Relative Fitness of Helicoverpa armigera (Lepidoptera: Noctuidae) on Seven Host Plants: A Perspective for IPM in Brazil

The cotton bollworm Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) is a widespread pest of many cultivated and wild plants in Europe, Africa, Asia, and Australia. In 2013, this species was reported in Brazil, attacking various host crops in the midwestern and northeastern regions of the country and is now found countrywide. Aiming to understand the effects of different host plants on the life cycle of H. armigera, we selected seven species of host plants that mature in different seasons and are commonly grown in these regions: cotton (Gossypium hirsutum, “FM993”), corn (Zea mays, “2B587”), soybean (Glycine max, “99R01”), rattlepods (Crotalaria spectabilis), millet (Pennisetum glaucum, “ADR300”), sorghum (Sorghum bicolor, “AGROMEN70G35”), and cowpea (Vigna unguiculata, “SEMPRE VERDE”). The development time of immatures, body weight, survivorship, and fecundity of H. armigera were evaluated on each host plant under laboratory conditions. The bollworms did not survive on corn, millet, or sorghum and showed very low survival rates on rattlepods. Survival rates were highest on soybean, followed by cotton and cowpea. The values for relative fitness found on soybean, cotton, cowpea, and rattlepods were 1, 0.5, 0.43, and 0.03, respectively. Survivorship, faster development time, and fecundity on soybean, cotton, and cowpea were positively correlated. Larger pupae and greater fecundity were found on soybean and cotton. The results indicated that soybean, cotton, and cowpea are the most suitable plants to support the reproduction of H. armigera in the field.     

Adaptation of Habrobracon hebetor (Hymenoptera: Braconidae) to Rearing on Ephestia kuehniella (Lepidoptera: Pyralidae) and Helicoverpa armigera (Lepidoptera: Noctuidae)

Food characteristics strongly regulate digestive enzymatic activity of insects through direct influences on their midgut mechanisms. Insect performance is better on diets that contain nutrients in proportions that fit its digestive enzymes. Little is known about the influences of rearing history on parasitism success of Habrobracon hebetor Say. This research focused on the effect of nutrient regulation on survival, development, and parasitism of H. hebetor. Life history and digestive enzyme activity of fourth-stage larvae of H. hebetor were studied when reared on Ephestia kuehniella Zeller. This parasitoid was then introduced to Helicoverpa armigera (Hübner), and above-mentioned parameters were also studied in the first and fourth generations after transfer. In term of parasitism success, H. hebetor preferred E. kuehniella over He. armigera. When the first and fourth generations of He. armigera-reared H. hebetor were compared, the rearing history affected the life history and enzymatic activity of the parasitoid. A better performance of H. hebetor was achieved after it was reared on He. armigera for the four generations. Because, digestive α-amylase and general protease of the parasitoid were matched with the new host, it used reserve energy for a better performance. Thus, a better performance of H. hebetor could be obtained when the parasitoid was reared on its original host for at least four generations.     

Comparative Feeding Performance and Digestive Physiology of Helicoverpa armigera (Lepidoptera: Noctuidae) Larvae-Fed 11 Corn Hybrids

This study aimed to evaluate the feeding responses and digestive proteolytic and amylolytic activity of Helicoverpa armigera (Hübner) on 11 corn (Zea mays L.) hybrids at 25 ± 1°C, 65 ± 5% relative humidity (RH), and a photoperiod of 16:8 (L:D) h. The fourth- and fifth-instar larvae fed on hybrid K47*K19 had the highest weight of food consumption and those reared on hybrid KSC705 had the lowest value of food consumption. The highest weight gain of the larvae was observed when H. armigera were fed hybrid KLM78*MO17 and lowest when they were fed hybrids K36 * MO17, KSC705, and K35 * K36. Pupal weight of H. armigera was heaviest when larvae were fed hybrid K47*K19 and lightest when they were fed hybrid KSC705. The highest proteolytic activity of the fourth-instar larvae was observed when they were fed hybrid KSC705, and the lowest activity was observed when they were fed hybrid K47*A67. Fifth-instar larvae that fed on hybrid K47*K19 showed the highest proteolytic activity. Fourth-instar larvae that fed on hybrid K36*MO17 showed the highest amylase activity. The fifth-instar larvae fed on hybrid K47*A67 showed the maximum amylase activity and those reared on the K48*K18 showed the minimum activity. Our results indicated that K36 * MO17, KSC705, and K48 * K18 were the most unsuitable hybrids for feeding H. armigera.     

The Complete Mitochondrial Genome of Brachmia macroscopa (Lepidoptera: Gelechiidae) and Its Related Phylogenetic Analysis

The sweet potato leaf folder, Brachmia macroscopa, is an important pest in China. The complete mitogenome, which consists of 13 protein-coding genes (PCGs), 22 transfer RNA genes, two ribosomal RNA genes, and an A + T-rich region, was sequenced and found to be 15,394 bp in length (GeneBank no. KT354968). The gene order and orientation of the B. macroscopa mitogenome were similar to those of other sequenced lepidopteran species. All of the PCGs started with ATN as the canonical start codon except for cox1, which started with CGA. In regard to stop codons, most PCGs stopped at TAA except for cox2, which stopped at TA, and nad4, which stopped at a single T. Thirteen PCGs of the available species (33 taxa) were used to demonstrate phylogenetic relationships. The ditrysian cluster was supported as a monophyletic clade at high levels by using maximum likelihood and Bayesian methods. The apoditrysian group, covering the Gelechioidea, formed a monophyletic clade with a bootstrap value of 88% and a posterior probability of 1.00. The superfamily Gelechioidea was supported as a monophyletic lineage by a posterior probability of 1.00.     

Insecticidal Activity and Insect Repellency of Four Species of Sea Lily (Comatulida: Comatulidae) From Taiwan

The methanol and ethyl acetate (EA) extracts of four species of sea lily (Himerometra magnipinna, Comaster multifidus, Comanthina sp., and Comatella maculata) were evaluated for their insecticidal activity against Yellow-fever mosquito larvae (Aedes aegypti) and their repellency against adult Asian Tiger mosquitoes (Aedes albopictus). The 24-hr minimum inhibition concentration (MIC) data revealed that the extracts from H. magnipinna and the C. maculata were the most active, killing mosquito larvae at 12.5 ppm. The toxicity of the extracts from these four sea lilies in descending order was H. magnipinna (12.5 ppm), C. maculata (12.5 ppm), C. multifidus (100 ppm), and Comanthina sp. (200 ppm). Furthermore, no significant difference in toxicity was found using either EA or methanol as the extraction solvent. The MIC at 12.5 ppm is promising as an insecticide lead. The repellency study results show that EA is a better solvent for one species (H. magnipinna), but the methanol is a better solvent overall. The repellency of these sea lily extracts in descending order was Comanthina sp. MeOH (ED₅₀ at 0.32%), followed by H. magnipinna EA (ED₅₀ at 0.38%), C. multifidus MeOH (ED₅₀ at 0.57%), C. maculata MeOH (ED₅₀ at 0.76%), C. multifidus EA (ED₅₀ at 1.25%), and H. magnipinna MeOH (ED₅₀ at 1.67%). A compound with ED₅₀ <0.5% is considered to be a promising repellant. Among the studied sea lilies, both Comanthina sp. and H. magnipinna have potential to be further developed as mosquito control agents due to their favorable toxicity and repellency.     

Prevalence of Helicoverpa zea (Lepidoptera: Noctuidae) on late season volunteer corn in Mississippi: Implications on Bt resistance management

The southern United States has a long growing period between corn, Zea mays L., harvest and first winter frost, so volunteer corn which germinates after harvest has a growing period sufficient for corn earworm, Helicoverpa zea (Boddie) and fall armyworm, Spodoptera frugiperda (J. E. Smith) to feed on these plants. However, lower air temperatures can limit larval development on late season volunteer corn and thereby successful pupation. Here we explore the suitability of late season volunteer corn for larval development and the potential contribution of H. zea larvae to the overwintering population. Our survey revealed the occurrence of volunteer corn in high densities, with monthly mean densities ranging from 56,000 to 143,000 plants ha⁻¹. H. zea larvae were found feeding on both vegetative and reproductive stage plants while S. frugiperda were only found on vegetative stage plants. An analysis of H. zea growing degree day (GDD) accumulations based on Mississippi weather data from 1980 to 2010 revealed that sufficient GDD to reach prepupation would always be accumulated before first frost if oviposition occurred by 9 September, with the probability of successful pupation decreasing rapidly thereafter. However, most of the H. zea larvae were oviposited after this, and could not reach pupation. Because S. frugiperda cannot overwinter in Mississippi, their ability to pupate was not examined. Low suitability of whorl stage corn for H. zea development coupled with low larval densities during this stage effectively diminish the number of larvae that complete development on late season volunteer transgenic corn expressing genes from the soil bacterium, Bacillus thuringiensis (Bt). This limits the Bt resistance risk posed by larvae developing on late season volunteer corn in all but the most southern locations in the US.     

Identification and Physicochemical Properties of the Novel Hemolysin(s) From Oral Secretions of Helicoverpa armigera (Lepidoptera: Noctuidae)

Hemolysins cause the lysis of invading organisms, representing major humoral immunity used by invertebrates. Hemolysins have been discovered in hemolymph of Helicoverpa armigera larvae as immune factors. As oral immunity is great important to clear general pathogens, we presumed that hemolysins may be present in oral secretions (OS). To confirm this hypothesis, we conducted four testing methods to identify hemolysin(s) in larval OS of H. armigera, and analyzed physicochemical properties of the hemolysin in comparison with hemolytic melittin of Apis mellifera (L.) (Hymenoptera: Apidae) venom. We found hemolysin(s) from OS of H. armigera for the first time, and further identified in other lepidopteran herbivores. It could be precipitated by ammonium sulfate, which demonstrates that the hemolytic factor is proteinaceous. Labial gland showed significantly higher hemolytic activity than gut tissues, suggesting that hemolysin of OS is mainly derived from saliva secreted by labial glands. Physicochemical properties of hemolysin in caterpillar’s OS were different from bee venom. It was noteworthy that hemolytic activity of OS was only partially inhibited even at 100°C. Hemolytic activity of OS was not inhibited by nine tested carbohydrates contrary to bee venom melittin. Moreover, effects of metal ions on hemolytic activity were different between OS and bee venom. We conclude that there is at least a novel hemolysin in OS of herbivorous insects with proposed antibacterial function, and its hemolytic mechanism may be different from melittin. Our study enriches understanding of the potential role of hemolysins in insect immunity and provides useful data to the field of herbivorous insect-pathogen research.     

Susceptibility of Helicoverpa armigera from different host plants in northern China to Bacillus thuringiensis toxin Cry1Ac

Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) is a serious pest of cotton and many other crops in northern China. To evaluate the contribution of alternative hosts as an effective refuge for transgenic cotton expressing the Bacillus thuringiensis (Bt) Cry1Ac toxin, the susceptibility to this toxin was measured in progeny derived from field-collected H. armigera larvae and pupae from different hosts in the Xiajin’s region of the Shandong Province in northern China. During 2008-2010, progeny from a total of 258,56,184 and 160 single-pair crosses derived from wheat (first-generation), Bt cotton (second-generation), Bt cotton (third-generation), and corn (third-generation) were screened on Cry1Ac diets, respectively. Based on relative average development rates (RADR) of H. armigera larvae in these F₁ tests, the second and third-generation moths emerging from Bt cotton fields were more tolerant to the Bt toxin than the first and third-generation moths emerging from wheat and corn each year. These results suggest that there is significant variation in susceptibility to Bt toxins among H. armigera populations derived from different host crops. Alternate crops, such as corn, that maintain Bt susceptible populations of H. armigera could be used as refugia to minimize the evolution of resistance to Bt cotton.     

Acute toxicity and sublethal effects of methoxyfenozide and thiodicarb on survival,development and reproduction of Helicoverpa armigera (Lepidoptera: Noctuidae)

The cotton bollworm Helicoverpa armigera (Hubner) is one of the most destructive pest insects in Iran and many other countries. In this study, lethal and sublethal effects of methoxyfenozide, and thiodicarb were evaluated against H. armigera larvae that fed on insecticide-treated artificial diet. The effects of methoxyfenozide and thiodicarb were assessed in 3rd instars. Methoxyfenozide and thiodicarb showed LC₅₀ values of 4 and 639 mg a.i./ml, respectively, in H. armigera larvae. Sublethal effects on development, adult longevity, and reproduction were observed in H. armigera larvae that survived exposure to an LC₃₀ of the insecticides. Larvae that were exposed to an LC₃₀ concentration of methoxyfenozide exhibited lower pupal weight and increased larval and pupal developmental times compared with thiodicarb treated larvae or control larvae. Adults that were exposed as larvae to an LC₃₀ concentration of methoxyfenozide or thiodicarb showed reduced fecundity (35% and 30%, respectively) compared to control adults. The tested insecticides significantly reduced adult longevity. The longevity of adult females that as larvae were treated with an LC₃₀ concentration of methoxyfenozide or thiodicarb was reduced by 28% and 23%, respectively, in comparison to control females. We predict that the combination of lethal and sublethal effects of the insecticides, especially methoxyfenozide, will induce significant effects on field population dynamics of H. armigera.     

Manual topping decreases bollworm infestations in cotton cultivation in Mali

In sub-Saharan areas of Africa, cotton growers no longer cut the shoot tips from plants (topping), although manual topping was promoted at the start of the 20th century to improve yield and, surprisingly, to reduce pest incidence. In these areas, the bollworms Helicoverpa armigera Hübner, Earias spp., and Diparopsis watersi Rothschild are responsible for the majority of cotton yield losses, and the use of pyrethroids has resulted in resistance in field populations of H. armigera. In the face of these problems and given the scarcity of literature on the effects of topping on pest control, we assessed bollworm infestation levels in 12 trials comparing manual topping and non-topping cotton plots in Mali over a six-year period (2002, 2003, and 2005 to 2008). Topping was performed at the emergence of the 15th sympodial branch or at 10 days after the first flower opening. Our results showed no significant difference in seed cotton yields between topped and non-topped cotton. Bollworm infestations (all species) were always lower on topped cotton and 7 out of 12 trials showed significantly lower infestations on topped cotton. In plots of topped cotton, we recorded an average of 56% fewer H. armigera larvae, 68% fewer Earias spp. larvae, and 71% fewer D. watersi larvae with respectively 5, 4, and 3 out of 12 trials with significant differences in favor of topping and no significant difference in favor of non-topping. To our knowledge, our study is the first to report decreases in D. watersi larval infestation with cotton topping. Further studies are required to understand the mechanisms involved in these effects and to ensure that topping is economically attractive for farmers.     

Chemical Composition,Antifeedant, Repellent,and Toxicity Activities of the Rhizomes of Galangal,Alpinia galanga Against Asian Subterranean Termites,Coptotermes gestroi and Coptotermes curvignathus (Isoptera: Rhinotermitidae)

Dual choice bioassays were used to evaluate the antifeedant property of essential oil and methanolic extract of Alpinia galanga (L.) (locally known as lengkuas) against two species of termites, Coptotermes gestroi (Wasmann) and Coptotermes curvignathus (Holmgren) (Isoptera: Rhinotermitidae). A 4-cm-diameter paper disc treated with A. galanga essential oil and another treated with either methanol or hexane as control were placed in a petri dish with 10 termites. Mean consumption of paper discs (miligram) treated with 2,000 ppm of essential oil by C. gestroi was 3.30 ± 0.24 mg and by C. curvignathus was 3.32 ± 0.24 mg. A. galanga essential oil showed significant difference in antifeedant effect, 2,000 ppm of A. galanga essential oil was considered to be the optimum concentration that gave maximum antifeedant effect. The essential oil composition was determined using gas chromatography-mass spectrometry. The major component of the essential oil was 1,8-cineol (61.9%). Antifeedant bioassay using 500 ppm of 1,8-cineol showed significant reduction in paper consumption by both termite species. Thus, the bioactive agent in A. galangal essential oil causing antifeeding activity was identified as 1,8-cineol. Repellent activity shows that 250 ppm of 1,8-cineol caused 50.00 ± 4.47% repellency for C. gestroi, whereas for C. curvignathus 750 ppm of 1,8-cineol was needed to cause similar repellent activity (56.67 ± 3.33%). C. curvignathus is more susceptible compare to C. gestroi in Contact Toxicity study, the lethal dose (LD₅₀) of C. curvignathus was 945 mg/kg, whereas LD₅₀ value for C. gestroi was 1,102 mg/kg. Hence 1,8-cineol may be developed as an alternative control against termite in sustainable agriculture practices.     

Monitoring resistance of Helicoverpa armigera to different insecticides used in cotton in Spain

Helicoverpa armigera is the key pest of cotton in Spain, resulting in many insecticide treatments against it. The resistance status of H. armigera to different insecticides currently used in cotton was evaluated in Spain in two different seasons, 1999 and 2004. Four populations were tested in total, two in each season. Toxicological bioassays were conducted in the laboratory, and performed on third instar larvae by topical application of the insecticides. LD₅₀'s were estimated by probit analysis and resistance factors (RF) were calculated at the LD₅₀ level. Four insecticides were evaluated, but only endosulfan reached a moderate resistance level (RF = 11.4), and the others (methomyl, chlorpyrifos and lambda-cyhalothrin) showed low resistance (RF between 1.9 and 6.0). Such results indicate the generally low resistance of H. armigera to most of the insecticides used against this pest in cotton in Spain. Possible explanations for this situation are discussed.     

Frequency of Bt resistance alleles in Helicoverpa armigera in 2007-2009 in the Henan cotton growing region of China

Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) is a key insect pest of cotton in the Henan cotton growing region of China. In this region, cotton is grown on small acreages in rich agricultural landscapes, contrary to cropping systems in the United States or Australia. Under such cropping regimes, naturally occurring refuges (with non-Bt plants) may be sufficient to prevent H. armigera resistance development to Bt toxins. In order to gain a timely understanding of the evolution of resistance of H. armigera to Bt toxin after continuous cultivation of Bt cotton for c. 10 years, we assessed the frequency of alleles conferring resistance to Cry1Ac toxin in field populations of H. armigera sampled from Xinxiang County in Henan province during 2007-2009. Screening F₁ and F₂ generations from isofemale lines, derived from female moths trapped in the field, were used with a discriminating dose of Cry1Ac diet to estimate the frequency of resistance alleles. Totals of 625, 516 and 488 isofemale lines were screened for the F₁ generation in 2007, 2008 and 2009, respectively. Resistance gene frequency in Xinxiang fluctuated between 0.0000 and 0.0005, and it did not increase significantly from 2007 to 2009. Based on the relative average development rates (RADR) of H. armigera larvae in F₁ tests, no substantial increase in Cry1Ac tolerance was found in the Xinxiang region over the 3-yr period.     

Interaction between host plant resistance and biological activity of Bacillus thuringiensis in managing the pod borer Helicoverpa armigera in chickpea

The legume pod borer Helicoverpa armigera (Hubner) (Lepidoptera: Noctuidae) has developed high levels of resistance to conventional insecticides, and therefore, efforts are being made to develop transgenic chickpea expressing toxin genes from the bacterium Bacillus thuringiensis (Bt) for controlling this pest. However, there is an apprehension that acid exudates in chickpea might interfere with the biological activity of Bt. Therefore, we studied the biological activity of Bt (Biolep^R) on four chickpea genotypes with different levels of resistance to H. armigera under field conditions, and by incorporating lyophilized leaf and pod tissue into the artificial diet with and without Bt. The pH of the acid exudates varied from 2.1 to 2.9, and malic and oxalic acids were the major components of the acid exudates in different chickpea genotypes. There was no survival of H. armigera larvae in chickpea plants sprayed with 0.1, 0.2 and 0.5% Bt. There was a significant reduction in larval survival, larval and pupal weights and fecundity, and prolongation of larval and pupal periods in chickpea plots sprayed with Bt (0.05%) as compared to the unsprayed plots. Biological activity of Bt was lower on artificial diets with leaf or pod powder of chickpea genotypes, which might be because of a low intake of Bt toxins due to the antifeedant effects of acid exudates in the chickpea or reduction in biological activity of Bt due to the interaction of biochemical constituents in chickpea with the Bt toxins. Larval survival, larval and pupal weights, pupation and adult emergence were significantly lower on diets with leaf or pod powder of the H. armigera-resistant genotypes than on the susceptible check. Chickpea genotypes with resistance to H. armigera acted in concert with Bt to cause adverse effects on the survival and development of this insect. The results suggested that development of transgenic chickpeas expressing toxin genes form Bt will be quite effective for controlling of the pod borer, H. armigera.     

Identification and Function Analysis of enolase Gene NlEno1 from Nilaparvata lugens (St?l) (Hemiptera:Delphacidae)

The enolase [EC 4.2.1.11] is an essential enzyme in the glycolytic pathway catalyzing the conversion of 2-phosphoglycerate (2-PGE) to phosphoenolpyruvate (PEP). In this study, a full-length cDNA encoding α-enolase was cloned from rice brown planthopper (Nilaparvata lugens) and is provisionally designated as NlEno1. The cDNA sequence of NlEno1 was 1,851 bp with an open reading frame (ORF) of 1,305 bp and encoding 434 amino acids. The deduced protein shares high identity of 80–87% with ENO1-like protein from Hemiptera, Diptera, and Lepidoptera speices. The NlEno1 showed the highest mRNA expression level in hemolymph, followed by fat body, salivary gland, ovaries and egg, and showed trace mRNA levels in testis. The mRNA of NlEno1 showed up-regulated level in virulent N. lugens population Mudgo, IR56 and IR42 when compared with TN1 population. Injection of double-stranded RNA (dsRNA) of NlEno1 into the adults significantly down-regulated the NlEno1 mRNA level along with decreased eggs and offspring. Moreover, injection of NlEno1-dsRNA decreased mRNA level of Vitellogenin (Vg) gene. These results showed that the NlEno1, as a key glycolytic enzyme, may play roles in regulation of fecundity and adaptation of N. lugens to resistant rice varieties.