High dose refuge strategies and genetically modified crops – reply to Tabashnik et al.

Cultivating non‐toxic conventional crops (refuges) in the proximity to transgenic crops that produce Bacillus thuringienesis (Bt) toxins is widely recommended to delay pest adaptation to these toxins. Using a spatially structured model of resistance evolution, Vacher and co‐workers (Vacher, C., Bourguet, D., Rousset, F., Chevillon, C. & Hochberg, M.E. 2003. J. Evol. Biol. 16 : 378–387.) show that the percentage of refuge fields required for the sustainable control of pests can be reduced through intermediate levels of refuge field aggregation and by lowering the toxin dose produced by Bt plants. Tabashnik, B.E., Gould, F. & Carrière, Y. (2004 J. Evol. Biol doi: 10.1111/j1420–9101.2004.00695.x) call into question the results of Vacher et al. (2003) concerning the effect of toxin dose. They argue that these results arise from invalid assumptions about larval concentration–mortality responses for the insect considered, the cotton pest Heliothis virescens. We show here that the models presented by Vacher et al. (2003) and Tabashnik et al. (2004) both show inaccuracies in their definitions of genotypic fitness. The level of dominance estimated by Tabashnik et al. (2004) from larval mortality rates data is irrelevant to resistance evolution, and the fitness cost of resistance evolution, and the fitness cost of resistance is inaccurately integrated into their framework. Neverthless, the comments of Tabashnik et al. (2004) are very helpful in elucidating the definitions of genotypic fitness used in Vacher et al. (2003) and in pointing out the essential factors in predicting the evolution of insect resistance to Bt transgenic crops, namely, accurate estimations of the fitness cost of resistance, of the dominance level of this cost, and of the variations in the dominance level of the advantage conferred by the resistance with Bt toxin dose.     

Effects of Rag1 aphid‐resistant soybean on mortality,development, and preference of brown marmorated stink bug

The brown marmorated stink bug, Halyomorpha halys (Stål) (Hemiptera: Pentatomidae), poses a new threat to soybean, Glycine max (L.) Merrill (Fabaceae), production in the north central USA. As H. halys continues to spread and increase in abundance in the region, the interaction between H. halys and management tactics deployed for other pests must be determined. Currently, the soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), is the most abundant and damaging insect pest of soybean in the region. Aphid‐resistant soybean, mainly with the Rag1 gene, is commercially available for management of A. glycines. Here, experiments were performed to evaluate the effects of Rag1 aphid‐resistant soybean on the mortality, development, and preference of H. halys. In a no‐choice test, mortality of H. halys reared on Rag1 aphid‐resistant soybean pods was significantly lower than when reared on aphid‐susceptible soybean pods (28 vs. 53%). Development time, adult weight, and proportion females of surviving adults did not differ when reared on Rag1 aphid‐resistant or aphid‐susceptible soybean pods. In choice tests, H. halys exhibited a preference for Rag1 aphid‐resistant over aphid‐susceptible soybean pods after 4 h, but not after 24 h. Halyomorpha halys exhibited no preference when tested with vegetative‐stage or reproductive‐stage soybean plants. The preference by H. halys for Rag1 aphid‐resistant soybean pods and the decreased mortality when reared on these pods suggests that the use of Rag1 aphid‐resistant soybean may favor this emerging pest in the north central USA.     

Spider mite infestations reduce Bacillus thuringiensis toxin concentration in corn leaves and predators avoid spider mites that have fed on Bacillus thuringiensis corn

Perceived benefits of insecticidal transgenic crops include reduced usage of broad‐based insecticides, and therefore lower risk to non‐target organisms. Numerous studies have documented low or no direct toxicity of Bacillus thuringiensis (Bt)‐derived toxins against non‐target organisms, but there has been less research on (a) effects of secondary pest infestations on Bt expressing in crops and (b) behavioural responses by predators feeding on host arthropods from Bt crops – both topics are investigated in this study. We quantified predation by the obligate spider mite predator Phytoseiulus persimilis of carmine spider mites (Tetranychus cinnabarinus), reared on Bt or non‐Bt corn (Zea mays). Both no‐choice and two‐choice studies were conducted. In addition, we quantified toxin levels in corn leaves with/without spider mite infestation. Under no‐choice conditions, P. persimilis consumed non‐Bt spider mites at a faster rate than Bt spider mites. Under two‐choice conditions, P. persimilis spent more time in the vicinity of non‐Bt spider mites than near Bt spider mites. Corn infested with spider mites exhibited lower toxin levels than non‐infested plants. These results suggest potentially complex interactions among non‐target herbivores, their natural enemies and Bt crops.     

Preference of Bt‐resistant and susceptible Busseola fusca moths and larvae for Bt and non‐Bt maize

The sustainability of genetically engineered insecticidal Bacillus thuringiensis Berliner (Bt) maize, Zea mays L. (Poaceae), is threatened by the evolution of resistance by target pest species. Several Lepidoptera species have evolved resistance to Cry proteins expressed by Bt maize over the last decade, including the African maize stem borer, Busseola fusca (Fuller) (Lepidoptera: Noctuidae). The insect resistance management (IRM) strategy (i.e., the high‐dose/refuge strategy) deployed to delay resistance evolution is grounded on certain assumptions about the biology and ecology of a pest species, for example, the interactions between the insect pest and crop plants. Should these assumptions be violated, the evolution of resistance within pest populations will be rapid. This study evaluated the assumption that B. fusca adults and larvae select and colonize maize plants at random, and do not show any preference for either Bt or non‐Bt maize. Gravid female B. fusca moths of a resistant and susceptible population were subjected to two‐choice oviposition preference tests using stems of Bt and non‐Bt maize plants. Both the number of egg batches as well as the total number of eggs laid on each stem were recorded. The feeding preference of Bt‐resistant and susceptible neonate B. fusca larvae were evaluated in choice test bioassays with whorl leaf samples of specific maize cultivars. Although no differential oviposition preference was observed for either resistant or susceptible female moths, leaf damage ratings indicated that neonate larvae were able to detect Bt toxins and that they displayed feeding avoidance behaviour on Bt maize leaf samples.     

Induced volatiles in soybean and pigeon pea plants artificially infested with the neotropical brown stink bug, Euschistus heros, and their effect on the egg parasitoid, Telenomus podisi

Herbivory is known to increase the emission of volatiles, which attract natural enemies to herbivore‐damaged plants in laboratory and agricultural systems. We report on signalling through volatiles induced by Euschistus heros (F.) (Heteroptera: Pentatomidae) in two legumes that influence the attraction and retention of the egg parasitoid Telenomus podisi (Ashmead) (Hymenoptera: Scelionidae). Air‐borne extracts obtained from two host plants of E. heros, soybean, Glycine max, and pigeon pea, Cajanus cajan (Leguminosae), produce a different blend of emitted volatiles when attacked by adult males or females and nymphs of the pest species, compared with the undamaged plants. The same results were obtained when the plants were treated with extracted saliva of E. heros which had been mechanically introduced into the plants. This indicates that some substance in the saliva contributed to the release of the volatiles. Bioassays in a Y‐tube olfactometer with female T. podisi and treated plants confirmed the significant preference of the egg parasitoid for plants attacked by either males, females, or fourth instar nymphs of the pest species. On the other hand, volatile extracts obtained from soybean subjected to the velvetbean caterpillar, Anticarsia gemmatalis, a non‐host species for T. podisi, showed a different blend of volatiles compared to those obtained from plants subjected to E. heros. Additionally, the volatiles obtained from this plant–host complex were not attractive to T. podisi. These results indicate that E. heros causes an increase in the emission of specific plant volatiles, and that the induction is possibly caused by an elicitor present in the pest saliva. The possibility that these plant volatiles play an important role in the attraction and retention of the egg parasitoid T. podisi is discussed.     

Lethal and sublethal effects of Embelia ribes and two commercial pesticides on a generalist predator

Even as new substances show promise as biopesticides for controlling pests due to their natural properties and high effectiveness in inhibiting pests, their side effects on non‐target organisms must nevertheless be evaluated before they can be included into integrated pest management systems. In this study, a crude extract from dried leaves of Embelia ribes was evaluated together with two commercial pesticides: azadirachtin (a natural product) and amitraz (a synthetic acaricide). We examined both lethal and sublethal effects on the predatory potential of the lynx spider Oxyopes lineatipes, which is among the most dominant predator in tropical agricultural agroecosystems. We found that the spider's mortality increased with rising concentration of both commercial products, azadirachtin and amitraz, but not with rising concentration of the extracts from E. ribes. The greatest mortality occurred when amitraz was used. That material caused almost 100% spider mortality in the doses recommended for field spraying. Azadirachtin significantly reduced the rate at which O. lineatipes captured prey, while there was no significant difference in capture rates among spiders exposed to a control treatment and the E. ribes treatment. Considering its absence of unfavourable impacts on O. lineatipes in terms of mortality and predatory activity, the plant extract from E. ribes shows promise as a new biopesticide material. In contrast, azadirachtin, which has been considered as safe for non‐target organisms, exhibited slight lethal effect only in higher concentrations and strong sublethal effect by reducing spiders’ predation rate.     

Susceptibility of Chrysochus auratus,a natural enemy of spreading dogbane,to insecticides used in wild blueberry production

Insect pest management in wild blueberry (Vaccinium angustifolium Aiton) usually involves insecticidal sprays, which may have detrimental effects on non‐target beneficial insects. Dogbane beetle (Chrysochus auratus Fabricius) (Coleoptera: Chrysomelidae) feeds almost exclusively on spreading dogbane (Apocynum androsaemifolium L.), an increasing weed problem in wild blueberry production. Because C. auratus is an important natural enemy of spreading dogbane, we assessed its susceptibility to several insecticides it may be exposed to during insect pest management. In laboratory bioassays, we found adult dogbane beetles were highly susceptible to field rates of phosmet (Imidan) and acetamiprid (Assail) by direct topical contact and ingestion of treated foliage, whereas no mortality was seen with spirotetramat (Movento) and chlorantraniliprole (Altacor). Topical applications of spinetoram (Delegate) did not cause significant mortality of beetles, but high mortality to beetles was found when they ingested spinetoram‐treated foliage. The results suggest that while some insecticides used in blueberry management will be hazardous to C. auratus, options are available that will cause little harm to this natural enemy.     

Spatial soil heterogeneity has a greater effect on symbiotic arbuscular mycorrhizal fungal communities and plant growth than genetic modification with Bacillus thuringiensis toxin genes

Maize, genetically modified with the insect toxin genes of Bacillus thuringiensis (Bt), is widely cultivated, yet its impacts on soil organisms are poorly understood. Arbuscular mycorrhizal fungi (AMF) form symbiotic associations with plant roots and may be uniquely sensitive to genetic changes within a plant host. In this field study, the effects of nine different lines of Bt maize and their corresponding non‐Bt parental isolines were evaluated on AMF colonization and community diversity in plant roots. Plants were harvested 60 days after sowing, and data were collected on plant growth and per cent AMF colonization of roots. AMF community composition in roots was assessed using 454 pyrosequencing of the 28S rRNA genes, and spatial variation in mycorrhizal communities within replicated experimental field plots was examined. Growth responses, per cent AMF colonization of roots and AMF community diversity in roots did not differ between Bt and non‐Bt maize, but root and shoot biomass and per cent colonization by arbuscules varied by maize cultivar. Plot identity had the most significant effect on plant growth, AMF colonization and AMF community composition in roots, indicating spatial heterogeneity in the field. Mycorrhizal fungal communities in maize roots were autocorrelated within approximately 1 m, but at greater distances, AMF community composition of roots differed between plants. Our findings indicate that spatial variation and heterogeneity in the field has a greater effect on the structure of AMF communities than host plant cultivar or modification by Bt toxin genes.     

Entomopathogenic nematodes fail to parasitize the woolly apple aphid Eriosoma lanigerum as their symbiotic bacteria are suppressed

The restriction of effective insecticides has facilitated the woolly apple aphid (WAA) Eriosoma lanigerum to become a major pest in apple orchards in Western Europe. It has also promoted alternative control strategies such as the use of entomopathogenic nematodes (EPN). We evaluated the control capacity of six commercially available EPN, viz. Heterorhabditis bacteriophora, Heterorhabditis megidis, Steinernema carpocapsae, Steinernema feltiae, Steinernema glaseri and Steinernema kraussei. We assessed the potential of these EPN to colonize and parasitize E. lanigerum in an in vitro multiwell test. Only S. carpocapsae caused higher mortality (20–40%) than the control treatment (water). However, the mortality observed with S. carpocapsae was found to be a test artefact and not induced by its specific entomopathogenic activity. A similar mortality range was recorded when applying the non‐entomopathogenic nematode Pratylenchus thornei in the same multiwell test set‐up. This result warrants careful interpretation of parasitism in these artificial test conditions. The failure of EPN activity was supported in further experiments by frequently finding S. carpocapsae inside living WAA. The presence of the EPN had no effect on aphid reproduction as numbers of ‘large’ embryos in EPN‐colonized and non‐colonized females were similar. In addition, the dauer juveniles did not recover in E. lanigerum reflecting that S. carpocapsae could not develop inside the WAA. We further demonstrated that growth of the EPN‐symbiotic bacteria Xenorhabdus nematophila and Photorhabdus luminescens is inhibited by the body fluid of the WAA, and we speculate that this antibacterial activity is the cause of the unsuccessful parasitization of the WAA by the EPN. This antibiosis inside the body of E. lanigerum would prevent production of the endotoxins by the bacterial symbionts that are essential for entomopathogenicity and insect control.     

Fitness of Bt‐resistant cabbage loopers on Bt cotton plants

Development of resistance to the insecticidal toxins from Bacillus thuringiensis (Bt) in insects is the major threat to the continued success of transgenic Bt crops in agriculture. The fitness of Bt‐resistant insects on Bt and non‐Bt plants is a key parameter that determines the development of Bt resistance in insect populations. In this study, a comprehensive analysis of the fitness of Bt‐resistant Trichoplusia ni strains on Bt cotton leaves was conducted. The Bt‐resistant T. ni strains carried two genetically independent mechanisms of resistance to Bt toxins Cry1Ac and Cry2Ab. The effects of the two resistance mechanisms, individually and in combination, on the fitness of the T. ni strains on conventional non‐Bt cotton and on transgenic Bt cotton leaves expressing a single‐toxin Cry1Ac (Bollgard I) or two Bt toxins Cry1Ac and Cry2Ab (Bollgard II) were examined. The presence of Bt toxins in plants reduced the fitness of resistant insects, indicated by decreased net reproductive rate (R₀) and intrinsic rate of increase (r). The reduction in fitness in resistant T. ni on Bollgard II leaves was greater than that on Bollgard I leaves. A 12.4‐day asynchrony of adult emergence between the susceptible T. ni grown on non‐Bt cotton leaves and the dual‐toxin‐resistant T. ni on Bollgard II leaves was observed. Therefore, multitoxin Bt plants not only reduce the probability for T. ni to develop resistance but also strongly reduce the fitness of resistant insects feeding on the plants.     

Interaction of acid exudates in chickpea with biological activity of Bacillus thuringiensis towards Helicoverpa armigera

The gram pod borer, Helicoverpa armigera, is one of the most important constraints to chickpea production. High acidity of chickpea exudates is associated with resistance to pod borer, H. armigera; however, acidic exudates in chickpea might influence the biological activity of the bacterium, Bacillus thuringiensis (Bt), applied as a foliar spray or deployed in transgenic plants for controlling H. armigera. Therefore, studies were undertaken to evaluate the biological activity of Bt towards H. armigera on chickpea genotypes with different amounts of organic acids. Significantly lower leaf feeding, larval survival and larval weights were observed on ICC 506EB, followed by C 235, and ICCV 10 across Bt concentrations. Leaf feeding by the larvae and larval survival and weights decreased with an increase in Bt concentration. However, rate of decrease in leaf feeding and larval survival and weights with an increase in Bt concentration was greater on L 550 and ICCV 10 than on the resistant check, ICC 506EB, suggesting that factors in the resistant genotypes, particularly the acid exudates, resulted in lower levels of biological activity of Bt possibly because of antifeedant effects of the acid exudates. Antifeedant effects of acid exudates reduced food consumption and hence might reduce the efficacy of Bt sprays on insect‐resistant chickpea genotypes or Bt‐transgenic chickpeas, although the combined effect of plant resistance based on organic acids, and Bt had a greater effect on survival and development of H. armigera than Bt alone.     

Impacts of Bt maize inoculated with rhizobacteria on development and food utilization of Mythimna separata

The current trend of increasing proportion of cultivation of transgenic Bt crops is pushing towards dramatic destabilization of the agroecosystem, thus raising severe concerns about the sustainability of transgenic Bt crops as an effective management tool for the control of target insect pests in the future. Rhizobacteria is the key biological regulator to ameliorate soil‐nitrogen utilization efficiency of crop plants, especially transgenic Bt crops. A laboratory study quantified the impacts of transgenic Bt maize (Line IE09S034 with Cry1Ie vs. non‐Bt maize cv. Xianyu335) inoculated with Azospirillum brasilense (AB) and Azotobacter chroococcum (AC) on the growth, development and food utilization of a target lepidopteran insect, Mythimna separata. The results showed that the inoculation of rhizobacteria significantly prolonged the larval lifespan and pupal duration, increased RCR and AD, reduced pupal weight, pupation rate, fecundity, RGR, ECD and ECI, and shortened adult longevity of M. separata fed on transgenic Bt maize, while exact opposite trends were found in these measured indexes of growth, development and food utilization for M. separata fed on non‐Bt maize inoculated with AB and AC compared with the buffer control in both years. Thus, the results clearly depicted that the inoculation of rhizobacteria had opposite influences on the growth, development and food utilization of M. separata fed on transgenic Bt maize. Presumably, rhizobacteria inoculation can be used to stimulate plant–soil‐nitrogen uptake and promote plant growth for transgenic Bt maize and non‐Bt maize, simultaneously increasing Bt toxin production and enhancing resistance efficiency against target lepidopteran pests for transgenic Bt maize.     

Mix‐planting pubescent and glabrous cassava affects abundance of Typhlodromalus aripo and its prey mite Mononychellus tanajoa

There is an increasing awareness that vegetation diversity can affect herbivore and natural enemy abundance and that plants can play a major role in directly manipulating natural enemy abundance for protection against herbivore attacks. Using data from cassava fields, we aimed at (i) testing the capacity of the predatory mite Typhlodromalus aripo to control the herbivorous mite Mononychellus tanajoa in a chemical exclusion trial; and (ii) testing, based on the differential preference by T. aripo for cassava cultivars, how combinations of two morphologically different cassava cultivars with differential suitability to the predator can improve its population densities on the non‐favourable cultivar, thereby reducing M. tanajoa densities with subsequent increases in cassava yield. The study was conducted in a cassava field in Benin, West Africa. The experiments confirmed that T. aripo effectively suppresses M. tanajoa populations on both cultivars and showed, in the no‐predator‐exclusion experiments, that cultivar combinations have significant effects on M. tanajoa and T. aripo densities. Indeed, T. aripo load on the non‐preferred cultivar was lowest in subplots where the proportion of T. aripo‐preferred cultivar was also low, while, and as expected, M. tanajoa load on the non‐preferred cultivar showed decreasing trends with increasing T. aripo densities. The possible mechanisms by which cultivar mixing could increase predator load on the non‐favourable cultivar were discussed. Our data showed that appropriate cultivar combinations effectively compensate for morphologically related differences in natural enemy abundance on a normally predator‐deficient cultivar, resulting in lower pest densities on the non‐favourable cultivar. In practical terms, this strategy could, in part, enhance adoption of cultivars that do not support sufficient levels of natural enemies for pest control.     

Colonization preference of Euschistus servus and Nezara viridula in transgenic cotton varieties,peanut, and soybean

Producers of Bt cotton, Gossypium hirsutum L. (Malvaceae), in the southeastern USA face significant losses from highly polyphagous stink bug species. These problems may be exacerbated by crop rotation practices that often result in cotton, peanut, Arachis hypogaea L., and soybean, Glycine max (L.) Merrill (both Fabaceae), growing in close proximity to one another. Because all of these crops are hosts for the major pest stink bug species in the region, we experimentally examined colonization preference of these species among the crops to clarify this aspect of their population dynamics. We planted peanut, soybean, Bt cotton, and glyphosate‐tolerant (RR) non‐Bt cotton at three sites over 3 years in replicated plots ranging from 192 to 1 323 m² and calculated odds ratios for colonization of each crop for Nezara viridula (L.) and Euschistus servus (Say) (both Hemiptera: Pentatomidae). In four of five experiments, both E. servus and N. viridula preferred soybean significantly more often than Bt cotton, non‐Bt cotton, and peanut. Neither N. viridula nor E. servus showed any preference between non‐Bt and Bt cotton in any experiment. Both species had higher numbers in Bt and non‐Bt cotton relative to peanut; this was not significant for any single experiment, but analyses across all experiments indicated that N. viridula preferred Bt and non‐Bt cotton significantly more often than peanut. Our results suggest that soybean in the landscape may function as a sink for stink bug populations relative to nearby peanut and cotton when the soybean is in the reproductive stage of development. Stink bug preference for soybean may reduce pest pressure in near‐by crops, but population increases in soybean could lead to this crop functioning as a source for later‐season pest pressure in cotton.     

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

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

Bt rice could provide ecological resistance against nontarget planthoppers

Genetically engineered (GE) rice lines expressing Lepidoptera‐active insecticidal cry genes from the bacterium Bacillus thuringiensis (Bt) have been developed in China. Field surveys indicated that Bt rice harbours fewer rice planthoppers than non‐Bt rice although planthoppers are not sensitive to the produced Bt Cry proteins. The mechanisms underlying this phenomenon remain unknown. Here, we show that the low numbers of planthoppers on Bt rice are associated with reduced caterpillar damage. In laboratory and field‐cage experiments, the rice planthopper Nilapavata lugens had no feeding preference for undamaged Bt or non‐Bt plants but exhibited a strong preference for caterpillar‐damaged plants whether Bt or non‐Bt. Under open‐field conditions, rice planthoppers were more abundant on caterpillar‐damaged non‐Bt rice than on neighbouring healthy Bt rice. GC–MS analyses showed that caterpillar damage induced the release of rice plant volatiles known to be attractive to planthoppers, and metabolome analyses revealed increased amino acid contents and reduced sterol contents known to benefit planthopper development. That Lepidoptera‐resistant Bt rice is less attractive to this important nontarget pest in the field is therefore a first example of ecological resistance of Bt plants to nontarget pests. Our findings suggest that non‐Bt rice refuges established for delaying the development of Bt resistance may also act as a trap crop for N. lugens and possibly other planthoppers.     

Impacts of nitrogen fertilizer on major insect pests and their predators in transgenic Bt rice lines T2A‐1 and T1C‐19

Nitrogen is a critical factor for plant development and nitrogen input is one of the important tactics to enhance the development and yield of crops. Nevertheless, nitrogen input could influence the occurrence of insects positively or negatively. Nitrogen is also one of the main elements composing the insecticidal crystal (Cry) protein. Cry protein production could affect nitrogen partitioning in Bt plants and as such nitrogen input may influence insect pest management in transgenic Bt rice, Oryza sativa L. (Poaceae). To test this possibility, we evaluated the impacts of nitrogen regimes on the main insect pests and their predators on two Bt rice lines, T2A‐1 and T1C‐19, expressing Cry2A and Cry1C, respectively, and their non‐transgenic parental counterpart MH63. The results showed that Cry proteins with different nitrogen regimes have enough insecticidal activity on rice leaffolder, Cnaphalocrocis medinalis Guenée (Lepidoptera: Crambidae), in both laboratory and field experiments. Laboratory studies indicated that relevant parameters of ecological fitness in brown planthopper, Nilaparvata lugens (Stål) (Hemiptera: Delphacidae), a non‐target insect pest, were significantly affected by nitrogen input both on Bt and MH63 rice lines. Nymphal survival, female adult longevity, and egg hatchability in N. lugens differed significantly among rice varieties. The experiments conducted in rice fields also demonstrated that nitrogen was positively correlated with the abundance of N. lugens on Bt rice, similar to that on MH63 rice. The abundances of two predators – the wolf spider Pirata subpiraticus (Boesenberg & Strand) (Araneae: Lycosidae) and the bug Cyrthorhinus lividipennis Reuter (Hemiptera: Miridae) – were significantly affected by rice growth stages but not by nitrogen input and rice varieties. In conclusion, the above results indicate that high nitrogen regimes for Bt rice (T2A‐1 and T1C‐19) and non‐Bt rice (MH63) cannot facilitate the management of insect pests.     

Volatiles from non‐host aromatic plants repel tea green leafhopper Empoasca vitis

The tea green leafhopper, Empoasca vitis Göthe (Hemiptera: Cicadellidae), is a serious pest of tea crops in China. The effectiveness of five aromatic non‐host plants, Corymbia citriodora (Hook.) (Myrtaceae), Ocimum basilicum L. (Lamiaceae), Lavandula pinnata L. (Lamiaceae), Ruta graveolens L. (Rutaceae), and Rosmarinus officinalis L. (Lamiaceae), was investigated to determine their ability to suppress E. vitis on tea plants. Volatile organic compounds derived from L. pinnata, R. officinalis, and C. citriodora were found to repel leafhoppers and to mask the host's odors. Intercropping L. pinnata and C. citriodora with tea plants significantly reduced the E. vitis population levels in the tea plantation associated with higher population densities of generalist predators. The volatile compounds from the five non‐host plants were collected by headspace absorption under field conditions, and the 10 major components were identified. Qualitative and quantitative differences were found among the five odor profiles. Moreover, the emission dynamics of non‐host volatiles were monitored. Non‐host volatile emissions showed two peak periods, one in the spring and one in the autumn period. These peaks were almost consistent with the population dynamics of E. vitis. Our findings suggested that this newer approach to ecologically based, sustainable pest management implemented via intercropping with non‐host aromatic plants such as L. pinnata and C. citriodora offers a promising alternative to chemical control of the leafhopper population.     

Crop dominance exerts specific effects on foliage‐dwelling arthropods in Bacillus thuringiensis cotton

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Influence of volatile compounds from herbivore‐damaged soybean plants on searching behavior of the egg parasitoid Telenomus podisi

Parasitoids use herbivore‐induced plant volatiles (HIPVs) to locate their hosts. However, there are few studies in soybean showing the mechanisms involved in the attraction of natural enemies to their hosts and prey. The objective of this study was to evaluate the influence of volatile organic compounds (VOCs) of soybean, Glycine max (L.) Merr. (Fabaceae) (cv. Dowling), that were induced after injury caused by Euschistus heros (Fabricius) (Hemiptera: Pentatomidae), on the searching behavior of the egg parasitoid Telenomus podisi Ashmead (Hymenoptera: Scelionidae). Four HIPVs from soybean, (E,E)‐α‐farnesene, methyl salicylate, (Z)‐3‐hexenyl acetate, and (E)‐2‐octen‐1‐ol, were selected, prepared from standards at various concentrations (10^?6 to 10^?1 m ), and tested individually and in combinations using a two‐choice olfactometer (type Y). Telenomus podisi displayed a preference only for (E,E)‐α‐farnesene at 10^?5 m when tested individually and compared to hexane, but they did not respond to the other compounds tested individually at any concentration or when combinations of these compounds were tested. However, the parasitoids stayed longer in the olfactometer arm with the mixture of (E,E)‐α‐farnesene + methyl salicylate at 10^?5 m than in the arm containing hexane. The results suggest that (E,E)‐α‐farnesene and methyl salicylate might help T. podisi to determine the presence of stink bugs on a plant. In addition, bioassays were conducted to compare (E,E)‐α‐farnesene vs. the volatiles emitted by undamaged and E. heros‐damaged plants, to evaluate whether (E,E)‐α‐farnesene was the main cue used by T. podisi or whether other minor compounds from the plants and/or the background might also be used to locate its host. The results suggest that minor volatile compounds from soybean plants or from its surroundings are involved in the host‐searching behavior of T. podisi.