Development of a synthetic plant volatile-based attracticide for female noctuid moths. II. Bioassays of synthetic plant volatiles as attractants for the adults of the cotton bollworm, Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

This paper reports olfactometer studies to determine the attractiveness of synthetic equivalents of plant volatiles to Helicoverpa armigera (Hübner) moths. Synthetic volatiles identified from host plants and other volatiles reported in literature as attractants to various noctuid species were tested in a two-choice olfactometer. Of 34 single chemicals tested, only seven were significantly attractive, and six were significantly repellent. However, when presented as blends of two or more volatiles, 21 of 31 blends tested were significantly attractive, and only one was significantly repellent. The most attractive blends were those containing four to six components, including aromatic volatiles primarily found in flowers, especially 2-phenylethanol and phenylacetaldehyde and volatiles found primarily in leaves, including green leaf volatiles and terpenoids. Some general principles in blending volatiles for developing attracticides for H. armigera are presented, including the need for blend complexity, the combination of volatiles from leaves and flowers, and early consideration of pragmatic factors such as price and toxicological profile.     

Development of a synthetic plant volatile-based attracticide for female noctuid moths. III. Insecticides for adult Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

We investigated the efficacy of insecticides combined with a plant volatile-based attractant for Helicoverpa armigera moths, under laboratory and field conditions. In the laboratory, 16 insecticides were assessed by the level of mortality and time to incapacitate and kill moths. The proboscis extension reflex technique was used for dosing moths. The pyrethroids, bifenthrin (only when synergised by the addition of piperonyl butoxide (PBO) but not without it) and cyfluthrin (with or without PBO), endosulfan, the carbamates methomyl and thiodicarb, and spinosad killed all moths tested at rates equivalent to, or less than, those which would be applied in cover sprays targeting larvae. The shortest time to moth incapacitation and death was observed with methomyl and thiodicarb. Spinosad produced very high mortality but moths took much longer to die. The two pyrethroids gave relatively slow kills, as did endosulfan. In a field trial, four insecticides were combined with the attractant and dead moths were collected daily from 1 to 4 days after application of the attracticide on 50 m rows of cotton. Significantly more dead moths ( H. armigera , H. punctigera and other noctuids) were found near the rows treated with attracticide containing methomyl compared with spinosad, fipronil and deltamethrin. For determining the impact of attracticides by recovering dead moths, quick acting insecticides are required to prevent moths flying away from the treated area to locations where they cannot be found. Methomyl and thiodicarb are suitable for this, but other insecticides especially spinosad could be used where quick action is not needed. Large numbers of moths were killed in the field trial, suggesting that attracticides for female Helicoverpa spp. moths could have significant impacts on local populations of these pests.     

Development of a synthetic plant volatile-based attracticide for female noctuid moths. I. Potential sources of volatiles attractive to Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

This paper is the first of a series which will describe the development of a synthetic plant volatile-based attracticide for noctuid moths. It discusses potential sources of volatiles attractive to the cotton bollworm, Helicoverpa armigera (Hübner), and an approach to the combination of these volatiles in synthetic blends. We screened a number of known host and non-host (for larval development) plants for attractiveness to unmated male and female moths of this species, using a two-choice olfactometer system. Out of 38 plants tested, 33 were significantly attractive to both sexes. There was a strong correlation between attractiveness of plants to males and females. The Australian natives, Angophora floribunda and several Eucalyptu s species were the most attractive plants. These plants have not been recorded either as larval or oviposition hosts of Helicoverpa spp., suggesting that attraction in the olfactometer might have been as nectar foraging rather than as oviposition sources. To identify potential compounds that might be useful in developing moth attractants, especially for females, collections of volatiles were made from plants that were attractive to moths in the olfactometer. Green leaf volatiles, floral volatiles, aromatic compounds, monoterpenes and sesquiterpenes were found. We propose an approach to developing synthetic attractants, here termed 'super-blending', in which compounds from all these classes, which are in common between attractive plants, might be combined in blends which do not mimic any particular attractive plant.     

Ants as egg predators of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) in Australian cotton crops

Abstract Helicoverpa armigera is a major pest of Australian cotton crops. To assess the impact of ant predation on H. armigera populations, the behaviour of four common ant taxa was observed in cotton crops in northern New South Wales over the 1999−2000 and 2001−02 seasons. Areas of cotton were artificially stocked with H. armigera eggs prior to observation. Pheidole spp. were the most frequently observed ants within the crop canopy in 1999−2000 and took the most H. armigera eggs. Iridomyrmex spp. were more frequently observed than Pheidole spp. in 2001−02 and also took some H. armigera eggs. Neither Paratrechina spp. nor Rhytidoponera metallica (Smith) took any H. armigera eggs, although both were seen in the crop canopy. Irrigation, cultivation and insecticide application disrupted foraging ants and limited their impact on H. armigera populations.     

Effects of Bt-toxin Cry1Ac on Propylaea japonica Thunberg (Col., Coccinellidae) by feeding on Bt-treated Bt-resistant Helicoverpa armigera (Hübner) (Lep., Noctuidae) larvae

Abstract:  Propylaea japonica is an important predatory insect of common cotton pests. To assess the ecological effects of transgenic Bt cotton, expressing Cry1Ac toxin, on this predator, we examined the life history parameters of P. japonica for two generations by feeding them with Bt-resistant Helicoverpa armigera . After ingesting Bt-treated Bt-resistant H. armigera larvae in the third and fourth instar, the body mass and body length of adult P. japonica decreased, a combined effect of poor prey quality and Cry1Ac Bt-toxin may account for these effects. However, larval survivorship and development in these two instars, pupal mortality, fecundity and adult longevity of P. japonica were not affected in both the generations. These results suggest that ingesting Bt-toxin Cry1Ac-treated pests in advanced larval stage might have no significant effect on the fitness of predator P. japonica .     

Oviposition and feeding avoidance in Helicoverpa armigera (Hübner) against transgenic Bt cotton

Helicoverpa armigera (Hübner), the major target pest of transgenic Bacillus thuringiensis (Bt) cotton, remains susceptible to Bt cotton in China at present. Behavioural avoidance by ovipositing females might lead to reduced exposure to Bt cotton and minimize selection for physiological resistance. We examined the behavioural responses of H. armigera to Bt and non‐Bt cottons to determine whether behavioural avoidance to Bt cotton may be present. In oviposition choice tests, the number of eggs on non‐Bt cotton plants was significantly higher than on Bt cotton plants. Similarly, in no‐choice tests, Bt cotton plants attracted significantly fewer eggs compared with non‐Bt cotton plants. H. armigera neonates showed higher dispersal and lower establishment on Bt cotton than on non‐Bt cotton. First instars were found to feed consistently on non‐Bt cotton leaves, creating large feeding holes, but only produced tiny feeding holes on Bt cotton leaves. The H. armigera population used in this study showed avoidance of oviposition and feeding on Bt cotton. Our results provide important insights into one possible mechanism underlying the durability of Bt cotton resistance and may be useful for improving strategies to sustain the effectiveness of Bt crops.     

Role of chemical cues from cotton in mediating host selection and oviposition behaviour in Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

We investigated the role of chemoreception in the host selection and oviposition behaviour of Helicoverpa armigera in the laboratory using five cotton genotypes and synthetic volatile terpenes. Female moths oviposited on substrates treated with methanol, ethanol, acetone and pentane extracts of leaves, squares and flowers of the cotton genotypes. Phytochemicals soluble in pentane were the most efficient in eliciting oviposition behaviour. In a two-way bioassay, pentane extracts of leaves or squares of a Multiple Host-plant Resistance genotype (MHR11), Deltapine commercial (DP90), and Smith Red Leaf (SRL) received significantly more eggs than solvent-treated controls. Extracts of squares of the native genotype Gossypium nelsonii did not receive more eggs. Females preferred DP90 and MHR11 to SRL and G. nelsonii. Female moths also laid more eggs on pentane extracts of MHR11 flowers than MHR11 leaves from preflowering, early flowering and peak-flowering plants. In a flight chamber, female moths used olfactory cues at short range to mediate oviposition and discrimination between host plants. Egg-laying, mated females were attracted at a distance (1.5 m) to volatile compounds released by whole plants and odours emanating from filter papers treated with synthetic volatile terpenes. Individually, the terpenes did not stimulate any significant oviposition response. However, there was a significant oviposition response to a mixture of equal volumes of the terpenes ( trans -β-caryophyllene, α-pinene, β-pinene, myrcene, β-bisabolol, and α-humulene). Conversely, antennectomised (moths with transected antennae), egg-laying, mated females did not stimulate any significant oviposition response. The significance of these findings in relation to H. armigera host-plant selection are discussed.     

Possible effect of genetically modified cotton on foraging habits of early instar Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) larvae

Abstract  We estimated survival and final location of neonate Helicoverpa armigera after 72 h on both genetically modified (GM) and conventional cotton at the squaring stage. On average, about 2.5% out of a total of 1620 initial larvae survived on two gene GM cotton (CS × 450, 289BGII®) compared with 39.5% out of a total of 810 initial larvae on Sicot 189, a conventional cotton variety. A disproportionate number of larvae were collected on squares of GM cotton, assuming they behave the same way on GM plants as on conventional plants but suffer higher rates of mortality. Our results imply that the behaviour of neonates on GM plants in terms of finding squares (and/or better survival on squares) may be contributing to the continued pest status of Helicoverpa on GM cotton from time to time. Experiments to determine if selection for changed behaviour in the field has or is occurring are urgently required.     

Biology of Campoletis chlorideae (Uchida) (Hym., Ichneumonidae) developing in Bt-treated, Bt-resistant Helicoverpa armigera (Hübner) (Lep., Noctuidae) larvae

Abstract:  Life history parameters in two generations of endoparasitoid Campoletis chlorideae (Uchida) were examined using Bacillus thuringiensis (Bt)-resistant Helicoverpa armigera (Hübner) larvae feeding on B. thuringiensis toxin Cry1Ac. In the laboratory, Bt toxin was fed to Bt-resistant host larvae continuously in case of Bt treatment and only before or after the host larvae were parasitized in Bt–P and P–Bt treatments, respectively. C. chlorideae pupae developed faster in Bt treatment than non-Bt treatment. The shortened pupal stage duration was mainly because of the feeding of host larvae on Bt-diet before being parasitized. Body length of adult male C. chlorideae developed inside Bt-treated Bt-resistant (Bt–Bt) H. armigera larvae significantly decreased, especially in host larvae feeding on Bt-diet after being parasitized. However, survival, pupal mortality and adult longevity of C. chlorideae were almost unaffected in Bt-resistant H. armigera larvae feeding on Bt-toxin. Furthermore, Bt-treated host larvae had the same effect on the F₁ progeny of C. chlorideae as the previous generation, and there was no significant difference between generations. This experiment suggests that there is very limited effect on the life history parameters in two generations of C. chlorideae parasitizing Bt–Bt H. armigera larvae. But both generations of C. chlorideae are affected when Bt-resistant H. armigera larvae fed on Bt toxin for different durations.     

棉花单宁-黄酮类化合物对棉铃虫的抗性潜力

陆地棉棉叶甲醇提取物包含了棉花的大部分单宁-黄酮类化合物。这些化合物对棉花多种病虫害具有抗性作用。将这些化合物定量加入人工饲料中饲养棉铃虫龄幼虫5d,结果表明,培儿茶素、缩合单宁、芸香苷和异槲皮苷在饲料中的浓度增加均与幼虫体重生长呈显著负相关,ED50分别为0．81％,0．49％,0．57％和0．83％;儿茶素则不存在这种关系;用含0．8％的浓度桔儿茶素、缩合单宁,芸香苷和异槲皮苷的饲料测试棉铃虫,发现随取食量的增加,相对生长率的增加显著低于对照(不含次牛物质),即这些物质属于慢毒剂,而儿茶素仅有阻食作用。     

Relationship between oviposition preference and offspring performance in Australian Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

Abstract We investigated the oviposition preference and larval performance of Helicoverpa armigera under laboratory conditions to determine if the oviposition preference of individual females on maize, cowpea and cotton correlates with offspring performance on the leaves of the same host plants. The host-plant preference hierarchy of females did not correlate with their offspring performance. Female moths chose host plants that contributed less to their offspring fitness. Plant effects accounted for the largest amount of variation in offspring performance, while the effects of female (family) was low. The offspring of most females (80%, n  = 10) were broadly similar, but 20% (two out of 10), showed marked difference in their offspring performance across the host-plant species. Similarly, there was no relation between larval feeding preference and performance. However, like most laboratory experiments, our experi-mental design does not allow the evaluation of ecological factors (for example, natural enemies, host abundance, etc.) that can play an important role in larval performance in the field. Overall, the results highlight the importance of carrying out preference performance analysis on the individual or family level, rather than pooling individuals to obtain average population data.     

中红侧沟茧蜂对不同龄期棉铃虫幼虫及其为害棉株的趋性反应

研究中红侧沟茧蜂Microplitis mediator Haliday对棉铃虫Helicoverpa armigera Hübner幼虫及其为害棉株的趋性行为, 可为高效应用中红侧沟茧蜂控制棉铃虫提供理论依据。本研究在实验室内以中红侧沟茧蜂雌蜂、1~5龄棉铃虫幼虫及其为害棉花植株（去虫或未去虫）为试验材料, 采用“Y”型昆虫嗅觉仪测定了中红侧沟茧蜂雌蜂对1~5龄棉铃虫幼虫及其为害棉株的趋性行为反应。结果表明: 中红侧沟茧蜂对3龄、4龄和5龄棉铃虫幼虫均没有明显的趋向行为反应, 1龄和2龄棉铃虫幼虫对中红侧沟茧蜂有显著的引诱作用。此外, 1~5龄棉铃虫幼虫为害后去虫棉株与健康棉株均对中红侧沟茧蜂有吸引作用。在未去虫的处理中, 虫害诱导棉株比健康植株更能吸引中红侧沟茧蜂, 与健康植株差异显著, 而且不同龄期棉铃虫幼虫为害处理间差异较大, 对中红侧沟茧蜂引诱作用的相对顺序为: 5龄为害植株>2龄和4龄为害植株>1龄和3龄为害植株 (2龄和4龄为害处理间差异不显著; 1龄和3龄为害处理间差异不显著)。本研究结果可为中红侧沟茧蜂在生物防治上的应用提供一定的理论指导和实践依据。