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.

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

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.

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.

An evaluation of selective Helicoverpa armigera control options in sweet corn

Helicoverpa armigera is a serious insect pest of sweet corn in Australia and is becoming increasingly difficult to manage with conventional chemical insecticides due to resistance problems. A number of alternative H. armigera control options were evaluated in sweet corn and compared with deltamethrin and no action (control). The alternative tactics evaluated were: heliothis nuclear polyhedrosis virus plus Trichogramma nr. brassicae releases; Bacillus thuringiensis; and Trichogramma alone. The H. tea nuclear polyhedrosis virus + Trichogramma plots had the lowest cob damage (6.0%), followed by the B. thuringiensis plots (12.0%), Trichogramma alone plots (20.2%), control plots (23.2%) and deltamethrin plots (53.5%). There was no evidence to suggest that the Trichogramma nr. brassicae releases had any impact on H. armigera egg mortality. However, there was a large natural population of Trichogramma pretiosum in all plots. The application of deltamethrin reduced the action of these wasps and predators, resulting in higher larval infestation and significantly more cob damage. The findings indicate that the pathogens heliothis nuclear polyhedrosis virus and B. thuringiensis can effectively control H. armigera when their action complements high natural levels of egg parasitism, and that they have potential for use in integrated pest management programs in sweet corn.

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.

Genetic shifts in Helicoverpa armigera Hubner (Lepidoptera : Noctuidae) over a year in the Dawson/Callide Valleys

Microsatellites were used to analyse 68 collections of Helicoverpa armigera in the Dawson/Callide Valleys in central Queensland. The study aimed to evaluate the genetic structure in this region over a 12-month period (September 2000-August 2001). The results detected genetic shifts in H. armigera collections, with genetic changes occurring month by month. Collections in any month were genetically distant from the preceding month's collections. There was no observed difference between collections of H. armigera from the Biloela region and those found in the Theodore region of central Queensland. The data support the current area-wide management strategies for H. armigera by reinforcing the importance and contribution of local management practices. The study also indicates a need for the continuation of regional or Australia-wide approaches to management of the low levels of immigration that are occurring, and for future high pest pressure years.

Sensitivity of Helicoverpa armigera nucleopolyhedrovirus polyhedra to sodium dodecyl sulfate

Sodium dodecyl sulfate (SDS) is commonly used to extract polyhedra from infected cells and diseased dead larval tissues. It was found, however, that 80% of Helicoverpa armigera nucleopolyhedrovirus (HaSNPV) polyhedra produced via cell culture were damaged after 30 min of 0.5% SDS treatment whereas only 20% of in vivo produced polyhedra were damaged by the same treatment. Transmission and scanning electron microscopy revealed that the damaged polyhedra had lost their polyhedron envelopes and virions were dislodged from the polyhedrin matrix, leaving empty spaces that were previously occupied by the occluded virions. Up to 20% in vitro produced polyhedra were resistant to SDS and remained intact, even after a 24 h exposure to SDS. This sensitivity to SDS was observed across a range of cell culture media, including serum supplemented media. Electron microscopy also revealed that the inferior polyhedron envelope of in vitro produced polyhedra is likely due to poor interaction between the polyhedron envelope, polyhedron envelope protein (PEP), and polyhedrin matrix. The PEP gene was cloned and sequenced and mutations in this gene were ruled out as an explanation. In vitro produced polyhedra that were passed through insect larva once were resistant to SDS, indicating that a critical component is lacking in insect cell culture medium used for producing HaSNPV or the cells growing in culture are inefficient in some ways in relation to production of polyhedra. (C) 2002 Elsevier Science (USA). All rights reserved.

Molecular cloning and insecticidal effect of Inga laurina trypsin inhibitor on Diatraea saccharalis and Heliothis virescens

Native Inga laurina (Fabaceae) trypsin inhibitor (ILTI) was tested for anti-insect activity against Diatraea saccharalis and Heliothis virescens larvae. The addition of 0.1% ILTI to the diet of D. saccharalis did not alter larval survival but decreased larval weight by 51%. The H. virescens larvae that were fed a diet containing 0.5% ILTI showed an 84% decrease in weight. ILTI was not digested by the midgut proteinases of either species of larvae. The trypsin levels were reduced by 55.3% in the feces of D. saccharalis and increased by 24.1% in the feces of H. virescens. The trypsin activity in both species fed with ILTI was sensitive to the inhibitor, suggesting that no novel proteinase resistant to ILTI was induced. Additionally, ILTI exhibited inhibitory activity against the proteinases present in the larval midgut of different species of Lepidoptera. The organization of the ilti gene was elucidated by analyzing its corresponding genomic sequence. The recombinant ILTI protein (reILTI) was expressed and purified, and its efficacy was evaluated. Both native ILTI and reILTI exhibited a similar strong inhibitory effect on bovine trypsin activity. These results suggest that ILTI presents insecticidal properties against both insects and may thus be a useful tool in the genetic engineering of plants. (c) 2012 Elsevier Inc. All rights reserved.

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.

Foraging behaviour of Helicoverpa armigera first instar larvae on crop plants of different developmental stages

Understanding how insect pests forage on their food plants can help optimize management strategies. Helicoverpa armigera (Hubner) (Lep., Noctuidae) is a major polyphagous pest of agricultural crops worldwide. The immature stages feed and forage on crops at all stages of plant development, damaging fruiting and non-fruiting structures, yet very little is known about the influence of host type or stage on the location and behaviour of larvae. Through semi-continuous observation, we evaluated the foraging (movement and feeding) behaviours of H. armigera first instar larvae as well as the proportion of time spent at key locations on mungbean [Vigna radiata (L.) Wilczek] and pigeon pea [Cajanus cajan (L.) Millspaugh] of differing developmental stages: seedling- and mature (flowering/pod fill)-stage plants. Both host type and age affected the behaviour of larvae. Larvae spent more time in the upper parts of mature plants than on seedlings and tended to stay at the top of mature plants if they moved there. This difference was greater in pigeon pea than in mungbean. The proportion of time allocated to feeding on different parts of a plant differed with host and age. More feeding occurred in the top of mature pigeon pea plants but did not differ between mature and seedling mungbean plants. The duration of key behaviours did not differ between plant ages in either crop type and was similar between hosts although resting bouts were substantially longer on mungbeans. Thus a polyphagous species such as H. armigera does not forage in equivalent ways on different hosts in the first instar stage.

Properties of a unique mutant of Helicoverpa armigera single-nucleocapsid nucleopolyhedrovirus that exhibits a partial many polyhedra and few polyhedra phenotype on extended serial passaging in suspension cell cultures

Serial passaging of wild-type Helicoverpa armigera, single-nucleocapsid (HaSNPV) in H. zea (HzAMI) illsect Cell Cultures results ill rapid selection for the few polyhedra (FP) phenotype. A unique HaSNPV mutant (ppC19) was isolated through plaque purification that exhibited a partial many polyhedra (MP) and FP phenotype. Oil serial passaging in suspension cell cultures, ppC19 produced fivefold more polyhedra than a typical FP mutant (FP8AS) but threefold less polyhedra than the wild-type virus. Most importantly, the polyhedra of ppC19 exhibited MP-like virion occlusion. Furthermore, ppC19 produced the same amount of budded virus (BV) as the FP mutant, which was fivefold higher than that of the wild-type virus. This selective advantage was likely to explain its relative stability in polyhedra production for six passages when compared with the wild-type Virus. However, subsequent passaging of ppC19 resulted in a steel) decline in both BV and polyhedra yields, which was also experienced by FP8AS and the wild-type virus Lit high passage numbers. Genomic deoxyribonueleic Licid profiling of the latter suggested that defective interfering particles (DIPS) were implicated in this phenomenon and represented another Undesirable mutation during serial passaging of HaSNPV Hence, a strategy to isolate HaSNPV Clones that exhibited MP-like polyhedra production but FP-like BV production, coupled with low multiplicities of infection during scale-up to avoid accumulation of DIPS, could prove commerically invaluable.

Host recognition by a polyphagous lepidopteran (Helicoverpa armigera): primary host plants, host produced volatiles and neurosensory stimulation

An important question in the host-finding behaviour of a polyphagous insect is whether the insect recognizes a suite or template of chemicals that are common to many plants? To answer this question, headspace volatiles of a subset of commonly used host plants (pigeon pea, tobacco, cotton and bean) and nonhost plants (lantana and oleander) of Helicoverpa armigera Hubner (Lepidoptera: Noctuidae) are screened by gas chromatography (GC) linked to a mated female H. armigera electroantennograph (EAG). In the present study, pigeon pea is postulated to be a primary host plant of the insect, for comparison of the EAG responses across the test plants. EAG responses for pigeon pea volatiles are also compared between females of different physiological status (virgin and mated females) and the sexes. Eight electrophysiologically active compounds in pigeon pea headspace are identified in relatively high concentrations using GC linked to mass spectrometry (GC-MS). These comprised three green leaf volatiles [(2E)-hexenal, (3Z)-hexenylacetate and (3Z)-hexenyl-2-methylbutyrate] and five monoterpenes (alpha-pinene,beta-myrcene, limonene, E-beta-ocimene and linalool). Other tested host plants have a smaller subset of these electrophysiologically active compounds and even the nonhost plants contain some of these compounds, all at relatively lower concentrations than pigeon pea. The physiological status or sex of the moths has no effect on the responses for these identified compounds. The present study demonstrates how some host plants can be primary targets for moths that are searching for hosts whereas the other host plants are incidental or secondary targets.