Comparison between volatile emissions from transgenic apples and from two representative classically bred apple cultivars

While most risk assessments contrast a transgenic resistant to its isogenic line, an additional comparison between the transgenic line and a classically bred cultivar with the same resistance gene would be highly desirable. Our approach was to compare headspace volatiles of transgenic scab resistant apple plants with two representative cultivars (the isogenic ‘Gala’ and the scab resistance gene-containing ‘Florina’). As modifications in volatile profiles have been shown to alter plant relationships with non-target insects, we analysed headspace volatiles from apple plants subjected to different infection types by gas chromatography-mass spectrometry. Marked differences were found between healthy and leafminer (Phyllonorycter blancardella) infested genotypes, where emissions between the transgenic scab resistant line and the two cultivars differed quantitatively in four terpenes and an aromatic compound. However, these modified odour emissions were in the range of variability of the emissions recorded for the two standard cultivars that proved to be crucial references.     

Divergent responses of two cereal aphids to previous infestation of their host plant

The role of pollen odour in resource location by the pollen beetle, Meligethes aeneus (Fabricius) (Coleoptera: Nitidulidae), a pollen-feeding insect regarded as a pest of oilseed rape, Brassica napus L., (Brassicaceae) crops, was investigated in a linear track olfactometer. Both male and female beetles were attracted to the odour of whole oilseed rape flowers, indicating that these insects can locate their host plants using floral odours as cues. The attractive odour of flowers was found to emanate from all floral parts tested: the petals/sepals, the anthers, and from pollen itself. Therefore, at least part of the attractive odour of oilseed rape flowers emanates from pollen. Beetles were more attracted to floral samples containing anthers than those without anthers when these odours were directly compared in a choice-test, and this indicates that there were detectable differences between them. Anthers and pollen may therefore release distinctive odours that are quantitatively and/or qualitatively different from the odour of the rest of the flower. These experiments support the hypothesis that pollen-seeking insects use pollen odour cues to locate this food source.     

Generalism Versus Specialism: Responses of Diadegma mollipla (Holmgren) and Diadegma semiclausum (Hellen), to the Host Shift of the Diamondback Moth (Plutella xylostella L.) to Peas

A population of the diamondback moth Plutella xylostella (L.) (Lepidoptera: Plutellidae) (DBM) was recently found to infest sugar snap- and snowpeas in the Rift Valley in Kenya, causing heavy damage. The influence of this host shift on host location preferences of two parasitoids was investigated: The indigenous Diadegma mollipla (Holmgren) regarded as a relative generalist, and Diadegma semiclausum(Hellen), regarded as highly specific to DBM. The attractiveness of different odour sources was compared for the two parasitoid species using a Y-tube olfactometer using naïve females. D. mollipla was not significantly attracted to any cabbage related odours but showed a significant preference for the DBM infested pea plant when tested against clean air. D. semiclausum was highly attracted to the undamaged cabbage plant and odours related to cabbage. On the other hand, peas infested with DBM, showed no attractiveness to this parasitoid. The results showed that specialisation of D. semiclausum is mediated by host plant signals, associated with crucifers, which are not encountered in DBM feeding on peas. For D. mollipla,although a frequent parasitoid on DBM in crucifers, volatiles emitted by these plants might not be used as primary cues for host location. This species may respond largely to chemicals yet unknown and associated with a variety of plant-herbivore interactions.     

Assessing the Effects of Bt Maize on the Predatory Mite Neoseiulus cucumeris

The investigation of Neoseiulus cucumeris in the context of the ecological risk assessment of insect resistant transgenic plants is of particular interest as this omnivorous predatory mite species is commercially available and considered important for biological control. In a multitrophic feeding experiment we assessed the impact of Bt maize on the performance of N. cucumeris when offered spider mites (Tetranychus urticae) reared on Bt (Bt11, Syngenta) or non-Bt maize (near isogenic line) and Bt or non-Bt maize pollen as a food source. Various parameters including mortality, development time, oviposition rate were measured. Spider mites were used as a prey for N. cucumeris, since these herbivores are known to contain similar levels of Cry1Ab toxin, when reared on Bt maize, as those found in the transgenic leaf material. In contrast, toxin levels in pollen of this transgenic cultivar are very low. No differences in any of the parameters were found when N. cucumeris was fed with spider mites reared on Bt and non-Bt maize. Pollen was shown to be a less suitable food source for this predator as compared to spider mites. Moreover, subtle effects on female N. cucumeris (9% longer development time and 17% reduced fecundity) were measured when fed with pollen originating from Bt maize as compared to non-Bt maize pollen. Our findings indicate that the predatory mite N. cucumeris is not sensitive to the Cry1Ab toxin as no effects could be detected when offered Bt-containing spider mites, and that the effects found when fed with Bt maize pollen can be assigned to differences in nutritional quality of Bt and non-Bt maize pollen. The significance of these findings is discussed with regard to the ecological relevance for risk assessment of transgenic plants.     

Response of natural populations of predators and parasitoids to artificially induced volatile emissions in maize plants (Zea mays L.)

1 In response to herbivore attack, maize plants (Zea mays L.) emit a specific blend of induced volatiles. Artificial damage and subsequent treatment of the damaged site with caterpillar regurgitant induces the same response. The induced volatile chemicals are known to be highly attractive to several parasitoids of herbivores in laboratory bioassays, but very limited information is available on how the plant odours affect entomophagous insects in the field. 2 Experiments were conducted to determine if induced maize volatiles attract parasitic and predatory insects under field conditions and whether they affect their spatial distribution. 3 In a preliminary field experiment with blue sticky traps near treated (damaged and treated with caterpillar regurgitant) and healthy plants, more entomophagous insects (total number of parasitic Hymenoptera, Anthocoridae and Syrphidae) were trapped near treated plants than near healthy plants. 4 In a second experiment, attraction to the induced volatiles was monitored with sticky traps placed next to treated and healthy maize plants in a regular maize field. No significant differences between the two treatments were found, but significantly more insects (parasitic wasps, thrips and anthocorid bugs) were trapped near to the top of plants than on traps placed near the mid‐stem. Displacement of these insect groups within the field seemed to occur principally over the canopy, but under severe weather conditions they travelled lower in the canopy. 5 In a third experiment, the effect of induced maize odours on the spatial distribution of predators and parasitoids was investigated by placing sticky traps at different distances from healthy and treated plants. The higher catches of parasitoids near treated plants and the increased presence of these insects on the downwind side of treated plants support the notion that herbivore‐induced maize odours attract natural enemies of maize pests in the field.     

Host odours enhance the responses of adult Rhyzopertha dominica to male-produced aggregation pheromone

The behavioural responses of adult male and female Rhyzopertha dominica (Coleoptera, Bostrichidae) to blends of host volatiles and male-produced aggregation pheromone were observed in a four-arm airflow olfactometer. The odour sources used were five pheromone-releasing males each on a single maize grain (lower maize-volatiles blend), five pheromone-releasing males on 500 g of maize (higher maize-volatiles blend) or the host volatiles emanating from 500g of maize (maize volatiles alone). Multiple-choice tests, in which volatiles from all three odour sources were presented in the exposure chamber at the same time, were used to study odour preferences of the males and females. Both sexes showed strongest attraction to the higher maize volatile blend but there were significant sex differences in response to the odour sources. Males spent significantly more time than females in the zone with only maize volatiles, and females spent significantly more time in the higher maize-volatiles zone. However, when odour sources were offered singly, females gave numerically greater responses than males to all sources although this difference was statistically significant for only the lower maize-volatiles blend. As males are more attracted than females to host odours alone it is suggested that they may be more highly adapted to seek out new hosts while females are more inclined to locate a food source by following the pheromone signals produced by males. However, both sexes responded most strongly to the odour source comprising aggregation pheromone with the higher proportion of maize volatiles.     

转Bt基因玉米的生态安全性研究进展

随着转基因作物的应用和推广 ,转 Bt基因作物释放后对生态环境及其它方面产生的潜在影响越来越受到重视。分别从生物活性杀虫晶体蛋白在土壤中的残留特性、杀虫晶体蛋白对土壤中非目标生物的影响、转 Bt基因玉米植株体成分的变化、转Bt基因玉米花粉中杀虫晶体蛋白的表达特性及其在田间和马力筋叶片上的散积状况、花粉中表达的杀虫晶体蛋白对君主斑蝶的毒性、君主斑蝶幼虫暴露在 Bt花粉中的概率及综合风险评价估算等方面对转 Bt基因玉米产生的杀虫晶体蛋白与土壤生态环境的相互作用、花粉对非目标生物影响的研究现状进行了综述。通过对转 Bt基因作物生态安全性的科学评价和广泛宣传 ,以确保生物技术的健康发展。     

Responses of <Emphasis Type="Italic">Phradis</Emphasis> parasitoids to volatiles of lavender, <Emphasis Type="Italic">Lavendula angustifolia</Emphasis>—a possible repellent for their host, <Emphasis Type="Italic">Meligethes aeneus</Emphasis>

Hymenopterous parasitoids of herbivorous insects can be useful biocontrol agents in integrated pest management strategies. However, the potential effects on these beneficials of new components in such strategies are often neglected. Essential oil of lavender, Lavendula angustifolia (Miller) (Lamiaceae), has recently been identified as a potential repellent in new control strategies being developed for the pollen beetle Meligethes aeneus (Fabricius) (Coleoptera: Nitidulidae), a major pest of oilseed rape, Brassica napus L. (Brassicaceae). We tested the electrophysiological and behavioural responses of two common parasitoids of M. aeneus: Phradis interstitialis (Thomson) and P. morionellus (Holmgren) (both Ichneumonidae) using coupled gas chromatography-electroantennodetection (GC-EAD) and olfactometry techniques. Both species elicited electrophysiological responses to lavender oil volatiles, including two compounds known to be repellent to M. aeneus. However, the parasitoids gave no significant responses to the odours of lavender oil in behavioural bioassays and there was no evidence to suggest that lavender-treated oilseed rape plants would reduce host habitat location by parasitoids of the target pest.     

Host location and discrimination mediated through olfactory stimuli in two species of Encyrtidae

The role of olfactory stimuli in host detection and evaluation was studied in two encyrtid Hymenoptera. The first, Epidinocarsis lopezi De Santis, is a monophagous parasitoid of the cassava mealybug Phenacoccus manihoti Matile-Ferrero, itself feeding exclusively on cassava, Manihot esculenta Crantz. The second, Leptomastix dactylopii Howard, is a monophagous parasitoid of the Citrus mealybug, Planococcus citri Risso, but this latter is highly polyphagous. The behaviour of females of both parasitoids (attaction and locomotion) was compared in a tubular olfactometer for the odours of their respective hosts on cassava and poinsettia. Tests were made using: 1) healthy host-plant alone; 2) host-plant infested with unparasitized mealybugs; 3) unparasitized mealybugs only; 4) host-plant infested with parasitized mealybugs and 5) parasitized mealybugs only. Only E. lopezi was attracted by the odour of the host-plant alone, but both species were attracted by the odour of an infested host-plant and that of unparasitized mealybugs. The odour of parasitized mealybugs, alone or on host-plant, induced an undirected activity. The attraction of E. lopezi to the odour of the host-plant alone could be linked to the monophagous diet of its host, whereas the attraction of the two species of parasitoids to the odours of infested host-plants and unparasitized mealybugs could be due to the fact that both parasitoids are specialists. The behavioural response of both species to the odour of parasitized mealybugs revealed a new aspect in host discrimination: the identification of parasitized hosts could be partly mediated through olfactory stimuli, and not only through gustatory stimuli.     

Tritrophic Choice Experiments with Bt Plants,the Diamondback Moth (Plutella xylostella) and the parasitoid Cotesia plutellae

Parasitoids are important natural enemies of many pest species and are used extensively in biological and integrated control programmes. Crop plants transformed to express toxin genes derived from Bacillus thuringiensis (Bt) provide high levels of resistance to certain pest species, which is likely to have consequent effects on parasitoids specialising on such pests. A better understanding of the interaction between transgenic plants, pests and parasitoids is important to limit disruption of biological control and to provide background knowledge essential for implementing measures for the conservation of parasitoid populations. It is also essential for investigations into the potential role of parasitoids in delaying the build-up of Bt-resistant pest populations. The diamondback moth (Plutella xylostella), a major pest of brassica crops, is normally highly susceptible to a range of Bt toxins. However, extensive use of microbial Bt sprays has led to the selection of resistance to Bt toxins in P. xylostella. Cotesia plutellae is an important endoparasitoid of P. xylostella larvae. Although unable to survive in Bt-susceptible P. xylostella larvae on highly resistant Bt oilseed rape plants due to premature host mortality, C. plutellae is able to complete its larval development in Bt-resistant P. xylostella larvae. Experiments of parasitoid flight and foraging behaviour presented in this paper showed that adult C. plutellae females do not distinguish between Bt and wildtype oilseed rape plants, and are more attracted to Bt plants damaged by Bt-resistant hosts than by susceptible hosts. This stronger attraction to Bt plants damaged by resistant hosts was due to more extensive feeding damage. Population scale experiments with mixtures of Bt and wildtype plants demonstrated that the parasitoid is as effective in controlling Bt-resistant P. xylostella larvae on Bt plants as on wildtype plants. In these experiments equal or higher numbers of parasitoid adults emerged per transgenic as per wildtype plant. The implications for integrated pest management and the evolution of resistance to Bt in P. xylostella are discussed.     

棉铃虫成虫对两种关键花香气味的联系性学习

为了明确棉铃虫对苯乙醛和乙酸苯甲酯两种关键花香气味的联系性学习行为,在室内分别测定了不同学习训练的棉铃虫雌雄成虫对两种关键花香气味的选择偏好性反应和雄成虫的触角电位反应。偏好性测定结果表明,无花香气味接触经历的棉铃虫对苯乙醛和乙酸苯甲酯的选择频次均无显著差异,且单纯的花香气味接触经历也不能诱导棉铃虫气味偏好性的显著改变,唯有花香气味接触经历伴随蔗糖溶液进行强化训练时,才能诱导气味偏好性的显著性改变。从对两种花香气味的联系性学习看,棉铃虫更容易将苯乙醛与蔗糖溶液进行联系性学习,而不容易将乙酸苯甲酯与蔗糖溶液进行联系性学习。从性别差异看,雌虫比雄虫具有更强的将花香气味与蔗糖溶液进行联系性学习的能力。触角电位测定结果表明,不同学习训练的棉铃虫雄成虫对两种花香气味和绿叶气味顺-3-己烯-1-醇的EAG反应值之间不存在显著性差异。总之,花香气味伴随食物资源的联系性学习经历,能够明显提高棉铃虫成虫对花香气味的选择偏好,可以此为基础进一步优化花香引诱剂的配方设计和应用技术。     

High susceptibility of Bt maize to aphids enhances the performance of parasitoids of lepidopteran pests

Concerns about possible undesired environmental effects of transgenic crops have prompted numerous evaluations of such crops. So-called Bt crops receive particular attention because they carry bacteria-derived genes coding for insecticidal proteins that might negatively affect non-target arthropods. Here we show a remarkable positive effect of Bt maize on the performance of the corn leaf aphid Rhopalosiphum maidis, which in turn enhanced the performance of parasitic wasps that feed on aphid honeydew. Within five out of six pairs that were evaluated, transgenic maize lines were significantly more susceptible to aphids than their near-isogenic equivalents, with the remaining pair being equally susceptible. The aphids feed from the phloem sieve element content and analyses of this sap in selected maize lines revealed marginally, but significantly higher amino acid levels in Bt maize, which might partially explain the observed increased aphid performance. Larger colony densities of aphids on Bt plants resulted in an increased production of honeydew that can be used as food by beneficial insects. Indeed, Cotesia marginiventris, a parasitoid of lepidopteran pests, lived longer and parasitized more pest caterpillars in the presence of aphid-infested Bt maize than in the presence of aphid-infested isogenic maize. Hence, depending on aphid pest thresholds, the observed increased susceptibility of Bt maize to aphids may be either a welcome or an undesirable side effect.     

Differentiation of innate but not learnt responses to host‐habitat odours contributes to rapid host finding in a parasitoid genotype

Abstract In parasitoids, host‐habitat odour can influence host searching within the habitat. This is the case in Leptopilina sp. (Hymenoptera, Figitidae), a Drosophila parasitoid searching for larvae by ovipositor probes. This a behaviour can be conditioned to fruit odours. In a previous study, the latency of probing to a fruit odour is reported to have a genetic variability within a laboratory strain. This suggests a link between rapidity of host discovery and fitness. In the present study, this hypothesis is tested by comparing responses to host‐habitat odours between two genotypes of Leptopilina heterotoma Thomson, from the Mediterranean coast (Antibes) and from Burgundy (Tailly). The two genotypes present contrasting rhythms and levels of locomotor activity linked to contrasting interspecific competition in their area of origin. The high activity observed in the Mediterranean genotype is interpreted as an adaptive response to a limited time‐window to win against a competitor species absent in Burgundy. The present study finds differentiation in innate but not learnt responses to host‐habitat odours. The more active genotype (Antibes) has a higher probability and a shorter latency of innate probing to the odours than the less active genotype (Tailly); Antibes females also find larvae and complete infestations more rapidly. Learning equalizes the probability and the latency of probing to the odours in both strains, and increases the probing duration. Innate responses to host‐habitat odours would allow time‐limited insects to increase their reproductive rate, when host predictability is high in the habitat. Selection of faster innate responses to host and habitat cues without evolution of learnt responses indicates that the initial host discovery is more crucial to fitness than subsequent ones.     

Herbivore-induced emissions of maize volatiles repel the corn leaf aphid, shape Rhopalosiphum maidis

When maize plants, Zea mays L., are mechanically damaged and the damaged sites are treated with caterpillar regurgitant, the plants will release a specific blend of volatiles. It is known that these volatiles can be attractive to natural enemies of herbivores. We hypothesise that the plant volatiles constitute part of the induced plant defence and that herbivores will be affected by the odours as well. In laboratory and semi-field studies this hypothesis was tested for the aphid Rhopalosiphum maidis (Fitch) (Rhynchota, Sternorrhyncha, Aphididae).In a Y-tube olfactometer significantly more aphids chose the odour of healthy, undamaged maize seedlings when tested against clean air or plants treated with regurgitant. Clean air was chosen more often when tested next to the odour of treated plants. This apparently repellent effect of the odour of treated plants was significant for winged aphids, but not for the wingless aphids.In field experiments aphids were released in the centre of circles of eight potted maize plants. Four plants in each circle were damaged and treated with caterpillar regurgitant while the other plants were left unharmed. At different intervals after aphid release, the number of aphids was counted on each plant. Significantly fewer winged and wingless aphids were found back on treated plants than on healthy plants.We suggest that herbivores may be repelled by the odours because they could indicate that: 1) the plant has initiated the production of toxic compounds; 2) potential competitors are present on the plant; 3) the plant is attractive to parasitoids and predators. Aphids may be particularly sensitive to induced maize volatiles because one of the major compounds emitted by the plant is (E)--farnesene, which is a common alarm pheromone for aphids. Collections and analyses of the odours emitted by crushed R. maidis confirmed that it too emits (E)--farnesene when stressed. The results are discussed in context of plant defence strategies and their possible exploitation for the control of pest insects.     

Habitats as Complex Odour Environments: How Does Plant Diversity Affect Herbivore and Parasitoid Orientation?

Plant diversity is known to affect success of host location by pest insects, but its effect on olfactory orientation of non-pest insect species has hardly been addressed. First, we tested in laboratory experiments the hypothesis that non-host plants, which increase odour complexity in habitats, affect the host location ability of herbivores and parasitoids. Furthermore, we recorded field data of plant diversity in addition to herbivore and parasitoid abundance at 77 grassland sites in three different regions in Germany in order to elucidate whether our laboratory results reflect the field situation. As a model system we used the herb Plantago lanceolata, the herbivorous weevil Mecinus pascuorum, and its larval parasitoid Mesopolobus incultus. The laboratory bioassays revealed that both the herbivorous weevil and its larval parasitoid can locate their host plant and host via olfactory cues even in the presence of non-host odour. In a newly established two-circle olfactometer, the weeviĺs capability to detect host plant odour was not affected by odours from non-host plants. However, addition of non-host plant odours to host plant odour enhanced the weeviĺs foraging activity. The parasitoid was attracted by a combination of host plant and host volatiles in both the absence and presence of non-host plant volatiles in a Y-tube olfactometer. In dual choice tests the parasitoid preferred the blend of host plant and host volatiles over its combination with non-host plant volatiles. In the field, no indication was found that high plant diversity disturbs host (plant) location by the weevil and its parasitoid. In contrast, plant diversity was positively correlated with weevil abundance, whereas parasitoid abundance was independent of plant diversity. Therefore, we conclude that weevils and parasitoids showed the sensory capacity to successfully cope with complex vegetation odours when searching for hosts.     

A six-arm olfactometer permitting simultaneous observation of insect attraction and odour trapping

Abstract.  Behavioural assays to study insect attraction to specific odours are tedious, time consuming and often require large numbers of replications. Olfactometer and flight tunnel tests can usually only be conducted with one or two odour sources at a time. Moreover, chemical information on the odour sources has to be obtained in separate analytical studies. An olfactometer was developed in which six odours can be tested simultaneously for their relative attractiveness while during the assays, part of each test odour can be trapped for further analyses. The effectiveness of this six-arm olfactometer was tested by observing the responses of the solitary endoparasitoid Cotesia marginiventris (Cresson) to host-induced odours from young maize plants. For statistical analyses, we used log-linear models were adapted to account for overdispersion and possible positional biases. Female wasps responded extremely well in tests where they were offered a single odour source, as well as in tests with multiple choices. The responses of wasps released in groups were the same as those released individually and it was found that females did not attract or repel each other, but males preferred arms in which females had been released. Dose–response tests with varying numbers of plants or host larvae on plants revealed that the wasps responded in a dose-related manner, thus showing that the system is well suited to measure relative preference. The clear choices of the insects amongst six possibilities provided substantial statistical power. Gas chromatographic analyses of sampled air revealed clean and effective odour trapping, which largely facilitates the comparison of results from behavioural assays with the actual blends of volatiles that were emitted by the various odour sources. Advantages and disadvantages compared to other methods are discussed.     

Electrophysiological and behavioural responses of mosquitoes to volatiles of Silene otites (Caryophyllaceae)

In order to understand the biological significance of flower odour for attraction of mosquitoes, electrophysiological responses to headspace flower odour samples of Silene otites (L.) Wibel were investigated on Culex pipiens pipiens biotype molestus Forskal 1775 and Aedes aegypti L. using coupled gas chromatographic-electroantennographic detection (GC-EAD). No remarkable differences in antennal responses to the odour compounds have been found between these two mosquito species. Further, the behavioural attractiveness of the electrophysiologically active compounds, singly or as multiple odour mixtures, was evaluated with bioassay experiments with C. pipiens molestus. In bioassays, C. pipiens responded to 14 electrophysiologically active compounds in different magnitudes (65–20%) and acetophenone, linalool oxide (pyranoid), phenyl acetaldehyde and phenylethyl alcohol were found as more attractive in comparison to the least attractive compound, hexanol. In two-stimulus choice test, mosquitoes were significantly more attracted to the mixture of the four most attractive compounds compared to the mixture of all 14 compounds. The results of present study confirm that floral odours are attractive cues for mosquitoes.     

Diversity and seasonal phenology of aboveground arthropods in conventional and transgenic maize crops in Central Spain

One of the major concerns regarding the release of Bt maize is its potential negative impact on non-target organisms present in this crop. In this paper, we compare the temporal phenology and community structure of the aboveground arthropods in commercial Bt maize fields in Central Spain with those of conventional maize crops, with or without an insecticide (imidacloprid) seed treatment, over a period of three years. Spiders, harvestmen, centipedes, ground beetles, rove beetles, carrion beetles, click beetles, earwigs and damsel bugs were captured in pitfall traps every year in sufficient number to provide meaningful phenological data. One predator spider and three omnivorous species of ground beetles have been consistently present in the maize fields: Pardosa occidentalis, Poecilus cupreus, Pseudophonus rufipes and Pseudophonus griseus, respectively. Rove beetles were caught to a lesser extent, with three dominant species: Acrotona aterrima, Philonthus varians and Platystethus nitens. The variability in activity–density patterns of the aboveground fauna was mainly influenced by the year, but no detrimental effects could be attributed to Bt maize. The only exception being the changes detected in rove beetles, although these differences were transitory and varied from year to year. No changes in species richness and diversity indices for spiders and ground beetles resulted from treatments. However, imidacloprid-treated maize caused a reduction in species richness of rove beetles, even though the abundance of the main species was not reduced. Our results suggest that Bt maize could be compatible with natural enemies that are common in maize fields in Spain.     

Landscape refuges delay resistance of the European corn borer to Bt-maize: a demo-genetic dynamic model

We constructed a reaction-diffusion model of the development of resistance to transgenic insecticidal Bt crops in pest populations. Kostitzin’s demo-genetic model describes local interactions between three competing pest genotypes with alleles conferring resistance or susceptibility to transgenic plants, the spatial spread of insects being modelled by diffusion. This new approach makes it possible to combine a spatial demographic model of population dynamics with classical genetic theory. We used this model to examine the effects of pest dispersal and of the size and shape of the refuge on the efficiency of the “high-dose/refuge” strategy, which was designed to prevent the development of resistance in populations of insect pests, such as the European corn borer, Ostrinia nubilalis Hübner (Lepidoptera, Crambidae). We found that, with realistic combinations of refuge size and pest dispersal, the development of resistance could be considerably delayed. With a small to medium-sized farming area, contiguous refuge plots are more efficient than a larger number of smaller refuge patches. We also show that the formal coupling of classical Fisher–Haldane–Wright population genetics equations with diffusion terms inaccurately describes the development of resistance in a spatially heterogeneous pest population, notably overestimating the speed with which Bt resistance is selected in populations of pests targeted by Bt crops.     

Effects of Bt maize on the herbivore <Emphasis Type="Italic">Spodoptera littoralis</Emphasis> (Lepidoptera: Noctuidae) and the parasitoid <Emphasis Type="Italic">Cotesia marginiventris</Emphasis> (Hymenoptera: Braconidae).

Recent studies have shown that transgenic insect resistant plants can have negative effects on non-target herbivores as well as on beneficial insects. The study of tritrophic interactions gives insight into the complex mechanisms of food webs in the field and can easily be incorporated into a tiered risk assessment framework. We investigated the effects of transgenic maize (Zea mays) expressing insecticidal proteins derived from Bacillus thuringiensis (Bt maize) on Spodoptera littoralis, a non-target herbivore, and on the hymenopteran parasitoid Cotesia marginiventris. In a laboratory study, S. littoralis larvae were reared for their whole lifespan on a mixture of leaves and stems from 2–4-week old Bt maize plants. S. littoralis survival, developmental times and larval weights were significantly affected by Bt maize diet. However, adult moths, which survived development on Bt maize, were the same size as the adults from the control group.C. marginiventris survival, developmental times and cocoon weights were significantly negatively affected if their S. littoralis host larva had been fed Bt maize. ELISA tests confirmed that S. littoralis larvae ingest high amounts of Cry1A(b) toxin while feeding on Bt maize. In S. littoralis pupae and in C. marginiventris cocoon silk, only traces of the toxin could be detected. No toxin was found in S. littoralis and C. marginiventris adults. Thus the toxin is not accumulating in the trophic levels and in fact appears to be excreted. Our results suggest that the effects on C. marginiventris when developing in susceptible S. littoralis larvae are indirect (host mediated). The biological relevance of those results and the significance of this study in risk assessment are discussed.