Role of chemical cues from cotton in mediating host selection and oviposition behaviour in Helicoverpa armigera (Hubner) (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-beta-caryophyllene, alpha-pinene, beta-pinene, myrcene, beta-bisabolol, and alpha-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 hostplant selection are discussed.

Ecology and behavior of first instar larval Lepidoptera

Neonate Lepidoptera are confronted with the daunting task of establishing themselves on a food plant. The factors relevant to this process need to be considered at spatial and temporal scales relevant to the larva and not the investigator. Neonates have to cope with an array of plant surface characters as well as internal characters once the integument is ruptured. These characters, as well as microclimatic conditions, vary within and between plant modules and interact with larval feeding requirements, strongly affecting movement behavior, which may be extensive even for such small organisms. In addition to these factors, there is an array of predators, pathogens, and parasitoids with which first instars must contend. Not surprisingly, mortality in neonates is high but can vary widely. Experimental and manipulative studies, as well as detailed observations of the animal, are vital if the subtle interaction of factors responsible for this high and variable mortality are to be understood. These studies are essential for an understanding of theories linking female oviposition behavior with larval survival, plant defense theory, and population dynamics, as well as modern crop resistance breeding programs.

Intra-specific variation for host plant use in Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae): Implications for management

The polyphagous moth Helicoverpa armigera (Hubner) is one of the world's most important agricultural pests. A number of existing approaches and future designs for management of H. armigera rely on the assumption that moths do not exhibit either genetically and/or non-genetically based variation for host plant utilization. We review recent empirical evidence demonstrating that both these forms of variation influence host plant use in this moth. The significance of this variation in H. armigera in relation to current and future pest management strategies is examined. We provide recommendations on future research needs and directions for sustainable management of H. armigera, under a framework that includes consideration of intra.-specific variation for host use relevant in this and other similar pest species. (C) 2004 Elsevier Ltd. All rights reserved.

Discrepancy between antennal and behavioural responses for enantiomers of -pinene: electrophysiology and behavior of Helicoverpa armigera (Lepidoptera)

The ability of adult cotton bollworm, Helicoverpa armigera (Hubner), to distinguish and respond to enantiomers of alpha-pinene was investigated with electrophysiological and behavioral methods. Electroantennogram recordings using mixtures of the enantiomers at saturating dose levels, and single unit electrophysiology, indicated that the two forms were detected by the same receptor neurons. The relative size of the electroantennogram response was higher for the (-) compared to the (+) form, indicating greater affinity for the (-) form at the level of the dendrites. Behavioral assays investigated the ability of moths to discriminate between, and respond to the (+) and (-) forms of alpha-pinene. Moths with no odor conditioning showed an innate preference for (+)-alpha-pinene. This preference displayed by naive moths was not significantly different from the preferences of moths conditioned on (+)-alpha-pinene. However, we found a significant difference in preference between moths conditioned on the (-) enantiomer compared to naive moths and moths conditioned on (+)-alpha-pinene, showing that learning plays an important role in the behavioral response. Moths are less able to distinguish between enantiomers of alpha-pinene than different odors (e.g., phenylacetaldehyde versus (-)-alpha-pinene) in learning experiments. The relevance of receptor discrimination of enantiomers and learning ability of the moths in host plant choice is discussed.

Replication of Heliothis virescens ascovirus in insect cell lines

Ascoviruses (AVs) infect larvae of various insect pests belonging to the family Noctuidae. The result of AV infection in the hosts is cleavage of infected cells into vesicles, a unique feature of AV infection. Since insect cell lines facilitate the study of virus life cycles, attempts were made to analyze Heliothis virescens AV (HvAV3e) infection in several cell lines and compare cell pathology to larval infection. In this study, replication and cytopathological effects of HvAV3e on four different cell lines were investigated. HvAV3e replication was confirmed in three noctuid cell lines from Spodoptera frugiperda (Sf9) and Helicoverpa zea (BCIRL-Hz-AM1 and FB33). However, the virus did not replicate in the non-noctuid insect cell line from Pieris rapae (Pieridae). Despite replication of the virus in the three permissive cell lines, the cytopathological effects of the virus were significantly different from that of larval infection.

Some factors affecting white stem borer Scirpophaga innotata (Walker) (Lepidoptera : Pyralidae) injury to rice

The amount of injury to rice caused by white stem borer Sciryophaga innotata depends on cultivar, and stage of plant and insect development, as well as insect abundance. Of the cultivars tested, IR64, IR42, Cisadane and Ketan. IR64 were the most susceptible and Ketan the least susceptible to feeding damage. Third and fourth instars consumed more stem dry matter than other stages, although yield reduction depended on the number of tillers injured. On the wider stemmed Ketan, fewer tillers were injured than the narrower IR64. Larvae are more likely to move among tillers in the third instar stage, which tends to coincide with maximum tillering and may result in more tillers injured and in yield reduction. Later instar larvae burrow downwards to the internode where they pupate. Larvae appear to move less among tillers in 'resistant' cultivars. Management strategies should target this pest at third instar and when its abundance in the field warrants control. Fewer than 10% of the neonates establish successfully on stems, and this mortality needs to be taken into account when deciding on control, as does the ability of rice plants to compensate for injury. (C) 1998 Published by Elsevier Science Ltd. All rights reserved.

Effects of egg load on the host-selection behaviour of Helicoverpa armigera (Hubner) (Lepidoptera : Noctuidae)

We examined the effect of age-specific fecundity, mated status, and egg load on host-plant selection, by Helicoverpa armigera under laboratory conditions. The physiological state of a female moth (number of mature eggs produced) greatly influences her host-plant specificity and propensity to oviposit (oviposition motivation). Female moths were less discriminating against cowpea (a low-ranked host) relative to maize (a high-ranked host) as egg load increased. Similarly, increased egg load led to a greater propensity to oviposit on both cowpea and maize. Distribution of oviposition with age of mated females peaked shortly after mating and declined steadily thereafter until death. Most mated females (88%) carried only a single spermatophore, a few females (12%) contained two. The significance of these findings in relation to host-plant selection by H. armigera, and its management, are discussed.

Demonstration of an ecdysis-triggering compound in the epitracheal gland of the cotton bollworm, Helicoverpa armigera (Hubner) (Lepidoptera : Noctuidae)

The recent report of an ecdysis-triggering hormone (ETH) in the tobacco hornworm Manduca sexta and several other Lepidoptera prompted the search for the homologous hormone in the pest noctuid, Helicoverpa armigera. In M. sexta, ETH is produced in a large cell that forms part of a three-cell epitracheal gland complex found near each of the the larval and pupal spiracles. The homologous glands were found in H. armigera and an ecdysis-triggering function of the gland contents was confirmed by the induction of premature ecdysis after injection of a crude gland extract into pupae.

Size of the first spring generation of Helicoverpa punctigera (Wallengren) (Lepidoptera : Noctuidae) and winter rain in central Australia

Serious infestations of Helicoverpa punctigera are experienced yearly in the eastern cropping regions of Australia. Regression analysis was used to determine whether the size of the first generation in spring (G(1)), which is comprised mostly of immigrants from inland Australia, was related to monthly rainfall in inland winter breeding areas. Data from two long series of light-trap catches at Narrabri in New South Wales (NSW) and Turretfield in South Australia (SA) were used in the analyses. The size of G1 at Narrabri in each year was significantly regressed on the amount of rainfall in western Queensland and NSW in May and June. The size of G1 at Turretfield each year was significantly regressed on the amount of rain in May, June and July in western Queensland and NSW and also in the desert of central Western Australia. Low r(2) values of the regressions suggest that rainfall data for more sites, as well as biological and other physical factors, such as temperature, evaporation, and prevailing wind systems, may need to be included to improve forecasts of the potential magnitude of the infestations in coastal cropping regions.

Life tables of the processionary caterpillar Ochrogaster lunifer Herrich-Schaffer (Lepidoptera : Thaumetopoeidae) at local and regional scales

A major ongoing debate in population ecology has surrounded the causative factors underlying the abundance of phytophagous insects and whether or not these factors limit or regulate herbivore populations. However, it is often difficult to identify mortality agents in census data, and their distribution and relative importance across large spatial scales are rarely understood. Were, we present life tables for egg batches and larval cohorts of the processionary caterpillar Ochrogaster lunifer Herrich-Schaffer, using intensive local sampling combined with extensive regional monitoring to ascertain the relative importance of different mortality factors at different localities. Extinction of entire cohorts (representing the entire reproductive output of one female) at natural localities was high, with 82% of the initial 492 cohorts going extinct. Mortality was highest in the egg and early instar stages due to predation from dermestid beetles, and while different mortality factors (e.g. hatching failure, egg parasitism and failure to establish on the host) were present at many localities, dermestid predation, either directly observed or inferred from indirect evidence, was the dominant mortality factor at 89% of localities surveyed. Predation was significantly higher in plantations than in natural habitats. The second most important mortality factor was resource depletion, with 14 cohorts defoliating their hosts. Egg and larval parasitism were not major mortality agents. A combination of predation and resource depletion consistently accounted for the majority of mortality across localities, suggesting that both factors are important in limiting population abundance. This evidence shows that O. lunifer is not regulated by natural enemies alone, but that resource patches (Acacia trees) ultimately, and frequently, act together to limit population growth.

Analysis of long-term light-trap data for Helicoverpa spp. (Lepidoptera : Noctuidae) in Australia: the effect of climate and crop host plants

Regression analyses of a long series of light-trap catches at Narrabri, Australia, were used to describe the seasonal dynamics of Helicoverpa armigera (Hubner). The size of the second generation was significantly related to the size of the first generation, to winter rainfall, which had a positive effect, and to spring rainfall which had a negative effect. These variables accounted for up to 96% of the variation in size of the second generation from year to year. Rainfall and crop hosts were also important for the size of the third generation. The area and tonnage of many potential host crops were significantly correlated with winter rain. When winter rain was omitted from the analysis, the sizes of both the second and third generations could be expressed as a function of the size of the previous generation and of the areas planted to lucerne, sorghum and maize. Lucerne and maize always had positive coefficients and sorghum a negative one. We extended our analysis to catches of H. punctigera (Wallengren), which declines in abundance after the second generation. Winter rain had a positive effect on the sizes of the second and third generations, and rain in spring or early summer had a negative effect. Only the area grown to lucerne had a positive effect on abundance. Forecasts of pest levels from a few months to a few weeks in advance are discussed, along with the improved understanding of the seasonal dynamics of both species and the significance of crops in the management of insecticide resistance for H. armigera.

Long range forecasts of the numbers of Helicoverpa punctigera and H-armigera (Lepidoptera : Noctuidae) in Australia using the Southern Oscillation Index and the Sea Surface Temperature

The use of long-term forecasts of pest pressure is central to better pest management. We relate the Southern Oscillation Index (SOI) and the Sea Surface Temperature (SST) to long-term light-trap catches of the two key moth pests of Australian agriculture, Helicoverpa punctigera (Wallengren) and H. armigera (Hubner), at Narrabri, New South Wales over 11 years, and for H. punctigera only at Turretfield, South Australia over 22 years. At Narrabri, the size of the first spring generation of both species was significantly correlated with the SOI in certain months, sometimes up to 15 months before the date of trapping. Differences in the SOI and SST between significant months were used to build composite variables in multiple regressions which gave fitted values of the trap catches to less than 25% of the observed values. The regressions suggested that useful forecasts of both species could be made 6-15 months ahead. The influence of the two weather variables on trap catches of H. punctigera at Turretfield were not as strong as at Narrabri, probably because the SOI was not as strongly related to rainfall in southern Australia as it is in eastern Australia. The best fits were again given by multiple regressions with SOI plus SST variables, to within 40% of the observed values. The reliability of both variables as predictors of moth numbers may be limited by the lack of stability in the SOI-rainfall correlation over the historical record. As no other data set is available to test the regressions, they can only be tested by future use. The use of long-term forecasts in pest management is discussed, and preliminary analyses of other long sets of insect numbers suggest that the Southern Oscillation Index may be a useful predictor of insect numbers in other parts of the world.

Toxicity of biorational insecticides to Helicoverpa spp. (Lepidoptera : Noctuidae) and predators in cotton field

Field trials on upland cotton (Gossypium hirstum L.) during its reproductive phase were used to assess the toxicity of several biorational pesticides and chemicals to Helicoverpa armigera (Hubner) and H. puntigera Wallengren, as well as major predators at Dalby, Queensland, Australia. Moderate rate-dependent control was obtained in plots treated with neem (Azadirachta indica A. Juss) seed extract-azadirachtin (Aza) at rates of 30, 60 and 90 g/ha. Plots treated with Talstar EC (bifenthrin) applications achieved the best results, followed by treatment with alternation of chemicals (methomyl, bifenthrin, thiodicarb and endosulfan) and biorational insecticides (neem oil, azadirachtin and Bacillus thuringiensis kurstaki var. Berliner). Predators, including lady beetles, lacewings, spiders and predatory bugs, were insensitive to Aza, tooseendanin (Tsdn) and BT applications. In contrast, chemicals were very destructive of predators. All treatments provided some protection from infestation of H. armigera and H. puntigera. The effect of Aza on Helicoverpa spp. was reflected in a relatively higher yield of seed cotton harvested from Aza-treated plots compared with the control, but chemical control achieved significantly higher yields than any other treatment.