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Insect herbivores face multiple challenges to their ability to grow and reproduce. Plants can produce a series of defenses that disrupt and damage the herbivore digestive system, which are heightened upon injury by insect feeding. Additionally, insects face threats from virulent microorganisms that can incur their own set of potential costs to hosts. Microorganisms that invade through the digestive system may function in concert with defenses generated by plants, creating combined assailments on host insects. In our study, we evaluated how tomato defenses interact with an enteric bacterial isolate, Serratia marcescens, in the corn earworm (Helicoverpa zea). We performed bioassays using different tomato cultivars that were induced by methyl jasmonate and larvae orally inoculated with a S. marcescens isolate. Untreated corn earworm larval mortality was low on constitutive tomato, while larvae inoculated with S. marcescens exhibited > 50% mortality within 5 days. Induction treatments elevated both control mortality (~ 45%) and in combination with S. marcescens (> 95%). Larvae also died faster when encountering induced defenses and Serratia. Using a tomato mutant, foliar polyphenol oxidase activity likely had stronger impacts on S. marcescens-mediated larval mortality. Induction treatments also elevated the number of bacterial colony-forming units in the hemolymph of larvae inoculated with Serratia. Larval mortality by S. marcescens was low (< 10%) on artificial diets. Our results demonstrate that plant chemical defenses enhance larval mortality from an opportunistic gut microbe. We propose that the combined damage from both the plant and microbial agent overwhelm the herbivore to increase mortality rates and expedite host death. [ABSTRACT FROM AUTHOR]

Insects possess specific immune responses to protect themselves from different types of pathogens. Activation of immune cascades can inflict significant developmental costs on the surviving host. To characterize infection kinetics in a surviving host that experiences baculovirus inoculation, it is crucial to determine the timing of immune responses. Here, we investigated time‐dependent immune responses and developmental costs elicited by inoculations from each of two wild‐type baculoviruses, Autographa californica multiple nucleopolyhedrovirus (AcMNPV) and Helicoverpa zea single nucleopolyhedrovirus (HzSNPV), in their common host H. zea. As H. zea is a semi‐permissive host of AcMNPV and fully permissive to HzSNPV, we hypothesized there are differential immune responses and fitness costs associated with resisting infection by each virus species. Newly molted 4th‐instar larvae that were inoculated with a low dose (LD15) of either virus showed significantly higher hemolymph FAD‐glucose dehydrogenase (GLD) activities compared to the corresponding control larvae. Hemolymph phenoloxidase (PO) activity, protein concentration and total hemocyte numbers were not increased, but instead were lower than in control larvae at some time points post‐inoculation. Larvae that survived either virus inoculation exhibited reduced pupal weight; survivors inoculated with AcMNPV grew slower than the control larvae, while survivors of HzSNPV pupated earlier than control larvae. Our results highlight the complexity of immune responses and fitness costs associated with combating different baculoviruses. [ABSTRACT FROM AUTHOR]

The effect of temperature on insect-plant interactions in the face of changing climate is complex as the plant, its herbivores and their interactions are usually affected differentially leading to an asymmetry in response. Using experimental warming and a combination of biochemical and herbivory bioassays, the effects of elevated temperatures and herbivore damage (Helicoverpa zea) on resistance and tolerance traits of Solanum lycopersicum var. Better boy (tomato), as well as herbivory performance and salivary defense elicitors were examined. Insects and plants were differentially sensitive towards warming within the experimental temperature range. Herbivore growth rate increased with temperature, whereas plants growth as well as the ability to tolerate stress measured by photosynthesis recovery and regrowth ability were compromised at the highest temperature regime. In particular, temperature influenced the caterpillars' capacity to induce plant defenses due to changes in the amount of a salivary defense elicitor, glucose oxidase (GOX). This was further complexed by the temperature effects on plant inducibility, which was significantly enhanced at an above-optimum temperature; this paralleled with an increased plants resistance to herbivory but significantly varied between previously damaged and undamaged leaves. Elevated temperatures produced asymmetry in species' responses and changes in the relationship among species, indicating a more complicated response under a climate change scenario. [ABSTRACT FROM AUTHOR]

Animals have ubiquitous associations with microorganisms, but microbial community composition and population dynamics can vary depending upon many environmental factors, including diet. The bacterial communities present in caterpillar (Lepidoptera) guts are highly variable, even among individuals of a species. Across lepidopteran species, it is unclear if the variation in their gut bacterial communities is due to ingested bacteria with diets or responses of gut bacteria to their diet. In this study, we aimed to understand whether bacteria establish and persist in the lepidopteran gut or just pass through the gut with food. We also examined whether bacterial establishment in lepidopteran guts depended on diet. We conducted a series of experiments using axenic and gnotobiotic insect rearing methods to address these objectives. We found that bacteria were established and maintained without replacement through the larval instars of the fall armyworm (Spodoptera frugiperda) and corn earworm (Helicoverpa zea). Gut bacterial titers increased when larvae were fed gamma-irradiated corn leaves but decreased when fed a wheat germ artificial diet. However, bacterial titers of larvae fed on a pinto bean artificial diet were similar to those consuming intact plants. We also observed that microbial titers of fall armyworm and other folivorous larvae were positively related to the host body size throughout larval development. Collectively, these results suggest that the populations of bacteria present in caterpillar guts are not simply a transient community passing through the system, but rather are a dynamic component of the caterpillar gut. Sensitivity of bacterial populations to the type of diet fed to lepidopterans suggests that not all diets are equally useful for reducing variance in community structure and interpreting insect-microbe interactions. [ABSTRACT FROM AUTHOR]

Microplitis croceipes is a solitary parasitoid that specializes on noctuid larvae of Helicoverpa zea and Heliothis virescens. Both the parasitoid and its hosts are naturally distributed across a large part of North America. When parasitoids deposit their eggs into hosts, venom and polydnaviruses (PDVs) are also injected into the caterpillars, which can suppress host immune responses, thus allowing parasitoid larvae to develop. In addition, PDVs can regulate host oral cues, such as glucose oxidase (GOX). The purpose of this study was to determine if parasitized caterpillars differentially induce plant defenses compared to non-parasitized caterpillars using two different caterpillar host/plant systems. Heliothis virescens caterpillars parasitized by M. croceipes had significantly lower salivary GOX activity than non-parasitized caterpillars, resulting in lower levels of tomato defense responses, which benefited parasitoid performance by increasing the growth rate of parasitized caterpillars. In tobacco plants, parasitized Helicoverpa zea caterpillars had lower GOX activity but induced higher plant defense responses. The higher tobacco defense responses negatively affected parasitoid performance by reducing the growth rate of parasitized caterpillars, causing longer developmental periods, and reduced cocoon mass and survival of parasitoids. These studies demonstrate a species-specific effect in different plant-insect systems. Based on these results, plant perception of insect herbivores can be affected by parasitoids and lead to positive or negative consequences to higher trophic levels depending upon the particular host-plant system. [ABSTRACT FROM AUTHOR]

Insect-associated microbes can contribute to the physiological and ecological functions of insects. Despite a few examples in beetles and piercing-sucking insects, the varied mechanisms of how insect-associated bacteria mediate plant-insect interactions are still not fully understood. The polyphagous herbivore Helicoverpa zea is a major agricultural pest that harbors certain microbes in their digestive systems. Enterobacter ludwigii is one of the gut-associated bacteria identified from field-collected caterpillars, and it has been shown to indirectly induce defenses in the dicot plant tomato by triggering the biosynthesis of salivary elicitors, but there are no clear mechanisms to show how gut microbes alter these salivary cues and how a different host plant responds to these inducible elicitors. Here, we conducted a series of assays to determine whether infection with E. ludwigii affects H. zea larval growth, immunity, and salivary responses and thus influences induced defenses of maize to herbivory. Inoculating lab-reared caterpillars with E. ludwigii, did not significantly affect the growth of caterpillars, but two immunity-related genes glucose oxidase (GOX) and lysozyme (LYZ) were more highly expressed in both salivary glands and midguts compared with MgCl2 solution-treated caterpillars. Oral elicitors were evaluated for their role in triggering maize-specific defense responses. Our results show that saliva and its main component protein glucose oxidase (GOX) from E. ludwigii-inoculated caterpillars played a role in inducing maize anti-herbivore responses. These findings provide a novel concept that introducing bacteria to an herbivore may be an important approach to pest control through alteration of insect immune responses and thus indirect induction of plant resistance. [ABSTRACT FROM AUTHOR]

Insect gut-associated microbes modulating plant defenses have been observed in beetles and piercing-sucking insects, but the role of caterpillar-associated bacteria in regulating plant induced defenses has not been adequately examined., We identified bacteria from the regurgitant of field-collected Helicoverpa zea larvae using 16S ribosomal RNA (rRNA) gene sequencing and matrix-assisted laser desorption/ionization time of flight (MALDI-TOF) mass spectrometry. A combination of biochemical, molecular, and confocal electron microscopy methods were used to determine the role of caterpillar-associated bacteria in mediating defenses in Solanum lycopersicum (tomato)., Laboratory-reared H. zea inoculated with one of the bacteria identified in field-collected H. zea, Enterobacter ludwigii, induced expression of the tomato defense-related enzyme polyphenol oxidase and genes regulated by jasmonic acid (JA), whereas the salicylic acid (SA)-responsive pathogenesis-related gene was suppressed. Additionally, saliva and its main component glucose oxidase from inoculated caterpillars played an important role in elevating tomato anti-herbivore defenses. However, there were only low detectable amounts of regurgitant or bacteria on H. zea-damaged tomato leaves., Our results suggest that H. zea gut-associated bacteria indirectly mediate plant-insect interactions by triggering salivary elicitors. These findings provide a proof of concept that introducing gut bacteria to a herbivore may provide a novel approach to pest management through indirect induction of plant resistance. [ABSTRACT FROM AUTHOR]

Naturally occurring insect viruses can modify the behaviour of infected insects and thereby modulate virus transmission. Modifications of the virus genome could alter these behavioural effects. We studied the distance moved and the position of virus-killed cadavers of fourth instars of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) infected with a wild-type genotype of H. armigera nucleopolyhedrovirus (HaSNPV) or with one of two recombinant genotypes of this virus on cotton plants. The behavioural effects of virus infection were examined both in larvae infected with a single virus genotype, and in larvae challenged with mixtures of the wild-type and one of the recombinant viruses. An egt-negative virus variant caused more rapid death and lower virus yield in fourth instars, but egt-deletion did not produce consistent behavioural effects over three experiments, two under controlled glasshouse conditions and one in field cages. A recombinant virus containing the AaIT-( Androctonus australis Hector) insect-selective toxin gene, which expresses a neurotoxin derived from a scorpion, caused faster death and cadavers were found lower down the plant than insects infected with unmodified virus. Larvae that died from mixed infections of the AaIT-expressing recombinant and the wild-type virus died at positions significantly lower, compared to infection with the pure wild-type viral strain. The results indicate that transmission of egt-negative variants of HaSNPV are likely to be affected by lower virus yield, but not by behavioural effects of egt gene deletion. By contrast, the AaIT recombinant will produce lower virus yields as well as modified behaviour, which together can contribute to reduced virus transmission under field conditions. In addition, larvae infected with both the wild-type virus and the toxin recombinant behaved as larvae infected with the toxin recombinant only, which might be a positive factor for the risk assessment of such toxin recombinants in the environment. [ABSTRACT FROM AUTHOR]

The original version of this article unfortunately contained a mistake. Fig. 3 and Fig. 4a were identical and the original version of Fig. 4a had been accidentally replaced. [ABSTRACT FROM AUTHOR]

[Display omitted] • Serratia -mediated lepidopteran larval mortality increases when hosts consume low-quality food. • Fall armyworm exhibited little mortality across diets, but corn earworm and beet armyworm had high levels of mortality on diluted diet. • Larval mortality were not driven by dilution in diet antimicrobials. Insect guts often harbor an abundance of bacteria. Many of these members are commensal, but some may emerge as opportunistic pathogens when the host is under stress. In this study, we evaluated how dietary nutritional concentration mediates a shift from commensal to pathogenic, and if host species influences those interactions. We used the lepidopterans (Noctuidae) fall armyworm (Spodoptera frugiperda), beet armyworm (Spodoptera exigua), and corn earworm (Helicoverpa zea) as hosts and a Serratia strain initially isolated from healthy fall armyworm. Diet concentration was altered by bulk reduction in nutritional content with dilution using cellulose. Our experiments revealed that low nutrient diet increased mortality from Serratia for beet armyworm and corn earworm. However, for fall armyworm, little mortality was observed in any of the diet combinations. Dietary nutrition and oral inoculation with Serratia did not change the expression of two antimicrobial peptides in fall and beet armyworm, suggesting that other mechanisms that mediate mortality were involved. Our results have implications for how pathogens may persist as commensals in the digestive tract of insects. These findings also suggest that diet plays a very important role in the switch from commensal to pathogen. Finally, our data indicate that the host response to changing conditions is critical in determining if a pathogen may overtake its host and that these three lepidopteran species have different responses to opportunistic enteric pathogens. [ABSTRACT FROM AUTHOR]