michx., genotypes 216 and 259). Trees were grown at the Aspen Free Air CO₂ Enrichment (FACE) facility near Rhinelander, WI, USA, and had been exposed to ambient or elevated concentrations of CO₂ and/or O₃. Levels of nutritional and secondary compounds were quantified through phytochemical analyses. The results showed that elevated O₃ levels increased FTC larval preferences for birch compared with aspen, whereas elevated CO₂ levels had the opposite effect. In assays with the two aspen genotypes, addition of both CO₂ and O₃ caused a shift in feeding preferences from genotype 259 to genotype 216. Consumption was unaffected by experimental treatments in assays comparing aspen and birch, but were increased for larvae given high O₃ foliage in the aspen genotype assays. Elevated levels of CO₂ and O₃ altered tree phytochemistry, but did not explain shifts in feeding preferences. The results demonstrate that increased levels of CO₂ and O₃ can alter insect host plant preferences both between and within tree species. Also, consequences of altered host quality (e.g., compensatory consumption) may be buffered by partial host shifts in situations when alternative plant species are available. Environmentally induced changes in host plant preferences may have the potential to alter the distribution of herbivory across plant genotypes and species, as well as competitive interactions among them.     

Efficacy of Beauveria bassiana for control of Tribolium castaneum with reduced oxygen and increased carbon dioxide

This study investigated the effect of atmosphere modification, a widely adopted means of insect control in stored products, on the efficacy of Beauveria bassiana for one of the most difficult to control pests, Tribolium castaneum. Oxygen reduction to 5% (±1%) as opposed to CO₂ elevation to 40% (±2%) for the first 72 h of fungus exposure resulted in significantly greater larval mortality than fungus exposure under ambient atmospheres. Both treatments reduced pupation of older larvae suggesting that slowed development may be a beneficial factor for fungal efficacy. CO₂ elevation but not O₂ reduction significantly affected the mortality of adult beetles that were exposed to the fungus. Carbon dioxide elevation significantly reduced B. bassiana’s germination and growth rates, but oxygen reduction did not.     

Dictyocaulus viviparus: Migration in agar of larvae subjected to a variety of physicochemical exposures

Dictyocaulus viviparus larvae were exposed to ox bile and CO₂ at intervals during their cultivation to the infective stage. Preinfective and young infective larvae were stimulated by CO₂. Bile slightly inhibited preinfective larvae, but stimulated the infective stage. Old coiled, resting infective larvae were stimulated by bile down to a concentration of 10 ppm of bile dry matter, by vertebrate biles of pig, sheep, newborn calf, cow, guinea pig, dog, and chicken, as well as by defatted bile dry matter and by glyco-, tauro-, glycodeoxy-, and taurodeoxycholates. Continuous bile exposure appeared necessary to maintain high larval activity. A high pCO₂ as well as a low redox potential potentiated the effect of bile, but had no effect alone. Exposure to pepsin-HCl and to trypsin had only a minor stimulatory effect.     

Parasitoid genus‐specific manipulation of orb‐web host spiders (Araneae,Araneidae)

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Effects of learning experience on behaviour of the generalist parasitoid Sclerodermus pupariae to novel hosts

Massicus raddei Blessig (Coleoptera: Cerambycidae), also referred to as the oak long‐horned beetle (OLB), is a non‐natural host for the generalist parasitoid Sclerodermus pupariae Yang et Yao (Hymenoptera: Bethylidae). To determine whether this generalist parasitoid might be a suitable agent for the control of OLB, the adaptive learning experience of adult female parasitoids to OLB larvae was investigated in the laboratory. A Y‐tube olfactometer bioassay was used to examine the effects of adaptive learning experience on the foraging ability of parasitoids for OLB larvae. The results indicated that parasitoids were significantly attracted by the volatiles of ash bark, Fraxinus velutina, with emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) larvae and larval frass, after exposure to ash bark mixed with EAB larval frass (learning condition A). In contrast, after exposure to oak bark, Quercus liaotungensis, mixed with OLB larval frass (learning condition C), parasitoids showed significant preference for the volatiles of oak bark with OLB larvae and larval frass. On the basis of the results of no‐choice tests, we found that parasitoids exposed to learning condition C had greater paralysis efficiency and higher OLB larvae parasitism rates than those exposed to learning condition A or no experience. Furthermore, parasitoids fed on OLB larvae in learning condition C had significantly greater paralysis efficiency and higher OLB larvae parasitism rates than other parasitoids tested. Parasitoids fed on EAB larvae in learning condition A had the lowest paralysis efficiency and OLB larvae parasitism rates among the parasitoids tested. These findings suggested that adaptive learning significantly enhanced the ability of a generalist parasitoid to utilize a novel host. This may provide a new approach to controlling non‐natural hosts using generalist parasitoids.     

The effects of enriched CO2 atmospheres on plant-insect herbivore interactions: growth responses of larvae of the specialist butterfly,Junonia coenia (Lepidoptera: Nymphalidae)

Summary Little is known about the effects of enriched CO₂ environments, which are anticipated to exist in the next century, on natural plant-insect herbivore interactions. To begin to understand such effects on insect growth and survival, I reared both early and penultimate instar larvae of the buckeye, Junonia coenia (Lepidoptera: Nymphalidae), on leaves from one of their major hostplants, plantain, Plantago lanceolata (Plantaginaceae), grown in either ambient (350 PPM) or high (700 PPM) CO₂ atmospheres. Despite consuming more foliage, early instar larvae experienced reduced growth on high CO₂-grown compared to ambient CO₂-grown leaves. However, survivorship of early instar larvae was unaffected by the CO₂ treatment. Larval weight gain was positively correlated with the nitrogen concentration of the plant material and consumption was negatively correlated with foliar nitrogen concentration, whereas neither larval weight gain nor consumption were significantly correlated with foliar water or allelochemical concentrations. In contrast, penultimate instar larvae had similar growth rates on ambient and high CO₂-grown leaves. Significantly higher consumption rates on high CO₂-grown plants enabled penultimate instar larvae to obtain similar amounts of nitrogen in both treatments. These larvae grew at similar rates on foliage from the two CO₂ treatments, despite a reduced efficiency of conversion of ingested food (ECI) on the low nitrogen, high CO₂-grown plants. However, nitrogen utilization efficiencies (NUE) were unaffected by CO₂ treatment. Again, for late instar larvae, consumption rates were negatively correlated with foliar nitrogen concentrations, and ECI was also very highly correlated with leaf nitrogen; foliar water or allelochemical concentrations did not affect either of these parameters. Differences in growth responses of early and late instar larvae to lower nitrogen, high-CO₂ grown foliage may be due to the inability of early instar larvae to efficiently process the increased flow of food through the gut caused by additional consumption of high CO₂ foliage.