Effects of larval host plants on over-wintering preparedness and survival of the cotton bollworm, Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

Laboratory colonies of cotton bollworm larvae, Helicoverpa armigera, kept at 20 degrees C under a photoperiod of L:D=10:14 were fed on five host plants (cotton, corn, kidney bean, tobacco and tomato) and an artificial diet (control) to determine the effects of larval host quality on survival and pupal over-wintering preparedness. A separate experiment showed that diapausing pupae weighed more and contained greater nutrient stores than did non-diapausing pupae. Diapausing pupae reared on different host plants showed significant differences in terms of over-wintering reserve storage, and degree of cold-hardiness (extent of low-molecular-weight substances and SCPs), and survivorship. The more nutrients the host plant had, the more the pupae weighed and the higher the levels of total lipids and glycogen. Body water content was also significantly affected by larval food quality. The mean pupal super-cooling capacities varied significantly from -16.7 to -18.9 degrees C according to host plants the larvae feed on, and these significantly related to water content, pupal weight, lipid and glycogen content, and the levels of glycerol. Levels of trehalose, glycerol, and inositol, which were mainly low-molecular-weight substances, showed no significant differences among different host plants, except for trehalose. Pupal mortality varied from 39.7% on corn to 3.3% on the artificial diet, which was significantly related to pupal weight, total lipid content, trehalose levels, and super-cooling points. These results suggest that larval food quality can affect survival and influence the over-wintering preparedness of the cotton bollworm.     

Effects of larval host plants on over-wintering physiological dynamics and survival of the cotton bollworm, Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

Laboratory colonies of cotton bollworm larvae, Helicoverpa armigera, were kept at 20 °C under a photoperiod of L:D = 10:14 and fed on three host plants (cotton, tobacco, kidney bean) and an artificial diet (control) to determine the dynamic effects of larval host quality on over-wintering physiology and mortality. Energy reserves (glycogen and lipid), super-cooling points (SCPs), low-molecular-weight sugars, temperature, and mortality were monitored from November 2002 to April 2003. Lipid content did not change much for each group during over-wintering, but differed according to larval host plants. Larval host plants obviously influence the amount of glycogen, as does time of year: glycogen was lowest in February and increased in early spring. During winter, the mean pupal SCPs increased the most in February, then decreased, and were also affected by larval host plant, i.e. over-wintering pupae reared on kidney bean had the highest SCPs. Levels of glycerol and inositol differed significantly among host plants and months, which peaked in February. Pupal mortality also varied according to larval host plants and time: pupae reached their highest mortality in March and showed host plant differences in January. Records show that February was the coldest month during the period we observed, which corresponded closely to changes in over-wintering characteristics.     

Modeling Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) damages on cotton

We studied and modeled damage caused by Helicoverpa armigera larvae on cotton with the aim of developing a coupled crop pest model. Two damage components were studied: the voracity (quantity of fresh matter and number of organs consumed) and feeding preferences (type of organ infested). The laboratory no-choice study of voracity on excised squares and bolls revealed that an H. armigera larva consumes 2,856 mg of fresh matter throughout its larval life, with the sixth instar consuming 86% of this quantity. This consumption rate corresponded to 23.6 squares, or 7.8 bolls. We developed equations to predict the quantity of fresh matter uptake from an individual plant organ, according to the organ mass and the larval instar. The field study of feeding preference confirmed previous findings that larvae prefer squares to bolls, with this preference decreasing as the larval age increases. However, no significant relationship was noted between the age of larvae and the size of infested organs within each organ class (square or boll). We developed a logistic model to predict the probability of a larva infesting a boll rather than a square. According to this model, the relative organ availability in the field and the larval instar were found to be significant factors.     

Ovicidal activity of Couroupita guianensis (Aubl.) against cotton bollworm Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

There is a great demand for food with increasing population of the world. Many chemical fertilisers and pesticides are used to increase crop productivity. Over use of pesticides and fertilisers has affected the environment and non-target organisms severely; minor pests have become major destructive pests. Hence, eco-friendly approaches are needed. The present study reports the effect of hexane, chloroform and ethyl acetate extract of Couroupita guianensis on the eggs of Helicoverpa armigera. All the extracts showed ovicidal activity, and among them hexane extract showed 64.28% ovicidal activity with LC₅₀ value of 2.62% and regression of (R ²) 83.5%. All the original data showed normality and homogeneity. The active hexane extract was fractionated; finally, eight fractions were obtained and studied at different concentrations. Among the fractions, fraction 8 showed 75% ovicidal activity with R ² of 90%. It showed LC₅₀ value of 322.94?ppm with χ ² value at 0.5%. C. guianensis could be used in pest management programme.     

Organophosphorus insecticides synergize pyrethroids in the resistant strain of cotton bollworm, Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) from West Africa

Helicoverpa armigera (Hübner) populations from West Africa recently developed resistance to pyrethroid insecticides through enhanced metabolism by mixed-function oxidases. The combination index method was used to study the synergism of pyrethroids by organophosphorus insecticides. Several mixtures of insecticides currently registered to control cotton pest complex in West Africa were tested, including: cypermethrin/ethion, cypermethrin/profenofos, deltamethrin/ triazophos, deltamethrin/chlorpyriphos, cyfluthrin/chlorpyriphos, and betacyfluthrin/chlorpyriphos. In the resistant strain, the organophosphorus insecticides significantly increased the toxicity of pyrethroids suppressing the resistance effect, either by additive or synergistic effects. Significant synergism was shown for the following mixtures: cypermethrin/ethion, deltamethrin/triazophos, and deltamethrin/chlorpyriphos. The use of synergism from these insecticide mixtures should prove to be an additional tool in the overall resistance management strategy because the pyrethroid resistance in H. armigera from West Africa is not yet stable, decreasing between cotton seasons and increasing with treatments. In absence of selection, the susceptibility of H. armigera to insecticides should be restored.     

Mechanisms of monocrotophos resistance in cotton bollworm,Helicoverpa armigera (Hübner)

Insensitive acetylcholinesterase was identified as a resistance mechanism by comparing biochemical analysis with a laboratory selected monocrotophos resistant cotton bollworm (RR: 200) and the susceptible strain. The cDNA encoding AChE was cloned by the method of RACE (rapid amplification of cDNA ends). The complete AChE gene deduced from the cDNA consisted of a putative signal peptide of 32 amino acid residues, a mature protein of 615 residues, 5' untranslated regions (UTR) of 315 bp and 3' UTR of 324 bp. The coding sequence had a high degree of homology to the AChE from other insect species reported in the GenBank. After comparing analysis of the entire AChE gene sequence from 5 resistant and 6 susceptible cotton bollworm individuals, nine mutations were identified. One of them, the Ala/Thr mutation, is likely to be responsible for the AChE insensitivity to monocrotophos.     

Structure and ultrastructure of the silk glands and spinneret of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

This study provides comprehensive documentation of silk production in the pest moth Helicoverpa armigera from gland secretion to extrusion of silk thread. The structure of the silk glands, accessory structures and extrusion apparatus are reported. The general schema of the paired silk glands follows that found for Lepidoptera. Morphology of the duct, silk press, muscle attachments and spigot are presented as a three-dimensional reconstruction and the cuticular crescent-shaped profile of the silk press is demonstrated in both open and closed forms with attendant muscle blocks, allowing advances in our knowledge of how the silk press functions to regulate the extrusion of silk. Growth of the spigot across instars is documented showing a distinctive developmental pattern for this extrusion device. Its shape and structure are related to use and load-bearing activity.     

Taxonomy and biogeography of the Nearctic Raphia Hübner (Lepidoptera,Noctuidae, Raphiinae)

The taxonomic status and biogeography of the North American Raphia species is reviewed using adult morphology, larval host plants, geographic phenotypic variation, and variation of mtDNA COI barcode sequences. Lack of diagnostic morphological differences, combined with relatively low mtDNA barcode divergences and clinal phenotypic variation in key geographic regions indicate that the six previously recognized species of North American Raphia are best interpreted as parapatric subspecies. Raphia frater abrupta Grote, stat. n., R. f. coloradensis Putnam-Cramer, stat. r., R. f. piazzi Hill, stat. n., and R. f. elbea Smith, stat. n., are accordingly revised to subspecies of R. frater Grote. Type locality restrictions are provided for Raphia abrupta and Raphia frater and a neotype is designated for Raphia frater var. coloradensis.     

Antifeedant activity of numb and salty taste compounds against the larvae of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

Helicoverpa armigera, an important polyphagous insect pest in agriculture, attacks more than 200 plant species of more than 30 families. Our previous study showed that the choice feeding percentages of H. armigera larvae to tobacco leaf discs treated with sweet, bitter, and hot taste substances were higher than the control leaf discs, while numb and salty substances could significantly inhibit their feeding. To quantitatively determine the synergistic effect of numb and salty substances, in this paper, the antifeeding activities of numb and salty substances and their mixtures blended in different doses or volume ratios were assayed on H. armigera larvae. The first bioassay was designed to elucidate the relative feeding preference of the larvae to the leaves from several common host species, each paired with tobacco leaf discs, and the result indicated that the most preferred host leaf by the larvae was tobacco leaf, followed by cotton and peanut leaves, suggesting that tobacco leaf was the most suitable matrix for the antifeeding bioassay, and the larval consumption of maize, pepper, or tomato leaves were significantly lower than that of tobacco leaf. The second bioassay was to test the choice feeding response of H. armigera larvae to tobacco leaf discs treated with Zanthoxylum bungeanum extracts obtained with different solvents, and the result showed that the antifeeding activity of the alcohol extracts was the strongest (93.38%), and the leaf consumption in the treatment and the control showed extremely significant difference (t = 4.23, t_0.01 = 3.25, P = 0.0022), followed by the dichloromethane extracts (47.64%), while the other three solvents (water, acetone, and n-hexane) could not extract the active antifeeding components from Z. bungeanum. The larval consumption of tobacco leaf discs treated with the alcohol extracts of Z. bungeanum and NaCl solution were significantly less than their corresponding controls. The mean larval consumption of the treated leaf discs decreased with ever-increasing dosage, and the consumption of tobacco leaf discs coated with different doses of alcohol extracts of Z. bungeanum or NaCl solution showed extremely significant difference (F_{alcohol extract of Z. bungeanum} = 3.88, F_0.01 = 3.58, P = 0.0064; F_{NaCl solution} = 54.29, F_0.01 = 3.58, P = 0.0000), with maximum antifeeding effects at a dosage of 30 μL per 1.5 cm ID leaf disc. We further tested the larval consumption of tobacco leaf discs treated with alcohol extracts of Z. bungeanum in saturated NaCl solution mixed in different volume ratios, and the result showed that the choice antifeeding percentages of the treatments with 15 μL or more Z. bungeanum alcohol extracts were higher than 90%, among which the mixture with 25:15 volume ratio of Z. bungeanum alcohol extracts and saturated NaCl solution exhibited the strongest antifeeding activity, and the mean consumed leaf area of tobacco leaf discs coated with this blend was only 0.10 mm². In the further test on feeding dose-response of the 25:15 mixture, the mean leaf consumption decreased linearly with ever-increasing dosage, with a regression equation y = ?3.9356x + 120.78(R² = 0.9998), and the 30 μL dose could completely inhibit H. armigera feeding.     

Karyomorphological effects of two new oil formulations on Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

The cotton bollworm Helicoverpa armigera is a serious pest of many economically important crops. Since this pest has become resistant to the conventional synthetic insecticides, newer compounds and formulations are being developed against this insect pest. Many natural compounds isolated from the plants were tested against this pest. Among them Hyptis suaveolens and Melochia corchorifolia showed insecticidal properties against H. arnigera. Based on bioefficacy studies, caryophyllene and β-sitosterol were isolated from H. suaveolens and M. corchorifolia respectively. The isolated natural compounds were further developed as formulations in various combinations with neem (Azadirachta indica) and karanj (Pongamia pinnata) oils. The present study indicated that the formulations influenced the karyomorphology of H. armigera.     

Ultrastructure of the antennal sensilla of Alabama argillacea (Hübner, 1823) (Lepidoptera: Erebidae)

Insects have several types of sensilla, the characterization of which has been fundamental to understanding the mechanisms of sensory perception in different species. This study aimed to describe the ultrastructure of the sensilla present on the antennae of Alabama argillacea (Hübner, 1823) (Lepidoptera: Erebidae), an important pest of cotton (Gossypium hirsutum L.) crops, as well as their possible variation between sexes. To do this, the antennae of males and females of A. argillacea were analyzed using scanning electron microscopy. Sensilla morphometry was assessed using photomicrographs, from which the lengths and basal and apical diameters of sensilla were measured using the ImageJ program. Seven types of sensilla were identified on the antennae of A. argillacea: sensilla trichodea, sensilla chaetica, sensilla auricillica, sensilla coeloconica, sensilla styloconica, sensilla basiconica, and sensilla Böhm bristles. Differences between the sensilla of males and females were found in their lengths and basal diameters in the distal and proximal regions. This suggests that sensilla functionality may not only vary from one species to another, but also between sexes within the same species. Thus, further transmission electron microscopy and single sensillum recording studies may provide more detailed information on the sensilla of A. argillacea and their functions.     

Effect of the populations of Alabama argillacea (Hübner) (Lepidoptera: Noctuidae) on cotton crop at two sowing distances

The objective of this research was to evaluate the incidence of damage by the leafworm Alabama argillacea (Hübner) on yields of cotton grown under rows spaced 0.38 m and 0.76 m at the irrigation area of Santiago del Estero, Argentina. The period evaluated was extended from first flower to first open boll. Treatments were T1 - without control of larvae, T2 - with control of larvae, T3 - control since first flower to the end of effective blooming, and T4 - with larval control since the end of effective blooming to first open boll. The effect of injuries on the crop was evaluated trough boll cotton yield. Larvae were sampled in a weekly basis and insects were present from the first flower until harvest. Populations of A. argillacea decreased crop yields in the two distances tested, by decreasing the weight or number of open bolls.     

Biopotential of Fleuggea leucopyrus (Koen.) Willd. against Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae)

In the past five decades, agricultural pests are controlled by synthetic pesticides which caused ill effects on non-target organisms and environment; also insect pests developed resistance and minor pests became major pests. By keeping this in mind, the present study was selected. Antifeedant and larvicidal activities and growth inhibitory effects of hexane, chloroform and ethyl acetate leaf extracts of Fleuggea leucopyrus (Koen.) Willd. against Helicoverpa armigera (Hbn.) were studied. Antifeedant and larvicidal activities were evaluated at 0.5, 1, 2.5 and 5% concentrations at laboratory conditions. Hexane and chloroform extracts at 5% concentration recorded significant antifeedant activity of 82.41 and 74.45%, respectively. Hexane extract recorded the least LC₅₀ and LC₉₀ values of 1.37 and 4.80%, respectively. Hexane extract recorded maximum larvicidal activity of 85.78% with LC₅₀ and LC₉₀ values of 1.69 and 4.94%, respectively. Ethyl acetate extract also had notable amount of larvicidal activity of 81.77% at 5% concentration. Hexane extract at 5 and 2.5% and ethyl acetate extract at 5% concentrations completely prevented the adult emergence of H. armigera. This plant could be further exploited to identify the active principle(s) responsible for the activities and to develop a novel pesticidal formulation. This is the first report of F. leucopyrus is studied for its bioefficacy against H. armigera.     

External antennal morphometry of Trichoplusia Ni (Hübner) (Lepidoptera: Noctuidae)

A detailed morphometric analysis of the antennae of male and female Trichoplusia ni (Hübner) is presented, based on light and scanning electron microscopic observations. These measurements of general antennal dimensions and the number, distribution, dimensions, and surface features of the antennal sensilla are applied in the consideration of olfactory transduction processes and in the derivation of a mathematical representation of the antennal surface. The surface area and the number of sensilla on the antennal flagellum are found to vary with the weight of the pupa and adult moth. The long Type 1 sensilla trichodea are hypothesized to be innervated by pheromone-sensitive olfactory cells on the basis of morphological comparisons with sensilla of other insects.     

Mechanisms of premating isolation between Helicoverpa armigera (Hübner) and Helicoverpa assulta (Guenée) (Lepidoptera: Noctuidae)

Helicoverpa armigera and Helicoverpa assulta are sympatric sibling species, and in the laboratory they can interbreed and produce viable offspring. To assess the contributions of temporal barriers and sexual barriers to premating isolation, we investigated both the temporal rhythms of calling behavior and pheromone titers of H. armigera and H. assulta females and the behavioral responses of males to conspecific and heterospecific calling females in a wind tunnel. Both H. armigera and H. assulta females called throughout the scotophase, and there was more calling during the second half of the scotophase than during the first half. Maximal pheromone titer and maximal calling activity in H. armigera synchronously occurred at the sixth hour into the scotophase, whereas, in H. assulta, the maximal pheromone titer occurred 2 h before the peak of calling. Pheromone blend ratios of the two species were opposite and, within each species, changes in the ratio within the scotophase and at different ages were relatively small. Males of both H. armigera and H. assulta responded strongly to their conspecific calling females in the wind tunnel and completed the whole courtship sequence. In contrast, they did not land and had no copulation attempts in response to heterospecific calling females. These results show that the two species do not have obvious temporal differences in calling behavior and pheromone production, and the specificity of sex pheromone blend emitted by females plays a key role in their premating isolation. In addition, we summarized the potential isolation mechanisms of H. armigera and H. assulta.     

Novel polymorphic microsatellite markers developed in the cotton bollworm Helicoverpa armigera （Lepidoptera：Noctuidae）

A novel set of five polymorphic di- or trinucleofide microsatellite loci suitable for population genetic study were developed from an enriched genomic library for the pest insect cotton bollworm, Helicoverpa armigera, and cross-amplifiability of these and other published loci was tested in a closely related species, the tobacco budworm, H.assulta. The expected heterozygosity at these loci ranges from 0.62 to 0.91 in the cotton bollworm. The observed allele numbers varies from 4 to 12 in the limited number of individuals tested. Although a large proportion of cloned microsatellite sequences are present in multi-copy in the cotton bollworm genome, the overwhelming majority of the finalized polymorphic diallelic loci are tri-nucleotide microsatellites - an unexpected outcome, which should facilitate subsequent genotyping analysis.     

Electrophysiological responses of female Helicoverpa armigera (Hübner) (Lepidoptera; Noctuidae) to synthetic host odours

Studies were conducted to investigate the electroantennographic (EAG) responses of adult female Helicoverpa armigera to a range of known and putative kairomone components. The studies show that at a given dose the EAG responses elicited by a series of straight-chain aliphatic primary alcohols were not dependent on volatility since butan-1-ol and pentan-1-ol elicited EAG responses that were significantly smaller than those elicited by hexan-1-ol. The amplitudes of responses to hexan-1-ol were found to be dose dependent with a dose of 10(-1) μmol at source in a non-volatile solvent eliciting the largest response. Similarly, changes in functionality in a range of C(6) straight-chain aliphatic compounds significantly changed the amplitude of response elicited, with aldehydes eliciting smaller responses than the related primary alcohols and saturated compounds eliciting higher responses than related unsaturated compounds. Of the range of nine host plant-produced terpenoids tested, ocimene and beta-phellandrene elicited the highest responses and of the six aromatic compounds tested phenylacetaldehyde and benzaldehyde elicited the largest responses, at the doses tested. The significance of these findings for analysis of floral odours by gas chromatography linked to electroantennography as a means of identifying kairomone components attractive to H. armigera are discussed.