Whereas considerable effort has been afforded in understanding the properties of galaxies, a full physical picture, connecting their baryonic and dark-matter content, super-massive black holes, and (metric) theories of gravity, is still ill-defined. Strong gravitational lensing furnishes a powerful method to probe gravity in the central regions of galaxies. It can (1) provide a unique detection-channel of dark-matter substructure beyond the local galaxy group, (2) constrain dark-matter physics, complementary to direct-detection experiments, as well as metric theories of gravity, (3) probe central super-massive black holes, and (4) provide crucial insight into galaxy formation processes from the dark matter point of view, independently of the nature and state of dark matter. To seriously address the above questions, a considerable increase in the number of strong gravitational-lens systems is required. In the timeframe 2010-2020, a staged approach with radio (e.g. EVLA, e-MERLIN, LOFAR, SKA phase-I) and optical (e.g. LSST and JDEM) instruments can provide 10^(2-4) new lenses, and up to 10^(4-6) new lens systems from SKA/LSST/JDEM all-sky surveys around ~2020. Follow-up imaging of (radio) lenses is necessary with moderate ground/space-based optical-IR telescopes and with 30-50m telescopes for spectroscopy (e.g. TMT, GMT, ELT). To answer these fundamental questions through strong gravitational lensing, a strong investment in large radio and optical-IR facilities is therefore critical in the coming decade. In particular, only large-scale radio lens surveys (e.g. with SKA) provide the large numbers of high-resolution and high-fidelity images of lenses needed for SMBH and flux-ratio anomaly studies.
1. To assess the risks of an insect-resistant transgenic plant for non-target arthropods, it is important to investigate the exposure of non-target species to the transgene product. Exposure of predators in the field depends on the toxin levels in food sources, their feeding ecology and that of their prey.2. To verify the transmission of Cry1Ab toxin through the food chain, and thus exposure of predators in the field, samples from different plant tissues, herbivores, and predators in Bt maize fields in Spain (Event 176) were collected at different periods over the season and the toxin content was measured using ELISA. Complementary laboratory studies were performed with the omnivorous predator Orius majusculus to assess the toxin uptake and persistence after feeding on variable Bt-containing food sources.3. Field results revealed that toxin content in some herbivores was negligible (aphids, thrips, leafhoppers) compared with those in spider mites. The latter herbivore only occurred after pollen shed and contained three times greater toxin levels than Bt maize leaves.4. Data confirmed that the Bt toxin can be transferred to predators, that is to say to Orius spp., Chrysoperla spp., and Stethorus sp. This only applied when Bt maize pollen or spider mites were available. The passage of Bt toxin to O. majusculus via these two food sources was also confirmed in the laboratory. Contrastingly, some predators in the field (hemerobiids, Nabis sp., Hippodamia sp., Demetrias sp.) contained no or negligible toxin levels even when pollen or spider mites were present.5. Besides essential information for exposure assessment of numerous arthropod predators, this study provides an insight into the feeding ecology of different arthropods in the maize system.
A major concern regarding the deployment of insect resistant transgenic plants is their potential impact on non-target organisms, in particular on beneficial arthropods such as predators. To assess the risks that transgenic plants pose to predators, various experimental testing systems can be used. When using tritrophic studies, it is important to verify the actual exposure of the predator, i.e., the presence of biologically active toxin in the herbivorous arthropod (prey). We therefore investigated the uptake of Cry1Ab toxin by larvae of the green lacewing (Chrysoperla carnea (Stephens); Neuroptera: Chrysopidae) after consuming two Bt maize-fed herbivores (Tetranychus urticae Koch; Acarina: Tetranychidae and Spodoptera littoralis (Boisduval); Lepidoptera: Noctuidae) by means of an immunological test (ELISA) and the activity of the Cry1Ab toxin following ingestion by the herbivores. Moreover, we compared the activity of Cry1Ab toxin produced by Bt maize to that of purified toxin obtained from transformed Escherichia coli, which is recommended to be used in toxicity studies. The activity of the toxin was assessed by performing feeding bioassays with larvae of the European corn borer (Ostrinia nubilalis (Hübner); Lepidoptera: Crambidae), the target pest of Cry1Ab expressing maize. ELISA confirmed the ingestion of Bt toxin by C. carnea larvae when fed with either of the two prey species and feeding bioassays using the target pest showed that the biological activity of the Cry1Ab toxin is maintained after ingestion by both herbivore species. These findings are discussed in the context of previous risk assessment studies with C. carnea. The purified Cry1Ab protein was more toxic to O. nubilalis compared to the plant-derived Cry1Ab toxin when applied at equal concentrations according to ELISA measurements. Possible reasons for these findings are discussed.
As a part of a risk assessment procedure, the impact of Bt maize expressing Cry1Ab toxin on the thrips Frankliniella tenuicornis (Uzel) (Thysanoptera: Thripidae) was investigated, and the potential risks for predators feeding on thrips on Bt maize were evaluated. The effects of Bt maize on F. tenuicornis were assessed by measuring life‐table parameters when reared on Bt and non‐Bt maize. The content of Cry1Ab toxin in different stages of F. tenuicornis reared on Bt maize and the persistence of the toxin in adults where determined in order to evaluate the possible exposure of predators when feeding on thrips. In addition, Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae) was used as a model predator to assess how the behaviour of prey and predator may influence the exposure of a natural enemy to the Bt toxin. Life‐table parameter results showed that F. tenuicornis was not affected when it was reared on Bt maize. This indicates that the potential for prey quality‐mediated effects on predators is low. Bt content was highest in thrips larvae and adults, and negligible in the non‐feeding prepupal and pupal stages. The persistence of the Cry1Ab toxin in adult F. tenuicornis was short, resulting in a decrease of 97% within the first 24 h. Predation success by young C. carnea larvae varied among the thrips stages, indicating that exposure of predators to Bt toxin can additionally depend on the prey stage. When combining the current knowledge of the susceptibility of major thrips predators with our findings showing no potential for prey quality‐mediated effects, relatively low toxin content in thrips as well as short persistence, it can be concluded that the risks for predators when feeding on thrips in or next to Bt maize fields are negligible.
Various studies have been conducted to assess the damage caused by secondary lepidopteran pests to transgenic Bt maize expressing Cry1Ab. However, to date little is known on the effects of transgenic maize on Spodoptera littoralis (Boisduval) (Lepidoptera: Noctuidae), a polyphagous herbivore which is considered a pest in Mediterranean maize growing areas. Here we present results on the effects of Bt maize (Bt-11) and Bt spray (Dipel) on the various life stage parameters of this herbivore. We further assess the expression of Cry1Ab in different leaves and leaf parts in maize at a given plant growth stage, and determine whether the feeding damage of 3rd instar S. littoralis is influenced by Bt toxin expression. Contrary to previous literature reporting that S. littoralis is not sensitive to Bt Cry1Ab toxin, our results show that insects fed on either transgenic or Bt sprayed plants were negatively affected. Young S. littoralis larvae (1st and 2nd instars) were found to be the most sensitive to the Bt toxin. This was represented by a higher mortality and a slower developmental time of larvae maintained on transgenic or sprayed plants when compared to insects maintained on control plants. Moreover, Bt maize had a stronger and prolonged detrimental effect on insects when compared to Bt spray in maize. This was revealed by the fact that insects maintained on transgenic plants from 3rd instar to pupation took longer to reach adult emergence compared to insects that were maintained on sprayed plants. This was likely due to the continuous exposure of insects to the toxin when kept on transgenic maize. ELISA results showed a variation in the amount of Bt toxin among different leaf sections in transgenic maize at a given plant growth stage. These differences in Bt toxin were primarily found in the youngest leaf of growing plants. Although the lowest amounts of Bt toxin were detected in the growing leaf section of young leaves, this difference did not appear to influence the feeding behavior of 3rd instar S. littoralis.
Earlier studies have shown that larvae of the green lacewing predator Chrysoperla carnea are negatively affected when preying on lepidopteran larvae that had been fed with transgenic maize expressing the cry1Ab gene from Bacillus thuringiensis. To test whether the observed effects were directly caused by the Cry1Ab toxin, we have developed a bioassay which allows us to feed high concentrations of the toxin directly to the predator. The results of these feeding studies show no direct toxic effect of Cry1Ab on C. carnea larvae. The amount of toxin ingested by first instar C. carnea in the present study was found to be a factor 10,000 higher than the concentration ingested when feeding on Bt-reared lepidopteran larvae, a treatment that was previously shown to have a negative impact on the predator. In addition, feeding first instar C. carnea with the Cry1Ab toxin did not affect the utilisation of subsequently provided prey. Furthermore, the quality of the prey provided to first instars did not affect the sensitivity of second and third instar C. carnea to the Bt-toxin. The presented results strongly suggest that C. carnea larvae are not sensitive to Cry1Ab and that earlier reported negative effects of Bt-maize were prey-quality mediated rather than direct toxic effects. These results, together with the fact that lepidopteran larvae are not regarded as an important prey for C. carnea in the field, led us to conclude that transgenic maize expressing Cry1Ab poses a negligible risk for this predator.
Bacillus thuringiensis (Bt) bioinsecticides are generally considered safe to beneficial insects. However, negative effects of transgenic Bt-expressing maize on an important predator, Chrysoperla carnea, have previously been reported. Here, we make an ecological assessment of the effects of Dipel, one of the most widely used Bt-sprays in agriculture, on C. carnea larvae. Indirect effects due to a reduction of prey were tested by rearing three prey species of C. carnea (the aphid Rhopalosiphum padi, the spider mite Tetranychus urticae, and Lepidoptera larva Spodoptera littoralis) on either maize plants sprayed with Dipel (at the recommended field concentrations) or on control plants. Effects of Dipel on C. carnea were assessed by performing greenhouse experiments in which chrysopid larvae were kept on Bt-sprayed or control plants and fed with herbivores reared on Bt-sprayed or control plants. Dipel had no effect on aphids; however, negative effects on spider mites were observed. Spider mites reared on Bt-sprayed plants had a significantly lower intrinsic rate of natural increase compared to those reared on control plants. Similarly, S. littoralis larvae were significantly affected by Dipel as the developmental time required by larvae which were fed Bt-sprayed plants was prolonged when compared to larvae on untreated plants. Negative effects on C. carnea larvae were also shown through prey-mediated exposure to Dipel. A significant increase in mortality, a prolonged developmental time and a slight decrease in weight was observed for C. carnea fed with ‘Bt-contaminated’ S. littoralis larvae. The effects of Dipel on C. carnea larvae shown in this study, are comparable to earlier published results obtained with Bt maize. The ecological relevance of these results is discussed in comparison with the possible risks that the deployment of Bt-expressing maize plants pose to the predator C. carnea.
1. Chrysoperla carnea is an important predatory insect in maize. To assess the ecological effects of Bt-maize, expressing the Cry1Ab protein, on larvae of this predator, the following factors were examined: (1) the performance of three prey herbivores (Rhopalosiphum padi, Tetranychus urticae, and Spodoptera littoralis) on transgenic Bt and non-transgenic maize plants; (2) the intake of the Cry1Ab toxin by the three herbivores; and (3) the effects on C. carnea when fed each of the prey species. 2. The intrinsic rate of natural increase (r(m)) was used as a measure of performance for R. padi and T. urticae. No difference in this parameter was observed between herbivores reared on Bt or non-transgenic plants. In contrast, a higher mortality rate and a delay in development were observed in S. littoralis larvae when fed Bt-maize compared with those fed the control maize plants. 3. The ingestion of Cry1Ab toxin by the different herbivores was measured using an immunological assay (ELISA). Highest amounts of Cry1Ab toxin were detected in T. urticae, followed by S. littoralis, and only trace amounts detected in R. padi. 4. Feeding C. carnea with T. urticae, which were shown to contain the Cry1Ab toxin, or with R. padi, which do not ingest the toxin, did not affect survival, development, or weight of C. carnea. In contrast, a significant increase in mortality and a delay in development were observed when predators were fed S. littoralis larvae reared on Bt-maize. 5. A combined interaction of poor prey quality and Cry1Ab toxin may account for the negative effects observed on C. carnea when fed S. littoralis. The relevance of these findings to the ecological risks of Bt-maize on C. carnea is discussed.
The relationship between host stage selection and foraging behaviour of Pholetesor bicolor Nees (Hymenoptera: Braconidae), a larval parasitoid of Phyllonorycter spp. (Lepidoptera: Gracillariidae), was investigated under laboratory conditions. The endophytic host develops through two larval stages with different feeding habits, accordingly named sap‐ and tissue‐feeders. The parasitoid was able to find and parasitise both larval stages, even though it is most successful in parasitising the sap‐feeder stage. The influence of experience in the parasitoid's searching behaviour was observed in a choice bioassay. Searching activity increased when either contact experience with the sap‐ or the tissue‐feeder host was given. Furthermore, the ability of the parasitoid to locate a sap‐ or a tissue‐feeder infested plant was influenced by the type of experience given prior to the bioassay. Naive females were less active, and were observed with equal frequency on sap‐feeder, tissue‐feeder and non‐infested plants. In contrast, females that were given previous contact experience with sap‐feeders (i.e., the host stage which provided the most successful parasitism) were observed foraging more often on plants infested by the sap‐feeders, than on those infested by tissue‐feeders or on non‐infested plants. Experience with a tissue‐feeder host had no detectable effect on host stage location and only enhanced P. bicolor's foraging activity. The advantages of learning in this tritrophic system are discussed.
Field parasitism of the egg parasitoid Trichogramma brassicae Bezdenko (Hymenoptera, Trichogrammatidae) (synonymous to T. maidis Pint. et Voegele) on Ostrinia nubilalis Hübner was compared to four single quality parameters (walking speed, fecundity on the factitious host Ephestia keuhniella Zeller, fecundity on the natural host O. nubilalis, and life span) previously measured in the laboratory and, a quality index calculated from three of the measured parameters. A single quality parameter (fecundity on E. keuhniella) and the calculated quality index showed a correlation to T. brassicae field parasitism, for different T. brassicae populations. The number of female Trichogramma released clearly influenced parasitism but not in the same proportion for all populations tested, reflecting that numbers released can only to some extent compensate for low quality in Trichogramma. Variation indegree hours above 18 °C also influenced field parasitism. When incorporating degree hours to the quality index a significant correlation R2 = 0.56 (P = 0.01) is obtained. In this work fecundity of T. brassicae on E. keuhniella can be as good an indicator for the potential field performance of T. brassicae, as the quality index which takes into account three quality parameters. Since environmental factors can obscure the potential performance of a population, i.e., one which under optimal laboratory conditions performs well, the quality index gives only a partial indication of how the released strains will perform in the field.
A laboratory and a field test for flight initiation ofTrichogramma brassicae Bezdenko (Hymenoptera, Trichogrammatidae) (synonymous toT. maidis Pintureau et Voegele) were developed with the aim to establish a simple, cheap and quick flight quality control method forTrichogramma producers. The flight quality of four strains ofT. brassicae reared onEphestia kuehniella Zeller eggs were compared. The material tested consisted of four strains: two strains reared for two (F2) and 39 to 42 (F39–42) generations onE. kuehniella eggs without storage treatment, a diapause strain reared six generations (F6) onE. kuehniella eggs and a commercial strain also reared onE. kuehniella eggs whose production and storage conditions were unknown. Clear differences in flight activity among strains were observed. Both, the F2 and commercial strain showed significantly better flight activity under laboratory conditions compared to the other strains. Flight field cage experiments were made for comparison between field and laboratory results. Similar differences among strains in field cage experiments were observed when compared to laboratory tests.