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The beet armyworm Spodoptera exigua (Hübner) is a serious polyphagous pest that infests vegetable crops worldwide and has rapidly developed resistance due to its long-term exposure to insecticides. The current resistance statuses to four insecticides exhibited by three field populations of beet armyworms collected in southern China from 2014 to 2018 were investigated. Monitoring data from five consecutive years demonstrated that all three tested S. exigua populations developed extremely high resistance to chlorantraniliprole in 2018 (220.58- to 2,597.39-fold). Two populations (Baiyun and Fengxian) developed low to moderate resistance to spinosad, whereas the Huangpi population remained susceptible (except in 2014, with RR of 6.11-fold). The RR of the Fengxian and Baiyun populations to indoxacarb steadily increased over the years, whereas that of the Huangpi population increased relatively slowly. The Baiyun and Fengxian populations developed moderate to high resistance to indoxacarb and methoxyfenozide, whereas the Huangpi population exhibited susceptibility to low resistance (1.06- to 6.45-fold) to indoxacarb and susceptibility to moderate resistance (1.53- to 14.22-fold) to methoxyfenozide. These results suggest that chlorantraniliprole should not be employed to control this pest in southern China. Reduced use of indoxacarb and methoxyfenozide or the use of alternating insecticides with low levels of resistance is recommended. Spinosad remains an effective insecticide for the management of S. exigua. To avoid the rapid development of insecticide resistance, rotations of insecticides with low levels of resistance and different modes of action based on the resistance patterns of S. exigua should be performed in southern China.

Brassinosteroids (BRs) and jasmonates (JAs) regulate plant growth, development, and defense responses, but how these phytohormones mediate the growth-defense tradeoff is unclear. Here, we identified the Arabidopsis (Arabidopsis thaliana) dwarf at early stages1 (dwe1) mutant, which exhibits enhanced expression of defensin genes PLANT DEFENSIN1.2a (PDF1.2a) and PDF1.2b. The dwe1 mutant showed increased resistance to herbivory by beet armyworms (Spodoptera exigua) and infection by botrytis (Botrytis cinerea). DWE1 encodes ROTUNDIFOLIA3, a cytochrome P450 protein essential for BR biosynthesis. The JA-inducible transcription of PDF1.2a and PDF1.2b was significantly reduced in the BRASSINOSTEROID INSENSITIVE1-ETHYL METHANESULFONATE-SUPPRESSOR1 (BES1) gain-of-function mutant bes1- D, which was highly susceptible to S. exigua and B. cinerea. BES1 directly targeted the terminator regions of PDF1.2a/PDF1.2b and suppressed their expression. PDF1.2a overexpression diminished the enhanced susceptibility of bes1- D to B. cinerea but did not improve resistance of bes1- D to S. exigua. In response to S. exigua herbivory, BES1 inhibited biosynthesis of the JA-induced insect defense-related metabolite indolic glucosinolate by interacting with transcription factors MYB DOMAIN PROTEIN34 (MYB34), MYB51, and MYB122 and suppressing expression of genes encoding CYTOCHROME P450 FAMILY79 SUBFAMILY B POLYPEPTIDE3 (CYP79B3) and UDP-GLUCOSYL TRANSFERASE 74B1 (UGT74B1). Thus, BR contributes to the growth-defense tradeoff by suppressing expression of defensin and glucosinolate biosynthesis genes.

Metaflumizone is a novel semicarbazone insecticide. It functions as a sodium channel blocker insecticide (SCBI) with excellent insecticidal activity on most economically important lepidopterous pests. This study assessed the resistance risk of Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) to metaflumizone in the laboratory and the effects of metaflumizone selection on toxicities to other insecticides. Spodoptera exigua collected from a field population at Huizhou in 2012 were successively challenged by metaflumizone to evaluate the risk of resistance evolution. Twelve generations of selection increased resistance to metaflumizone by 3.4-fold and threshold trait analysis revealed that the realized heritability (h2) of this resistance was 0.086. When h2 was equal to 0.086 and 90% of individuals were killed at each generation, LC50 to metaflumizone increased by 10-fold after 15 generations. The selection by metaflumizone did not increase the resistance to indoxacarb, chlorantraniliprole, spinosad, methomyl, or endosulfan, suggesting a lack of cross-resistance. However, metaflumizone challenge upheld the recession of resistance to emamectin benzoate, chlorfluazuron, and tebufenozide. The block of resistance drops by metaflumizone exposure implied a possible cross-resistance between metaflumizone and these three insecticides. These results contribute to integrated resistance management of S. exigua.

Aims Spodoptera exigua is a serious pest of many agricultural crops. However, the bacterial communities of S. exigua are poorly studied, particularly over their entire life cycle. We aimed to study the biodiversity of the microbiota across the life cycle of S. exigua and to provide a better and obtain insight into new pest control strategies. Methods and results The bacterial diversity across the life cycle of S. exigua was studied using Illumina MiSeq sequencing of 16S rRNA genes. Spodoptera exigua is dominated by Proteobacteria and Firmicutes, with a total relative abundance of 90·03%. Enterococcus (24·6%), Pseudomonas (12·2%) and Asaia (45·9%) were abundant and active in eggs, while Methylobacterium (18·7%) and Halomonas (16·5%) dominated freshly eclosed larvae. The 3rd and 5th instar larvae were dominated by Enterococcus (76·3 and 62·0%). Pupal stages had the highest microbial diversity. There was no significant difference between newly emerged males and females. Symbionts of eggs were extremely similar and probably vertically transmitted by males during mating. Conclusions The result showed that the bacterial community was affected by the host developmental stages. Our results also suggest that symbionts of egg mass are probably vertically transmitted control by male spawning adults. Significance and impact of the study Our study documents the symbiont bacteria across the life cycle of S. exigua. Understanding the microbial symbionts may provide clues to develop potential biocontrol techniques against this pest.

Rice leaffolders are important pests on rice in Asia, Oceania, and Africa, causing serious loss to rice production. There are two main rice leaffolders in China, namely Cnaphalocrocis medinalis (Guenée) and C. exigua (Butler) with the former having the ability of long-distance migration. To reveal the differences in the mitochondrial genomes (mitogenome) between them, we compared the completed mitogenome of C. exigua with three C. medinalis individuals. Although phylogenetic analysis based on the mitogenomic data strongly supported the close relationship between these two species, many differences were still being revealed. The results showed that the mitogenome of C. exigua was shorter in length (15,262 bp) and slight lower in AT content than that of C. medinalis. Except for the different start codons of nad3 and nad6 gene, we also found the cox1 gene had a typical start codon ‘ATG’ which suggested that the starting position of this gene must be reconsidered in the entire superfamily Pyraloidea. All tRNAs have a typical clover-leaf structure, except for the dihydrouridine (DHU) stem losing of trnS1, which has the atypical anticondon ‘TCT’ instead of ‘GCT’ in C. medinalis and most Pyraloidea species. Two intergenic regions (between trnY and cox1, nad3 and trnA) featured by AT repeats were only found in C. medinalis and even rarely appeared in reported Pyraloidea species. Furthermore, regardless of interspecific comparison or intraspecific comparison of these two species, protein coding genes, especially the atp8 genes, had quite different evolutionary rates.

Spodoptera exigua (Hübner) is a polyphagous pest that shifts its population to different hosts during its life cycle to receive nutritive advantages. Therefore, demographic evaluation of alternate hosts is important for effective pest management. Here, we have evaluated castor (Ricinus communis L.), cauliflower (Brassica oleracea L.), cotton (Gossypium hirsutum L.), okra (Abelmoschus esculentus L.), and spinach (Spinacia oleracea L.) for growth, survival, and population development of S. exigua. Development of early populations of S. exigua is best supported on castor where earlier instars had least mortalities (10%) compared with spinach (36%), although later instars and pupae had significantly higher mortalities (20.8%) on it. Spinach and okra, on the other hand, promote larval survivals in later instars. Little or no differences in stadia lengths were observed during early development of larvae and, interestingly, the longevity of female moths increased significantly when reared on castor, cauliflower, and spinach (12.3, 11.3, and 11.7 d, respectively), resulting into significantly higher fecundity. The survival curves of all five populations have clearly demonstrated a steep early decline in larval numbers when reared on okra and only 60% larvae could survive. These findings conclude that S. exigua when fed on spinach was greatly disadvantaged in terms of growth and development; hence, the pest's field population can be opportunistically controlled by spraying adjacent spinach fields. In addition, the results highlight the vulnerable stages in pest's life cycle in the field where we can apply integrated control strategies for its effective management.

In insects, the mode of Cry1A toxins action has been studied in detail and many receptors that participate in the process are known. Recent evidence has revealed that an ABC transporter (ABCC2) is involved in conferring resistance to Cry1A toxins and that ABCC2 could be a receptor of Cry1A. However, it is not known whether Cry1Ca interacts with the same receptor proteins as Cry1A. In this study, we report the cloning of an ABC transporter gene, SeABCC2b, from the midgut of Spodoptera exigua (Hübner) larvae, and its expression in Sf9 cells for a functional analysis. The addition of Cry1Ca and Cry1Ac to Sf9 cell culture caused swelling in 28.5% and 93.9% of the SeABCC2-expressing cells, respectively. In contrast, only 7.4% and 1.3% of the controls cells swelled in the presence of Cry1Ca and Cry1Ac. Thus, SeABCC2b-expressing Sf9 cells had increased susceptibility to Cry1Ca and Cry1Ac. Similarly, S. exigua cadherin (SeCad1b) expressed in Sf9 cells caused 47.1% and 1.8% of the SeCad1b-expressing cells to swell to Cry1Ca and Cry1Ac exposure. Therefore, Sf9 cells expressing SeCad1b were more sensitive to Cry1Ca than Cry1Ac. Together, our data suggest that SeABCC2b from S. exigua mediates Cry1Ac cytotoxicity and, in conjunction with SeCad1b, contributes to enhance Cry1Ca toxicity in Sf9 cells.

Spodoptera exigua (Hübner) is a very serious worldwide pest capable of causing severe economic losses in numerous agricultural crops. The need for an effective, highly virulent, pathogenic microorganism for use as a biological control agent against S. exigua larvae is particularly important. Heliothis virescens ascovirus 3 h (HvAV-3h)-containing hemolymph with a titer of 9.58 × 10(12) genome copies per ml was used to inoculate S. exigua larvae per os with a 1.06 × 10(10) dosage per larva for the first- to second instar and 9.58 × 10(9) genome copies per larva for the third- to fifth instars. Intrahemocoelic injections were also used with a dosage of 1.53 × 10(9) genome copies per larva for third- to fifth instar. The postinjection mortality, body weight, and food intake of the S. exigua larvae were observed and recorded. The corrected mortality rates for the first- through fifth instar inoculated per os were 21.88 ± 0.98, 22.22 ± 4.00, 8.89 ± 4.01, 6.66 ± 3.33, and 8.89 ± 2.94%, respectively. The early instars were significantly easier to infect with virus compared to the later instar. The corrected mortality of the third, fourth, and fifth instars inoculated by injection was 96.58 ± 3.42, 98.83 ± 1.17, and 97.78 ± 2.22%, respectively. Compared to the healthy larval population, survival time of the diseased larval population was considerably extended. In addition, food intake was greatly reduced, and the body weight remained fairly constant in the third- and fourth instar. The body weight declined in the fifth instar corresponding to a reduction in food intake.

A viral gene, CpBV-CST1, was identified from a polydnavirus Cotesia plutellae bracovirus (CpBV). Its protein product was significantly toxic to lepidopteran insects. This study generated a transgenic tobacco plant expressing CpBV-CST1 Expression of transgene CpBV-CST1 was confirmed in T1 generation (second generation after transgenesis) in both mRNA and protein levels. Young larvae of Spodoptera exigua (Hübner) suffered high mortalities after feeding on transgenic tobacco. All 10 T1 transgenic tobacco plants had no significant variation in speed-to-kill. In order to further explore insect resistance of these transgenic tobaccos, bioassays were performed by assessing antixenosis and antibiosis. S. exigua larvae significantly avoided T1 plants in a choice test. Larvae fed with T1 plant exhibited significant decrease in protease activity in the midgut due to consuming CpBV-CST1 protein produced by the transgenic plant. Furthermore, the transgenic tobacco exhibited similar insect resistance to other tobacco-infesting insects, including a leaf-feeding insect, Helicoverpa assulta, and a sap-feeding insect, Myzus persicae These results demonstrate that a viral cystatin gene can be used to develop insect-resistant transgenic plant, suggesting a prospective possibility of expanding the current transgenic approach to high-valued crops.

The joint toxicity of chlorantraniliprole, a novel insecticide that acts on ryanodine receptors, and sinigrin, a natural plant defense compound from brassicaceous vegetables, to the larvae of Spodoptera exigua (Hübner) was determined in this paper. Additionally, the joint effects of the two compounds on cytochrome P450 enzyme activity and on the expression levels of mRNA of three P450 genes (including CYP9A9, CYP6B, and CYP4G37) and an NADPH cytochrome P450 reductase gene (HQ852049) were investigated. The toxicity of the mixture of chlorantraniliprole and sinigrin to fourth-instar S. exigua larvae was 1.60-fold higher than the toxicity of the chlorantraniliprole-only treatment after 24 h. Induced by chlorantraniliprole and sinigrin, the specific activity of the P450 O-deethylase was affected in a time-, dose-, and organ-specific manner in fifth-instar S. exigua larvae. The effects were more pronounced in the midgut than in the fat body. The specific activity of the P450 O-deethylase in almost all treatments increased at 12, 24, and 36 h posttreatment compared with that in the control. Based on real-time PCR analyses, the expression levels of the P450 genes CYP9A9, CYP6B, and CYP4G37 and the NADPH cytochrome P450 reductase gene HQ852049 in fifth-instar S. exigua larvae were induced by chlorantraniliprole and sinigrin, and the trends were similar to the specific activity of the P450 O-deethylase. Therefore, the CYP9A9, CYP6B, and HQ852049 in the tested genes were the most inducible genes that were expressed when the S. exigua larvae were exposed to chlorantraniliprole and sinigrin.

Beet armyworm, Spodoptera exigua (Hübner), is a worldwide pest of many crops. Chemical insecticides are heavily used for its control in China, and serious resistance has been evolved in the field to a variety of insecticides including emamectin benzoate. Through repeated backcrossing to a susceptible strain (WH-S) and selection with emamectin benzoate, the trait conferring resistance to emamectin benzoate in a field-collected population of S. exigua (moderately resistant to emamectin benzoate and strongly resistant to pyrethroids and indoxacarb) was introgressed into WH-S to generate a near-isogenic resistant strain (WH-EB). Compared with WH-S, the WH-EB strain developed a 1,110-fold resistance to emamectin benzoate and a high level of cross-resistance to abamectin (202-fold), with low levels of cross-resistance to cypermethrin (10-fold) and chlorfluazuron (7-fold), but no cross-resistance to representatives of another six different classes of insecticides (chlorantraniliprole, chlorfenapyr, indoxacarb, spinosad, tebufenozide, and chlorpyrifos). Resistance to emamectin benzoate in WH-EB was autosomal, incompletely dominant, and polygenic. Limited cross-resistance in WH-EB indicates that emamectin benzoate can be rotated with other classes of insecticides to which it does not show cross-resistance to delay the evolution of resistance in S. exigua. The incompletely dominant nature of resistance in S. exigua may explain the rapid evolution of resistance to emamectin benzoate in the field, and careful deployment of this chemical within a resistance management program should be considered.

Nowadays, heavy metal pollution has become more and more serious. Cadmium is one of the heavy metals that have caused serious pollution. Spodoptera exigua Hubner (Lepidoptera: Noctuidae), one of important agricultural pests, has broken out more frequently in recent years. In this study, accumulation at different developmental stages and excretion of Cd were detected, and effect of cadmium (Cd concentrations were 0.2, 0.8, 3.2, 12.8, and 51.2 mg/kg, respectively) on three successive generations of beet armyworm was also studied using life table in the laboratory. Results showed that Cd2+ contents were significantly different when S. exigua was reared under different Cd2+ concentrations in the same generation in a significant dose-dependent manner. The concentration of Cd2+ increased with extension of stress time. Cd2+ concentration at different developmental stages was different, that is, larvae > pupa > adult. After emergence, Cd2+ concentration in both females and males decreased gradually, but that in paired adults decreased to a greater extent than that in unpaired ones. Beet armyworm could excrete heavy metals by means of faeces, prepupa exuviate, and puparium. Life table of the laboratory population was constructed, which showed that low doses of cadmium promoted the increase of population, whereas high doses inhibited the increase, meanwhile, with the extension of exposure length, the index of population trend (I) declined more rapidly. The study might provide references for the long-term and objective assessment of heavy metal pollution and its effect on the populations of important agricultural pests.

As a selective biological insecticide, spinosad has been used widely for the control of pests including beet armyworm, Spodoptera exigua (Hubner) (Lepidoptera: Noctuidae). To form effective pest control strategies, lethal and sublethal effects should be considered for a complete analysis of spinosad impact. However, few studies have been reported to investigate sublethal effects of spinosad on S. exigua. This study attempts to evaluate the lethal and sublethal effects of spinosad on this pest by recording and analyzing various toxicological and physiological parameters. The toxicity of spinosad against S. exigua was determined under laboratory conditions by oral exposure of late second-instar larvae to the compound. The LC50 values of spinosad to S. exigua at 48 and 72 h after treatment were 0.317 and 0.293 mg x kg(-1), respectively. Spinosad at sublethal concentrations significantly extended the developmental period of survivor larvae, and reduced larval wet weight. Postexposure effects were indicated by decreased pupation ratio and pupal weight, by prolonged prepupal and pupal periods and by decreased emergence ratio, fecundity and longevity of adults. The net replacement rate (Ro) tended to be lower in the exposed spinosad groups than those in the unexposed spinosad group. Intrinsic rate of population increase (r(m)) for the high-dose group (0.365) was significantly lower than the control (0.521) and the low-dose group (0.521), but the latter two were not significantly difference. These results suggest that the combination of lethal and sublethal effects of spinosad might affect S. exigua population dynamics significantly by decreasing its survival and reproduction, and by delaying its development.

Beet armyworm, Spodoptera exigua (Hubner), is a serious pest of vegetables in China, and its control is heavily dependent on chemical insecticides. The current resistance status of nine insecticides was investigated in 16 field populations collected from seven provinces of China during 2009–2012. Compared with the susceptible strain WH-S, some field populations evolved various levels of resistance to eight of the nine insecticides tested: emamectin benzoate (4- to 348-fold), indoxacarb (2- to 41-fold), spinosad (5- to 38-fold), chlorantraniliprole (2- to 44-fold), tebufenozide (2- to 87-fold), chlorfluazuron (3- to 31-fold), cypermethrin (79- to 1240-fold), and chlorpyrifos (8- to 3,080-fold), but no significant resistance was detected to chlorfenapyr (0.4- to 7-fold). This indicates that chlorfenapyr has no cross-resistance with these other currently used insecticides. Four consecutive years’ resistance screening at two places shows that resistance patterns were different between populations from Luhe (Jiangsu Province) and Fengxian (Shanghai), which are ≈300 km apart. Resistance levels to chlorpyrifos were much higher in populations from Luhe (877- to 3,080-fold) than from Fengxian (8- to 110-fold). Fengxian populations developed moderate levels of resistance to tebufenozide (13- to 87-fold), but no resistance in Luhe populations (2- to 6-fold). However, Luhe populations developed moderate levels of resistance to chlorfluazuron (21- to 31-fold), but there was no resistance in Fengxian populations (3- to 5-fold). It is suggested that local insecticide selection determined resistance patterns although S. exigua has long-distance migratory potential. Adaptive resistance management tactics (such as rotations) should be designed and implemented based on the resistance patterns of S. exigua for each geographic area.

Many studies have reported the effect of nitrogen (N) fertilization on plant constitutive defense responses; however, little is known about their effects on plant induced defense patterns and its consequence for insect herbivores. In our experiments, the effects of N availability on growth, nutritional quality (N content, protein/carbohydrate [P:C] ratio, modified gross energy [MGE]), and constitutive phenolics of corn, Zea mays L. were quantified. Moreover, the indirect effects of N fertilization on the beet armyworm, Spodoptera exigua Hübner through larval performance and feeding preference were examined. N fertilization increased plant growth, and depressed defense traits by increasing N content and the P:C ratio, as well as decreasing the constitutive concentration of phenolics. Subsequently, beet armyworm showed higher performance and preferentially fed on high-N corn because of its low defense traits. After beet armyworm feeding, high-N corn significantly deterred larval feeding, and had negative effects on the performance of beet armyworm through decreasing P:C ratio and increasing induced phenolics. On the contrary, there were no significant changes in P:C ratio and phenolics in low-N corn after feeding damage. Larval performance and preference were also not affected by induced compounds in low-N corn, which suggested that the expression of induced defense was dependent on N availability. The result indicates that N availability can exert a variety of bottom-up effect on plant defense patterns to influence insect population dynamics, and thereby may represent a source of variation in plant-insect interactions.

Being delivered as sprays or expressed in plant, Bacillus thuringiensis (Bt) crystalline proteins (Cry toxins) display insecticidal activities against numerous Lepidopteran, Dipteran, and Coleopteran larvae. Comparative study of toxicities of Bt Cry toxins between larvae and adults may afford important new insights into the interactions of the toxins with receptor proteins in host insect, and represent intriguing targets for the control of insect pests. However, the effectiveness of Bt Cry toxins in insect adults has paid less attention. In the present article, the effectiveness of Cry1Ac and Cry1Ca on lifespans and reproductive performance of Helicoverpa armigera (Hubner) and Spodoptera exigua (Hubner) adults were evaluated by in vivo experiments. Considering transgenic plants express modified, truncated versions of cry genes yielding active toxin fragment, we used activated Bt toxins at the concentration of 500, 100, and 20 microg/ml in a 10% sucrose aquous solution. At the highest concentration, Cry1Ac and Cry1Ca shortened 48.1 and 48.9% of H. armigera female lifespan, and 43.5 and 38.5% of S. exigua female lifespan, and they reduced 37.8 and 40.3%, and 50.5 and 47.4% of H. armigera and S. exigua male lifespans respectively. Bt toxins negatively affected copulation. Exposure to 500 microg/ml of Cry1Ac and Cry1Ca greatly reduced 50.0 and 46.8%, and 58.7 and 57.3% spermatophore acceptance by H. armigera and S. exigua females, respectively. Similarly, Cry1Ac and Cry1Ca exposure decreased 40.0 and 50.3%, and 61.3 and 60.0% of spermatophore transfer by H. armigera and S. exigua males, respectively. Moreover, exposure females rather than males to 500 microg/ml of Cry1Ac and Cry1Ca significantly dropped 57.5 and 57.5% of the number of eggs laid by H. armigera, and 35.4 and 45.8% of the number of egg masses deposited by S. exigua. In contrast, both Cry1Ac and Cry1Ca did not negatively influence the egg hatchability. At the middle and the lowest concentrations, however, the biological effects of Cry1Ac and Cry1Ca on lifespans and reproductive performance partially or completely disappeared.

Baculoviruses may interact with parasitoids in the same host. A previous study has shown that infection of larvae with Spodoptera litura nucleopolyhedrovirus (SpltNPV) was deleterious to the survival and development of Meteorus pulchricornis (Wesmael) (Hymenoptera: Braconidae). In this paper, the interactions between M. pulchricornis and Spodoptera exigua multiple nucleopolyhedrovirus (SeMNPV) in Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae), a permissive host of the virus and parasitoid, were investigated. The results showed that the effect of M. pulchricornis on SeMNPV and the effect of the virus on the parasitoid both depended on the concentration of the virus and the interval between viral infection and parasitism. Whether S. exigua was treated with the parasitoid and virus simultaneously or 1 day apart, the biological activities of 10(5), 10(6), and 10(7) OBs/mL SeMNPV were all significantly improved by M. pulchricornis. In contrast, the biological activity of 10(3) OBs/mL SeMNPV was significantly decreased when the host was exposed to the virus and parasitoid simultaneously. Regarding the impact of SeMNPV on M. pulchricornis, exposing the host to the parasitoid and SeMNPV with concentrations lower than 10(6) occlusion bodies (OBs)/mL produced no negative effects on the parasitoid. The results also showed that ingestion of SeMNPV by adult stage M. pulchricornis significantly increased the number of parasitoid offspring that successfully emerged from the host. Furthermore, M. pulchricornis was found to transmit SeMNPV among populations of S. exigua. Taken together, these findings indicate that M. pulchricornis integrated with an appropriate concentration of SeMNPV has the potential to improve the efficacy of biological control against S. exigua.

The insecticidal characteristics of two Spodoptera exigua multiple nucleopolyhedrovirus (SeMNPV) strains produced on two different S. exigua colonies were measured using the same two host colonies. These strains constitute the active ingredients of the biological insecticides Vir-ex and Spexit and were produced on insect colonies from Spain and Switzerland. Demographic characteristics of insects from each colony were examined before infection. Larval developmental time, larval survival, and adult sex ratio did not differ between the colonies, whereas mean pupal weight was significantly higher in the Spanish colony insects. After infection, susceptibility to virus occlusion bodies (OBs), time to death, larval weight at death, and total production of OBs/larva varied significantly depending on virus strain and the colony used. Vir-ex OBs produced in Spanish colony larvae had improved insecticidal characteristics in terms in lethal dose and speed of kill metrics than other strain-colony combinations. OB production was significantly higher in Spanish colony insects infected with Spexit compared with Vir-ex infected insects from the Swiss colony, with intermediate values for the other two strain-colony combinations. Virus strain and host colony origin were highly influential in determining the insecticidal characteristics of OBs and should be considered as key parameters that require optimization during the production of SeMNPV-based insecticides.

Baculoviruses and parasitoids are important biological control factors of insects in integrated pest management. Microplitis bicoloratus Chen (Hymenoptera: Braconidae: Microgastrina) is a solitary endoparasitoid of beet armyworm, Spodoptera exigua (Hubner) (Lepidoptera: Noctuidae) larvae. M. bicoloratus parasitized first to third instars of S. exigua and most effectively parasitized second instars. The survival rate of parasitoids emerging from S. exigua multiple nucleopolyhedrovirus (SeMNPV)-infected hosts decreased with increasing SeMNPV doses at second to fourth instars. In addition, the mortality rate of the host was >80% when coinfected with SeMNPV and M. bicoloratus, regardless of virus doses or the timing of virus treatment. Occlusion body production was significantly reduced in parasitized hosts. A comparison of host weights showed that hosts coinfected with SeMNPV and M. bicoloratus were significantly lighter than those infected with SeMNPV alone, suggesting that the decrease in virus yield resulted from a reduction in larval growth. The viral genome copy number in parasitized host was significantly lower than that in nonparasitized host at 48 and 72 h postinfection. These results suggest that SeMNPV and M. bicoloratus are compatible as S. exigua control agents.

We studied the effect of combining microbial pesticides with camptothecin (CPT) on the mortality of two lepidopteran insects: Trichoplusia ni (Hubner) and Spodoptera exigua (Hubner). CPT is an alkaloid that is often used as an anticancer agent. Here, CPT was evaluated as a microbial pesticide synergist of Bacillus thuringiensis (Bt) and insect baculovirus. The toxicity of CPT and its synergistic effects on two microbial pesticides were studied using the diet overlay method. Bioassay results showed that CPT significantly enhances the toxicity of Bt variety kurstaki to S. exigua and T ni. In addition, CPT strongly enhanced the infectivity of Autographa californica (Speyer) multinucleocapsid nucleopolyhedrovirus (AcMNPV) and S. exigua nucleopolyhedrovirus (SeMNPV). Using light microscopy, we found that CPT disrupts the peritrophic membrane of T. ni larvae and severely affects the structure of the midgut, resulting in an abnormal gut lumen morphology. We speculate that CPT increases toxicity by affecting the permeability of the peritrophic membrane.

Life table, reproduction, and population growth parameters of the beet armyworm, Spodoptera exigua (Hubner) (Lepidoptera: Noctuidae), were determined on nine soybean (Glycine max L.) cultivars (‘032’, ‘033’, ‘Hill’, ‘M4’, ‘M7’, ‘M9’, ‘M11’, ‘TMS’, and ‘Zane’) at 25°C. The age-specific survival rate of eggs and immature stages was higher on 033 in comparison with other cultivars. The gross fecundity rate ranged from 911 on M9 to 1,296 on Zane, but the differences were not statistically significant. The net fecundity rate was significantly higher on 033 (1,082 eggs per female) than on other cultivars. The gross fertility rate significantly differed on tested cultivars, and was the highest on Zane (1,257 eggs per female) and lowest on Hill (813 eggs per female). The net fertility rate varied from 532 on M11 to 1,082 eggs per female on 033. Both daily number of eggs and daily number of fertile eggs laid per female varied significantly among the different cultivars, and were the highest on TMS (90 for both parameter) and lowest on M7 (47 and 43), respectively. The values of net reproductive rate, intrinsic rate of increase, and finite rate of increase were significantly higher on 033. The shortest generation time (23 d), shortest doubling time (1.66 d), and highest percentage of female offspring (56%) also were obtained on 033. According to population growth parameters, 033 and Hill partially were susceptible and resistant, respectively, to S. exigua. The results of this study provide direction to design a more comprehensive integrated pest management program for this pest.

The beet armyworm, Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae), is an important pest of numerous crops, and it causes economic damage in China. Use of secondary metabolic compounds in plants is an important method used to control this insect as a part of integrated pest management. In this study the growth, development, and food utilization of three successive generations of S. exigua fed on three cotton gossypol cultivars were examined. Significantly longer larval life-spans were observed in S. exigua fed on high gossypol cultivar M9101 compared with those fed on two low gossypol cultivars, ZMS13 and HZ401. The pupal weight of the first generation was significantly lower than that of the latter two generations fed on ZMS13 group. Significantly lower fecundity was observed in the second and third generations of S. exigua fed on M9101 compared with S. exigua fed on ZMS13 and HZ401. The efficiency of conversion was significantly higher in the first and third generations fed on HZ401 compared with those fed on ZMS13 and M9101. A significantly lower relative growth rate was observed in the three successive generations fed on M9101 compared with those fed on ZMS13 and HZ401. Cotton cultivars significantly affected the growth, development, and food utilization indices of S. exigua, except for frass and approximate digestibility. Development of S. exigua was significantly affected by relative consumption rate and efficiency of conversion of ingested food, but not by relative growth rate or approximate digestibility, suggesting that diet-utilization efficiency was different based on food quality and generation. Measuring the development and food utilization of S. exigua at the individual and population levels over more than one generation provided more meaningful predictions of long-term population dynamics.

The toxicity and sublethal effects of methoxyfenozide were evaluated in third instars of the beet armyworm, Spodoptera exigua (Hubner) (Lepidoptera: Noctuidae), that fed on contaminated semisynthetic diet. The LC50 value was estimated at 0.23 mg of active ingredient (AI)/kg diet (range, 95% CI: 0.17–0.37) at 264 h after treatment. The effects on development, survival, and reproduction were observed in third instars of this pest that survived exposure to an LC25 concentration of methoxyfenozide. The larvae from the insecticide treatment exhibited lower pupal weights, an increase in both larval and pupal developmental times and a higher frequency of malformations of the wings in adults than untreated larvae. Adults from the methoxyfenozide treatment did not show reduced fecundity (mean cumulative number of eggs laid per female), but fertility as measured by the percentage of eggs hatched (fertility) was significantly reduced compared with untreated control insects. No significant effects were observed on pupal sex ratio. We concluded that the lethal and sublethal effects of methoxyfenozide are likely to have a significant impact on S. exigua populations on treated crops.

A low-cost simplified diet has been successfully developed for rearing Spodoptera exigua larvae under laboratory conditions. The cost of ingredients was lower than that of the standard diet based on a modified tobacco hornworm, Manduca sexta (L.), diet. The simplified diet fulfilled larval nutritional requirements without apparent adverse effects on the reproductive capacity of the insect. Survival, pupal sex ratio, and fecundity registered in insects that were reared on the simplified diet did not differ from those observed on the standard diet. The mean larval development period of insects that consumed the simplified diet was also similar to that of insects that consumed the standard diet, whereas weight of pupae and adult longevity were significantly higher in insects reared on simplified diet. Larvae consumed ≈11% more of the standard diet compared with the simplified diet and a corresponding increase was observed in the number of larvae that could be reared through to pupation on each liter of simplified diet. The production of S. exigua multiple nucleopolyhedrovirus (SeMNPV) occlusion bodies (OBs) in insects grown on each type of diet was also evaluated. Weights of larvae at inoculation and at death, OB yields and biological activity of OBs did not differ significantly for each type of diet. The simplified low-cost diet appears suitable for large-scale in vivo production of SeMNPV OBs.

Spodoptera exigua is one of the most renowned agricultural pest insects and relatively insensitive to Bacillus thuringiensis subsp. kurstaki strains which are widely used commercial products to control lepidopterans such as Heliothis armigera. In the current study, we have developed a new and efficient approach to screen and breed a B. thuringiensis subsp. kurstaki strain exhibiting high toxicity against S. exigua while retaining its high toxicity against H. armigera. UV and diethyl sulfate methods were used for mutagenesis, followed by an agar plug plate diffusion assay for preliminary screening of Zwittermicin A over-producing mutants, from which we obtained a mutant strain, designated here as B. thuringiensis subsp. kurstaki D1-23, with high toxicity against S. exigua. The toxicity of D1-23 against S. exigua and H. armigera was improved by 115.4 and 25.9%, respectively, compared to its parental commercial strain BMB005.

A Bacillus thuringiensis (Bt)-based formulation intended to increase feeding response in Spodoptera exigua (Hubner) (Lepidoptera: Noctuidae) was developed. Eight matrices were prepared from natural ingredients and confronted with fresh maize leaves to select one highly palatable to larvae. The selected matrix was combined with Bt HD-125 strain at three different concentrations (30, 70, and 100 g/kg) of the spore-crystal complex to elaborate three insecticidal formulations. Effects caused by the matrix, including palatability, any adverse effect, and reduction in toxic activity of the strain, were tested on the formulations. Results indicated that the blend of modified cornstarch, porcine gelatin, and dried corncob was the most preferred by larvae. Formulations obtained were evaluated in mortality bioassays with neonate larvae in laboratory, using artificial diet, and in the greenhouse, with maize, Zea mays L., plants to compare their effectiveness when applied to natural food. In laboratory tests, all formulations, including a commercial formulation, were ingested by larvae causing high mortality in neonate larvae; in the greenhouse tests, formulations were also ingested causing mortality in larvae, overcoming commercial formulation effectiveness. Results suggest that the selected matrix combined with Bt 70 g/kg is the most suitable formulation to achieve effective control because this concentration acts faster than the lower concentration and is equal than the higher concentration. This formulation, designed specifically against a particular pest, is highly effective and offers a high potential for effective pest control.

(-)-10-epi-Axisonitrile-3, a spirocyclic sesquiterpene isocyanide obtained from the marine sponge Geodia exigua, immobilized sperm of sea urchin and starfish to block fertilization at the minimum effective concentration of 0.4 microg/ml. On the other hand, fertilized eggs developed normally to the gastrula stage in the presence of a 250-times higher concentration of the isocyanide. Analysis by (31)P NMR revealed an accumulation of phosphocreatine and a depletion of inorganic phosphate in the isocyanide-treated sperm, suggesting that (-)-10-epi-axisonitrile-3 inhibited the phosphocreatine shuttle participating in the high-energy phosphate metabolism, thereby immobilizing sperm to block fertilization. No analogs of (-)-10-epi-axisonitrile-3 containing different functionalities or isocyanides with different carbon skeleton exhibited such activity.

Large numbers of a cockroach that resembled the German cockroach, Blattella germanica (L.), were observed during winter 2005–2006 in home turf settings in Weslaco, TX, 11 km from the U.S. border with northeastern Mexico. In June and July 2006, this cockroach was observed at night on the foliage of soybean, Glycine max (L.) Merr., in numbers often exceeding 50 per m row. Because of its similarity to B. germanica, but with the characteristic of flying frequently, we suspected that our observations might be of the Asian cockroach, Blattella asahinai Mizukubo. Using a combination of morphological characters, cuticular hydrocarbons, and sequences of the mitochondrial cytochrome oxidase II gene, we definitively identified this cockroach as B. asahinai. B. asahinai was frequently observed feeding on sentinel eggs of Helicoverpa zea (Boddie) and Spodoptera exigua (Hubner). This cockroach was the dominant egg predator in soybean during 2006, making up 36.4% of all predators observed feeding on eggs. B. as...

Cry1Ac protoxin (the active insecticidal toxin in both Bollgard and Bollgard II cotton [Gossypium hirsutum L.]), and Cry2Ab2 toxin (the second insecticidal toxin in Bollgard II cotton) were bioassayed against five of the primary lepidopteran pests of cotton by using diet incorporation. Cry1Ac was the most toxic to Heliothis virescens (F.) and Pectinophora gossypiella (Saunders), demonstrated good activity against Helicoverpa zea (Boddie), and had negligible toxicity against Spodoptera exigua (Hubner) and Spodoptera frugiperda (J. E. Smith). Cry2Ab2 was the most toxic to P. gossypiella and least toxic to S. frugiperda. Cry2Ab2 was more toxic to S. exigua and S. frugiperda than Cry1Ac. Of the three insect species most sensitive to both Bacillus thuringiensis (Bt) proteins (including H. zea), P. gossypiella was only three-fold less sensitive to Cry2Ab2 than Cry1Ac, whereas H. virescens was 40-fold less sensitive to Cry2Ab2 compared with CrylAc. Cotton plants expressing Cry1Ac only and both Cry1Ac and Cry2Ab2 proteins were characterized for toxicity against H. zea and S.frugiperda larvae in the laboratory and H. zea larvae in an environmental chamber. In no-choice assays on excised squares from plants of different ages, second instar H. zea larvae were controlled by Cry1Ac/Cry2Ab2 cotton with mortality levels of 90% and greater at 5 d compared with 30-80% mortality for Cry1Ac-only cotton, depending on plant age. Similarly, feeding on leaf discs from Cry1Ac/Cry2Ab2 cotton resulted in mortality of second instars of S.frugiperda ranging from 69 to 93%, whereas exposure to Cry1Ac-only cotton yielded 20-69% mortality, depending on plant age. When cotton blooms were infested in situ in an environmental chamber with neonate H. zea larvae previously fed on synthetic diet for 0, 24, or 48 h, 7-d flower abortion levels for Cry1Ac-only cotton were 15, 41, and 63%, respectively, whereas for Cry1Ac/Cry2Ab2 cotton, flower abortion levels were 0, 0, and 5%, respectively. Cry1Ac and Cry2Ab2 concentrations were measured within various cotton tissues of Cry1Ac-only and Cry1Ac/Cry2Ab2 plants, respectively, by using enzyme-linked immunosorbent assay. Terminal leaves significantly expressed the highest, and large leaves, calyx, and bracts expressed significantly the lowest concentrations of Cry1Ac, respectively. Ovules expressed significantly the highest, and terminal leaves, large leaves, bracts, and calyx expressed significantly (P < 0.05) the lowest concentrations of Cry2Ab2. These results help explain the observed differences between Bollgard and Bollgard II mortality against the primary lepidopteran cotton pests, and they may lead to improved scouting and resistance management practices, and to more effective control of these pests with Bt transgenic crops in the future.

To evaluate the relationship between reproduction and migratory flight, we examined ovaries, tethered flight, and egg laying in the beet armyworm, Spodoptera exigua (Hubner). Ovarian development in female S. exigua was classified into five stages in relation to the postemergence age of the moths. Oviposition peaked at ovarian stages III and IV. Mating accelerated ovarian development in this species, and thus the mated females developed faster than the virgin moths in ovaries. In virgin adults, flight capacity increased from the first day to the third day after emergence and remained at a high level until the moths were 7 d old. In contrast, the mated moths reached the maximum flight capacity in 2 d after emergence and maintained a high level until the age of 7 d. However, mating status could not influence the flight capacity of the beet armyworm, although there were general differences in flight capacity between virgin and mated adults at the same age. Moreover, significant correlations between the number of eggs laid and the flight capacity were not performed by the mated moths. These results showed that mating status and ovarian development did not seem to influence flight capacity in the beet armyworm, which was not in agreement with the definitions of the oogenesis-flight syndrome observed in many other migratory insects.

Benzylideneacetone (BZA) is a metabolite of gram-negative entomopathogenic bacterium Xenorhabdus nematophila, and it acts as an enzyme inhibitor against phospholipase A2 (PLA2). PLA2 catalyzes a committed biosynthetic step of eicosanoids, which mediate insect immune reactions to infection by microbial pathogens. This study tested a hypothesis that a putative immunosuppressive activity of BZA may enhance virulence of Bacillus thuringiensis against the fifth instars of Spodoptera exigua (Hubner) (Lepidoptera: Noctuidae). In in vitro conditions, BZA significantly inhibited hemocyte microaggregation induced by B. thuringiensis and impaired hemocyte-spreading behavior of S. exigua in a dose-dependent manner. Oral administration of BZA gave similar immunosuppressive effect on the hemocytes of the fifth instars. Although BZA itself did not possess any insecticidal activity on oral administration, when BZA was treated in a mixture with a low dose of B. thuringiensis spp. aizawai to fifth instars, the bacterial virulence was significantly enhanced. BZA also enhanced virulence of B. thuringiensis spp. kurstaki, which alone was of limited effectiveness against S. exigua. This study suggests that an immunosuppression by BZA is positively linked to potentiation of B. thuringiensis.

The Spodoptera exigua (Hubner) multiple nucleopolyhedrovirus (SeMNPV) is currently being tested as a biological insecticide for use in greenhouse crops in southern Spain. We performed a study in which semipurified SeMNPV occlusion bodies (OBs) were formulated in phosphate-buffered saline, pH 6.5, with 5% (vol:vol) glycerol and 0.15% (wt:vol) sorbic acid, and they were stored at -20, 4, or 25 degrees C during 18 mo. Initial aerobic counts (+/-SE) averaged 1.4 (+/-0.17) x 10(7) colony-forming units/ml after 17-h incubation at 37 degrees C. Aerobic counts of microorganisms that contaminated OB formulations stored at 250C decreased markedly over the period of the study, whereas only small decreases were observed in counts from OBs stored at 4 or -20 degrees C. The principal microbial contaminants of OB suspensions were Enterococcus spp., Enterobacteriaceae, and yeasts. Potential human pathogens (Salmonella, Shigella, and Vibrio species) were not detected, and populations of Staphylococcus aureus and Bacillus cereus were extremely low. Compared with newly formulated OBs, the estimated LD50 values of OBs stored at 25 degrees C increased by >16,666-fold over the 18 mo of storage, whereas LD50 values were not greatly affected by storage at 4 or -20 degrees C. Significant changes over time in OB concentrations were only observed in the 25 degrees C treatment. Complete degradation of viral DNA was observed at 25 degrees C but not in refrigerated or frozen OBs. We conclude that OB formulation with bacteriostatic or antioxidant additives, together with storage and distribution in refrigerated conditions, will likely result in an SeMNPV biopesticide shelf life that exceeds 18 mo.

Host size is considered a reliable indicator of host quality and an important determinant of parasitoid fitness. Koinobiont parasitoids attack hosts that continue feeding and growing during parasitism. In contrast with hemolymph-feeding koinobionts, tissue-feeding koinobionts face not only a minimum host size for successful development but also a maximum host size, because consumption of the entire host is often necessary for successful egression. Here we study interactions between a generalist tissue-feeding larval endoparasitoid, Hyposoter didymator Thunberg (Hymenoptera: Ichneumonidae) and two of its natural hosts, Spodoptera exigua Hübner and Chrysodeixis chalcites Esper (Lepidoptera: Noctuidae). Larvae of C. chalcites are up to three times larger than corresponding instars of S. exigua and also attain much higher terminal masses before pupation. We hypothesized that the range of host instars suitable for successful parasitism by H. didymator would be much more restricted in the large host C. chalcites than in the smaller S. exigua. To test this hypothesis, we monitored development of H. didymator in all instars of both host species and measured survival, larval development time, and adult body mass of the parasitioid. In contrast with our predictions, C. chalcites was qualitatively superior to S. exigua in terms of the survival of parasitized hosts, the proportion of parasitoids able to complete development, and adult parasitoid size. However, in both hosts, the proportion of mature parasitoid larvae that successfully developed into adults was low at the largest host sizes. Our results suggest that qualitative, as well as quantitative, factors are important in the success of tissue-feeding parasitoids.

Xenorhabdus sp. (X. sp.) and Photorhabdus temperata subsp. temperata (Ptt) are the symbiotic bacteria of the entomopathogenic nematodes, Steinernema monticolum and Heterorhabditis megidis, respectively. When the nematodes enter target insects, the symbiotic bacteria are released into the insect hemocoel. The bacteria cause septicemia and kill the insect host. When ingested, the bacteria caused high mortality of third-instar larvae of Spodoptera exigua, but did not show such high pathogenicity to the fifth-instar larvae. The high mortalities in the third-instar larvae seem to be caused by antibiotic activity against Bacillus cereus, a gut symbiont required for optimal development of S. exigua. To increase their pathogenicity in the fifth instar, the bacteria should be delivered into the hemocoel. To this end, Bacillus thuringiensis aizawai (Bt) as a synergist was used to facilitate entry of the bacteria from the gut lumen into the hemocoel of S. exigua by its disruption of the insect gut epithelium. The bacterial mixture treatment was highly synergistic against the fifth-instar larvae of S. exigua. The synergistic effects were shown by the successful infection of X. sp. or Ptt in the insect hemocoel. This research shows a possibility that Xenorhabdus or Photorhabdus can be applied to kill S. exigua by oral treatment in a mixture with Bt.

A meridic diet was supplemented with avidin at various concentrations to determine its effects on growth and mortality of three lepidopteran insects: Helicoverpa zea (Boddie), Spodoptera exigua (Hubner), and Anticarsia gemmatalis (Hubner). All insects were placed on diet immediately after hatching and observed until death or pupation occurred. At a concentration of 10 ppm, avidin had little or no effect on growth and mortality compared with the control. However at a concentration of 100 ppm almost all tested insects were killed. H. zea was further tested by adding sublethal concentrations of Bt (Cry1Ac) in the diet containing avidin. The synergistic effect was significant, with mortality increasing to 44.4% over additive mortality (21.6%) of Bt and avidin.

DNA marker analysis has mapped a quantitative trait locus for soybean, Glycine max (L.) Merr., resistance to the corn earworm, Helicoverpa zea (Boddie) (Lepidoptera: Noctuidae), on the USDA soybean genetic linkage map near the classical gene Pb, which conditions pubescence tip. This study was initiated to determine the effect of pubescence tip on resistance to H. zea larvae and to examine the effect on beet armyworm, Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae), and soybean looper, Pseudoplusia includens (Walker) (Lepidoptera: Noctuidae), larvae. The effect of blunt (pb) and sharp (Pb) pubescence tip was tested in antixenosis and antibiosis bioassays on H. zea, S. exigua, and P. includens larvae with near-isolines and insect-resistant and -susceptible genotypes differing in pubescence tip morphology. Sharp pubescence tip significantly reduced defoliation (antixenosis) from H. zea, S. exigua, and P. includens and weight gain (antibiosis) of H. zea. The weight gain of P. includens was unaffected, and S. exigua weight gain was significant for one pair of near-isolines differing in pubescence tip but not the other. The results indicate that sharp pubescence tip would be beneficial to introgress into elite soybean germplasm due to its association with resistance to H. zea, S. exigua, and P. includens.

A new Helicosporidium spp. isolate recently purified from an aquatic weevil, Cyrtobagus salviniae Calder & Sands, was capable of infecting and reproducing in three heterologous noctuid hosts. Regardless of host species, oral treatment of Heliocoverpa zea (Boddie), Spodoptera exigua (Hubner), or Trichoplusia ni (Hubner) early instars with cyst preparations of Helicosporidium spp. resulted in ≈50% infection of the challenged larvae. The sex ratio did not differ between infected and control groups, suggesting the existence of a natural, nonsex-related resistance to the disease. Injection of Helicosporidium spp. into the hemocoels of late instars resulted in virtually 100% infection, indicating that resistance is related to the ingestion of the pathogen and therefore affiliated with midgut-mediated barriers. The pathogen’s development was not interrupted by metamorphosis; likewise, the infection did not necessarily interrupt the insects’ development. When treated as early instars, 50–90% of the infected larvae formed pupae, of which 20–30% emerged as adults. However, a high proportion of the infected adults (62–86%) had malformed wings, and their longevity was reduced compared with that of healthy adults. Infected S. exigua adults that seemed to be morphologically healthy were able to mate and produce viable offspring. The disease was detected in five of 12 groups of progeny produced by infected adults. However, the relative infection rate in the filial generation was low (2–5%). To our knowledge, this is the first evidence for a vertical transmission of helicosporidial infection.

The influence of an optical brightener, Tinopal LPW, on the activity of a purified genotype of the nucleopolyhedrovirus (SeMNPV) of the beet armyworm, Spodoptera exigua (Hubner), was determined in second to fifth instar (L2-L5) S. exigua. When mixed with viral occlusion bodies (OB) 1% Tinopal LPW significantly reduced the median lethal dose (LD50) of the virus in all instars compared with insects treated with SeMNPV alone. Levels of enhancement, as determined by LD50 values, ranged from 2.6- to 580-fold, depending on the instar. The greatest enhancement occurred on the two later instars, L4 (70-fold) and L5 (580-fold), which show a much higher resistance to SeMNPV infection than earlier instars. The median time to death (MTD) values were not significantly different in any instar among larvae treated with SeMNPV + Tinopal LPW and those treated with SeMNPV alone. Larval development in SeMNPV + Tinopal LPW treated larvae was retarded, in second and fourth instars, compared with controls or larvae tr...

In the current study, we tested the effect of peanut, Arachis hypogaea L. (Leguminoseae), stem infection by the white mold fungus, Sclerotium rolfsii Saccodes (Basidiomycetes), on the oviposition preference of beet armyworms (BAW), Spodoptera exigua Hubner (Lepidoptera: Noctuidae) and on the host-searching behavior by a BAW larval parasitoid Cotesia marginiventris. We found that, in choice tests, adult BAW oviposited more on white mold-infected plants than on healthy plants. We also found evidence that this preference is mediated by plant volatiles and other biochemical changes in plant chemistry caused by fungal infection. When plants were exposed to BAW feeding, the parasitoid C. marginiventris landed more frequently on infected than on healthy plants. We conducted wind tunnel choice experiments to determine whether the more frequent landing by the wasps was mediated by the volatiles emitted by healthy and white mold-infected plants in response to BAW damage. In these wind tunnel experiments, wasps were more responsive to volatiles from plants infected with the white mold compared with healthy ones, when both types of plants were exposed to damaged by BAW caterpillars. Thus, white mold-infected peanut plants were preferred by BAW for oviposition, but, when damaged by BAW larvae, infected plants were also more attractive to one of the BAW natural enemies. To our knowledge, this is the first time that the effect of pathogen-induced biochemical changes in plants on parasitoid behavior has been evaluated.

In choice tests, beet armyworm (BAW), Spodoptera exigua, larvae feed preferentially on leaves from peanut plants, Arachis hypogaea L., previously infected by the white mold fungus, Sclerotium rolfsii Saccodes (mitosporic fungi) (Cardoza et al. 2002). In this study we determined that third instar BAW caterpillars allowed to feed on S. rolfsii-infected plants had significantly higher survival, produced significantly heavier pupae, and had shorter time to pupation than those allowed to feed on healthy plants. Leaf tissue from white mold infected peanut plants contained similar levels of soluble and insoluble protein, but significantly higher levels of soluble sugars. In addition, white mold-infected plants had significantly lower starch content and total soluble phenolics compared with leaves from healthy plants. Levels of jasmonic acid were similar in plants attacked by either the fungus or BAW, but were significantly higher in plants that were infected by the fungus and then fed on by BAW. Salicyl...

Geothelphusa dehaani (White, 1874) is sympatric with G. exigua Suzuki and Tsuda, 1994, in Ohsumi Peninsula and with G. marmorata Suzuki and Okano, 2000, in Yakushima Island, southern Kyushu, Japan. Geothelphusa dehaani is widely distributed in Japan, whereas G. exigua and G. marmorata are endemic in riverine habitats at their respective locations. Visual and trapping surveys were conducted to examine the habitat preference and elucidate the factors maintaining sympatry among the three species. Results showed that G. exigua and G. marmorata are mainly aquatic, whereas G. dehaani is amphibious. Geothelphusa dehaani is more associated with land habitats under sympatry but equally extends in water and on land when singly constituting a population. All species exhibit varying degrees of nocturnalism, but G. marmorata is more active at night than either G. dehaani or G. exigua. Differences in habitats and activity patterns among the Geothelphusa species were attributed to their microenvironments and th...

The management of agroecosystems affects intricately linked assemblages of organisms, and nontarget species are not necessarily unimpacted. We examined the effect of Bt-cotton and of lepidopteran prey (Spodoptera exigua Hubner) that had ingested it on the adult survivorship of four important heteropteran predators of cotton pests. Longevity significantly decreased for Orius tristicolor White and Geocoris punctipes Say (by 28 and 27% of the control value, respectively), whereas no effect was found for Nabis sp. and Zelus renardii Kolenati. This finding contrasts with the results of previous studies in which Orius spp. and G. punctipes were either fed only plant material or nonlepidopteran prey. S. exigua is a lepidopteran with low susceptibility to the Bt toxin expressed in cotton and therefore exemplifies the possible effect on predators of lepidopteran pests that would become resistant to Bt. The importance of Bt toxin type, the difference between plants and prey and between different prey species as routes of ingestion of Bt toxins, and the need for studies assessing the population and ecosystem-level effects of Bt cotton are discussed.

We examined the effect of irrigating alfalfa (Medicago sativa L.) with selenium- contaminated water on the oviposition response, larval feeding preference, development and survival of the beet armyworm, Spodoptera exigua Hubner, a generalist herbivore. Alfalfa was grown in sand cultures under three levels of sodium selenate irrigation: (1) control with no Se added; (2) a low rate of 0.0066 g sodium selenate/60 liters water; (3) and a high rate of 0.20 g sodium selenate/60 liters water. The low concentration treatment resulted in 2.88 0.52 g Se/g plant dry weight and did not affect percent survival to adult eclosion compared with the control at 1.26 0.11 g Se/g dry weight. The high rate generated 305.81 52.14g Se/g dry weight of alfalfa and signiÞcantly fewer insects survived compared with insects fed control alfalfa at 1.11 0.12 g Se/g dry weight. High Se levels, but not low levels, decreased the relative growth index for larvae. In two-choice bioassays (treated/control) neonate larvae did not discriminate between control and Se-treated plants at high or low levels. Fourth instars did not discriminate between plants with low Se levels and control plants, but preferred to consume plants with high, usually lethal concentrations of Se. Females preferred ovipositing on plants with low Se concentrations over control plants, but did not discriminate between plants with high Se levels and untreated controls. This indicates that although females and late instars may be able to differentiate between Se-treated and control alfalfa they do not avoid plants containing high con- centrations of Se. Thus, alfalfa with high Se-treatment levels is resistant to S. exigua, and may serve as a population "sink," where females oviposit and few offspring survive to reproduce.

The toxicity of a number of emamectin benzoate homologues and photodegradates to five species of Lepidoptera was investigated using diet and foliar bioassays. The emamectin benzoate homologues B1a and B1b were equally toxic in the diet and foliar assays to Spodoptera exigua (Hubner), Heliothis virescens (F.), Tricoplusia ni (Hubner), and Spodoptera frugiperda (J. E. Smith), within each of these species. Plutella xylostella (L.) was the most sensitive species to emamectin benzoate. The AB1a photodegradate of emamectin benzoate was as toxic as the parent compound in the diet assay. However, in the foliage assay AB1a was 4.4-fold less toxic to S. exigua than the parent compound. The MFB1a photodegradate of emamectin benzoate was as toxic as the parent compound to P. xylostella, and 3.1 to 6.2 times as toxic as the parent compound to the other species in the diet assay. The order of toxicity of the photodegradates were AB1a > MFB1a > FAB1a > 8,9-Z-MAB1a > PAB1a.

M . L I U , Q . X . C A I , H . Z . L I U , B . H . Z H A N G , J . P . Y A N A N D Z . M . Y U A N . 2002. Aims: To investigate the distribution of chitinase in Bacillus thuringiensis strains, and the enhancing effects of the chitinase-producing B. thuringiensis strains on insecticidal toxicity of active B. thuringiensis strain against Spodoptera exigua larvae. Methods and Results: The chitinolytic activities of B.thuringiensis strains representing the 70 serotypes were investigated by the whitish opaque halo and the colorimetric method. Thirtyeight strains produced different levels of chitinase at pH 7AE0, and so did 17 strains at pH 10AE0. The strain T04A001 exhibited the highest production, reaching a specific activity of 355 U ml )1 in liquid medium. SDS-PAGE and Western blotting showed that the chitinase produced by some B. thuringiensis strains had a molecular weight of about 61 kDa. The bioassay results indicated that the chitinase-producing B. thuringiensis strains could enhance the insecticidal activity of B. thuringiensis strain DL5789 against S. exigua larvae, with an enhancing ratio of 2AE35-fold. Conclusion: This study demonstrated that chitinase was widely produced in B. thuringiensis strains and some of the strains could enhance the toxicity of active B. thuringiensis strain. Significance and Impact of the Study: This is the first investigation devoted exclusively to analyse the distribution of chitinase in B. thuringiensis. It infers that the chitinase produced by B. thuringiensis might play a role in the activity of the biopesticide.

Digestive enzymatic activity in three geographic strains (Miandiab, Kalposh and Moghan regions) of Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) reared on different sugar beet cultivars (Dorothea, Rozier, Persia and Perimer) was studied under laboratory conditions (25 ± 1 °C, 65 ± 5% RH, and a photo period of 16:8 (L:D) h photoperiod). The results of this study demonstrated that digestive protease and amylase activity of S. exigua larvae was affected by both geographic origin of the pest and host plant cultivar. Three strains reared on the same sugar beet cultivars demonstrated different levels of proteolytic and amylolytic activities in fourth and fifth instars. The highest proteolytic and amylolytic activity, in most cases, was observed in larvae collected from Kalposh region. Among different sugar beet cultivars, the highest protease activity in three strains was observed on cultivars Rozier and Perimer. Nevertheless, the highest amylase activity was seen on cultivar Dorothea, and the lowest activity was seen on cultivar Rozier. This study suggested that variations in digestive enzymatic activity of three geographic strains of S. exigua might be attributed to local adaptation with their local host plant and environmental conditions inherent by larvae.

The sesquiterpene carboxylic acids (SCA), (+)-(E)-α-santalen-12-oic, (−)-(E)-endo-α-bergamoten-12-oic, and (+)-(E)-endo-β-bergamoten-12-oic acid, are produced in glandular trichomes of Lycopersicon hirsutum f. typicum Humb. & Bonpl. accession (LA) 1777, which is highly resistant to a range of pests of cultivated tomatoes, L. esculentum Mill. Exposure of the larvae of two key tomato pests, tomato fruitworm [Helicoverpa zea (Boddie)] and beet armyworm [Spodoptera exigua (Hubner)], to these compounds in diets and on leaf surfaces resulted in reduced development rates and survival and deterred feeding. These effects were observed when levels of SCA, in artificial insect diet, applied to leaflets of susceptible cultivars, or synthesized in trichomes of leaves of plants, exceeded 2 mg SCA/g of diet or fresh leaf weight. This study suggests that cultivated tomatoes capable of synthesizing SCA, at 2 mg SCA/g of leaf tissue or greater, on their leaves and fruit would display enhanced host plant resistance...

Consumption rates, development times, and life table parameters of the beet armyworm, Spodoptera exigua (Hubner), were determined on five host plants: cabbage, Brassica oleracea capitata L; cotton, Gossypium hirsutum L.; bell pepper, Capsicum annuum L.; pigweed, Amaranthus retroflexus L.; and sunflower, Helianthus annuus L. Mean total leaf weight consumed by larvae was highest in cabbage (2.7 g) and lowest in pigweed (1.6 g). The feeding index (pupal weight divided by total weight of leaf tissue consumed) was highest on pigweed, followed by cotton, pepper, sunflower, and cabbage. On all host plants, significant relationships were found between amount of leaf tissue consumed and resulting pupal weight. Likewise, significant relationships were found between pupal weight and subsequent adult fecundity on all host plants. The highest percentage of female progeny was recorded in S. exigua reared on pigweed (62.2%) and the lowest for larvae reared on cabbage (43.6%). Duration of the larval stage was shortest on pigweed (12.4 d) and longest on pepper (18.0 d). Larval survival was highest on pigweed (94.4%) and lowest on cabbage (67.1%). Three key statistics were used to assess performance of S. exigua on the different host plants: 1) feeding index, 2) intrinsic rate of increase, r; and 3) growth index (percentage immature survival divided by immature development time). Using these measures, S. exigua performance was best on pigweed, worst on cabbage, and intermediate on cotton, pepper, and sunflower. We discuss the implications of these findings for control of S. exigua.

The beet armyworm, Spodoptera exigua (Hubner), is a sporadic but devastating secondary pest of cotton. Scouting cotton for egg masses is commonly recommended for identifying potential outbreaks and for proper timing of insecticide applications. However, there is disagreement regarding where to look on the plant for beet armyworm eggs. We investigated and quantified placement of egg masses by laboratory colony females within cotton and pigweed (a preferred wild host plant) canopies of different heights. In cotton, almost all egg masses were deposited on the undersides of leaves, and ≈80% of the egg masses were consistently located in the upper 50% of the cotton canopy, and horizontally within the inner 50% of the canopy around the central axis. Although this trend was consistent among all categories of plant height tested, variation about the means decreased with increasing height. A smaller sample of wild females indicated similar vertical placement, but horizontal placement extending further distally in the canopy. Our results indicate that scouting for egg masses on the underside of leaves in the upper half of the canopy will recover ≈80% of the egg masses present on the plants. In pigweed, egg masses were commonly laid within the inner 50% of the canopy, but along the upper 80% of the vertical axis. As in cotton, variation about the means was less in taller plants. The number of eggs per egg mass was 29% less in cotton than in pigweed. This positional information will aid in further efforts to investigate, predict, and manage beet armyworm populations in cotton and noncotton hosts.