Polyphagous Caterpillar Research Articles

High intraspecific variability and previous experience affect polyphenol metabolism in polyphagous Lymantria mathura caterpillars.

Polyphagous insect herbivores feed on multiple host-plant species and face a highly variable chemical landscape. Comparative studies of polyphagous herbivore metabolism across a range of plants is an ideal approach for exploring how intra- and interspecific chemical variation shapes species interactions. We used polyphagous caterpillars of Lymantria mathura (Erebidae, Lepidoptera) to explore mechanisms that may contribute to its ability to feed on various hosts. We focused on intraspecific variation in polyphenol metabolism, the fates of individual polyphenols, and the role of previous feeding experience on polyphenol metabolism and leaf consumption. We collected the caterpillars from Acer amoenum (Sapindaceae), Carpinus cordata (Betulaceae), and Quercus crispula (Fagaceae). We first fed the larvae with the leaves of their original host and characterized the polyphenol profiles in leaves and frass. We then transferred a subset of larvae to a different host species and quantified how host shifting affected their leaf consumption and polyphenol metabolism. There was high intraspecific variation in frass composition, even among caterpillars fed with one host. While polyphenols had various fates when ingested by the caterpillars, most of them were passively excreted. When we transferred the caterpillars to a new host, their previous experience influenced how they metabolized polyphenols. The one-host larvae metabolized a larger quantity of ingested polyphenols than two-host caterpillars. Some of these metabolites could have been sequestered, others were probably activated in the gut. One-host caterpillars retained more of the ingested leaf biomass than transferred caterpillars. The pronounced intraspecific variation in polyphenol metabolism, an ability to excrete ingested metabolites and potential dietary habituation are factors that may contribute to the ability of L. mathura to feed across multiple hosts. Further comparative studies can help identify if these mechanisms are related to differential host-choice and response to host-plant traits in specialist and generalist insect herbivores.

Nutrition of host plants influence the infectivity of nucleopolyhedrovirus to polyphagous caterpillar, Hyphantria cunea

Plants play an important role in interactions between insect herbivores and their pathogens. The ability of host plants to modify the infectivity of entomopathogens in herbivorous insects has been widely documented. However, the plants’ nutrients have always been neglected as a factor contributing to variation in the susceptibility of insect herbivores to entomopathogens. The fall-webworm (FWW), Hyphantria cunea Drury, is a typical polyphagous caterpillar, and the Hyphantria cunea nucleopolyhedrovirus (HycuNPV) is a distinctly specialized baculovirus for the FWW, which is safe for other organisms and has been effectively used as a biological insecticide against H. cunea in China. In this study, we investigated the nutrient components of four host plant species, i.e., Prunus serrulate, Cerasus serrulate, Camptotheca acuminata, and Populus deltoides, and their effects on the susceptibility of H. cunea larvae to HycuNPV. The HycuNPV-infected larvae fed on P. deltoides leaves exhibited higher survival rates, longer survival times, more food intake, and gained larger body size. These biological parameters were positively correlated with the nitrogen components of host plant leaves. Moreover, the larval antioxidant enzymes exhibited different responses to HycuNPV. HycuNPV infection significantly triggered the catalase (CAT) and prophenoloxidase (PPO) enzyme activity levels of H. cunea larvae. The uninfected larvae fed on poplar leaves induced a robust increase in the POD activity, which could scavenge extra reactive oxygen species and provide a protective effect against the HycuNPV. In conclusion, the plant-mediated effects of HycuNPV on the FWW have been investigated in this study. The nitrogen content in dietary was an essential factor in determining the insect herbivore susceptibility to entomopathogenic viruses, and it helped explain variations in the susceptibility of pests to the entomopathogenic viruses and aid in developing more robust tolerance monitoring assays in the lab that reflect the performance of pests in the field.Graphical

A chromosome-level genome assembly of the beet armyworm Spodoptera exigua

BackgroundThe beet armyworm Spodoptera exigua is a polyphagous caterpillar that causes serious damage to many species of crops and vegetables. To gain insight into how this polyphagous insect differs from less harmful oligophagous species, we generated a chromosome-level assembly and compared it to closely related species with the same or different feeding habits. ResultsBased on Illumina and Pacific Biosciences data and Hi-C technology, 425.6 Mb of genome sequences were anchored and oriented into 31 linkage groups, with an N50 length of 14.8 Mb. A total of 24,649 gene models were predicted, of which 97.4% were identified in the genome assembly. Chemosensory genes are vital for locating food: of the four main families, odorant-binding proteins, chemosensory proteins and olfactory receptors showed little difference, whereas gustatory receptors are greatly expanded in S. exigua. Examination of other polyphagous insects confirmed this difference from oligophagous congeners and further identified the bitter receptor subfamily as being particularly affected. ConclusionOur high-quality genome sequence for beet armyworm identified a key expansion of the bitter gustatory receptor subfamily in this and other pests that differs crucially from more benign relatives and offers insight into the biology and possible future means of control for these economically important insects.

Fungal consortium of two Beauveria bassiana strains increases their virulence, growth, and resistance to stress: A metabolomic approach.

The use of two or more microorganisms in a microbial consortium has been increasingly applied in the biological control of diseases and pests. Beauveria bassiana is one of the most widely studied fungal species in biological control, yet little is known about its role in fungal consortiums. In a previous study, our group found that a consortium formed by two strains of B. bassiana had significantly greater biocontrol potential against the polyphagous caterpillars Duponchelia fovealis (Lepidoptera: Crambidae) than either strain on its own. In this study, we use GC-MS and LC-MS/MS to evaluate and discuss the metabolomics of the consortium. A total of 21 consortium biomarkers were identified, corresponding to 14 detected by LC-MS/MS and seven by GC-MS. Antioxidant and anti-inflammatory mechanisms are the main properties of the metabolites produced by the consortium. These metabolites can depress the insect’s immune system, increasing its vulnerability and, hence, the fungal virulence of the consortium. In light of these results, we propose an action model of insect mortality due to the metabolites secreted by the consortium. The model includes the inhibition of defense mechanisms such as pro-inflammatory interleukin secretion, cell migration, cell aggregation, Dif, Dorsal and Relish gene transcription, and JAK/STAT and JNK signaling pathways. It also promotes the cleaning of oxidative molecules, like ROS, NOS, and H2O2, and the induction of virulence factors.

Prevalence and Impact of the Polyphagous Caterpillar of Cabbage White Butterfly (Pieris brassicae Linnaeus, 1758) On The Marrow Stem Kale (Brassica oleracea L. Var Virdis)

Kale is one of the important and chiefly consumed vegetable crops of Kashmir valley. The pest infestation on the Brassica oleracea L. var virdis (Brassicales: Brassicaceae) by the polyphagous caterpillar of Pieris brassicae Linnaeus, 1758 (Lepidoptera: Pieridae) is presented in this study. The monitoring of pest infestation was assessed in the months from July-August 2021 from four different locations in the district Shopian, Kashmir, India. The present study thoroughly monitored the life stages of the polyphagous caterpillar of Pieris brassicae and its impact on the host plant. Among the larval stages of the pest, most damages was observed on the leaves by 2nd and 3rd instar larvae followed by the 4th and 5th instars respectively in non-insecticide treatment; with a higher rate of leaf damage (3.15 %) and larval incidence (1.13 %), while a lower rate of leaf damage (0.2 %) and larval incidence (0.04 %) was observed in insecticide control.

CYP4G8 is responsible for the synthesis of methyl-branched hydrocarbons in the polyphagous caterpillar of Helicoverpa armigera

Insect cuticular hydrocarbons (CHCs) have dual functions as physical barrier and chemical signals. The last step of CHC biosynthesis is known to be catalyzed by cytochrome P450 CYP4G in a number of insects. Until recently, studies on CYP4Gs in the context of functional evolution are rare. In this study, we analyzed sequence similarity and temporal-spatial expression patterns of the five CYP4G genes in the cotton bollworm Helicoverpa armigera, an important agricultural pest and also typical representative of lepidopteran insects. Moreover, the CRISPR/Cas9-induced knockout was used to clarify the roles of the five CYP4Gs in CHC biosynthesis. Temporal-spatial expression patterns revealed that CYP4G8 was highly expressed at all developmental stages and in most tissues examined. Larvae with CYP4G8 knocked out could not produce methyl-branched CHCs and failed to pupate, while larvae with the other four CYP4G genes knocked out (4G1-type-KO) showed no significant changes in their CHC profiles, weight gain and survival. Comparative transcriptomics revealed that knocking out CYP4G8 affected the global gene expression in larvae, especially down-regulated the expression of genes in the fatty acid biosynthetic pathway, while no significant change in 4G1-type-KO transcriptome was observed. These findings indicate that the five members of the CYP4G subfamily have undergone functional divergence: CYP4G8 maintains the essential function in CHC biosynthesis, while the function of the other four CYP4G genes remains unclear. Intriguingly, CYP4G8 has evolved to be a P450 enzyme responsible for the synthesis of larval methyl-branched hydrocarbons. The observation that CYP4G8 knockout is lethal strongly suggest that CYP4G8 may serve as a candidate target for the development of insecticidal agents for the control of cotton bollworms.

Polyphagous caterpillars of Spodoptera litura switch from a trap crop to the main crop, improve fitness, and shorten generation time

Trap crops are used for pulling the pest load from the main crops toward themselves. Here, we report a switching by a polyphagous pest, Spodoptera litura, from trap crop castor to main crop eggplant during its mid-larval development. In the eggplant–castor mixed field, adults oviposit exclusively on castor. However, 3rd and 4th instar castor-grown larvae migrate to eggplants. Host choice assays for adults and larvae conducted in the controlled environment and the nutritional indices of castor and eggplant-feeding larvae revealed that the natural enemies did not influence the host switch. The host switch was rather associated with larvae’s enhanced digestion and metabolism. It shortened the development time and increased the eclosion success on eggplant. Mother-selected castor is suitable for the early instars and eggplant is for the later instars, indicating that the host switching was not because of the mother’s erroneous host choice but has evolved as an advantageous behavior. S. litura’s ability to switch hosts in the same ontogenic phase and reduce the generation time can contribute to an additional generation within a cropping season and endanger the critical growth stage of fruiting. Our findings support the ‘mother knows the best’ concept and annex that the larvae can also possess the ability to know the best. Such ability in different ontogenic stages can be fundamental to the exceptional adaptability of species like S. litura.

Synergism of the Bacillus thuringiensis Cry1, Cry2, and Vip3 Proteins in Spodoptera frugiperda Control.

The polyphagous caterpillar, Spodoptera frugiperda, has been controlled with either chemical insecticides or transgenic plants such as Bt maize that expresses the cry and/or vip genes of the Bacillus thuringiensis (Bt) bacterium. Despite the efficiency of Bt toxins in lepidopteran control, populations resistant to Bt plants have emerged in different locations around the world. Thus, understanding how combined proteins interact against pests can assist resistance control and management. This work demonstrated the toxicity of Cry1Ab, Cry1Ac, Cry1Ca, Cry1Ea, Cry2Aa, Cry2Ab, Vip3Aa, and Vip3Ca in single and combined assays against S. frugiperda neonatal larvae. All protein mixtures had synergistic action in the control of the larvae. The Vip3Aa + Cry1Ab mixture had the highest toxicity, sequentially followed by Vip3Aa + Cry2Ab, Cry1Ab + Cry2Ab + Vip3Aa, Cry1Ea + Cry1Ca, Cry1Ab + Cry2Ab, Vip3Ca + Cry1Ea, and Vip3Ca + Cry1Ca. Cry1Ab, Cry1Ac, Cry2Ab, and Vip3Aa bound to more than one site on the brush border membrane vesicles (BBMV) of S. frugiperda. The Cry1Ab and Cry1Ac proteins share binding site, while Cry1Ab does not share binding site with the Cry2Aa and Cry2Ab proteins. The Vip3Aa protein does not share receptors with the tested Cry1 and Cry2. The results suggest that combination these tested proteins may increase toxicity against S. frugiperda neonates.

Herbivore-Induced Defenses in Tomato Plants Enhance the Lethality of the Entomopathogenic Bacterium, Bacillus thuringiensis var. kurstaki.

Plants can influence the effectiveness of microbial insecticides through numerous mechanisms. One of these mechanisms is the oxidation of plant phenolics by plant enzymes, such as polyphenol oxidases (PPO) and peroxidases (POD). These reactions generate a variety of products and intermediates that play important roles in resistance against herbivores. Oxidation of the catecholic phenolic compound chlorogenic acid by PPO enhances the lethality of the insect-killing bacterial pathogen, Bacillus thuringiensis var. kurstaki (Bt) to the polyphagous caterpillar, Helicoverpa zea. Since herbivore feeding damage often triggers the induction of higher activities of oxidative enzymes in plant tissues, here we hypothesized that the induction of plant defenses would enhance the lethality of Bt on those plants. We found that the lethality of a commercial formulation of Bt (Dipel® PRO DF) on tomato plants was higher if it was applied to plants that were induced by H. zea feeding or induced by the phytohormone jasmonic acid. Higher proportions of H. zea larvae killed by Bt were strongly correlated with higher levels of PPO activity in the leaflet tissue. Higher POD activity was only weakly associated with higher levels of Bt-induced mortality. While plant-mediated variation in entomopathogen lethality is well known, our findings demonstrate that plants can induce defensive responses that work in concert with a microbial insecticide/entomopathogen to protect against insect herbivores.

Immediate and long-term facilitative effects of cattle grazing on a polyphagous caterpillar

Mammalian herbivores induce changes in the composition, abundance, architecture and chemistry of vegetation which can affect insects in their habitat. Many studies addressed the long-term effects of mammalian grazing on insect herbivores, yet few examined the effects during grazing (or right after it takes place). We investigated the immediate and long-term effects of cattle grazing on the abundance and distribution of the herbivorous spring webworm caterpillar (Ocnogyna loewii), via excluding cattle (by fencing) within a grazed paddock. In addition, we estimated the caterpillar density in replicated grazed and non-grazed paddocks (maintained as so for dozens of years), in moderate and heavy grazing intensities. Since the caterpillars develop during the cold winter months, we predicted that cattle grazing would positively affect them by reducing plant height and increasing their exposure to direct warm sunlight. Therefore, we examined caterpillar preference for sun-exposed areas using shade-manipulation experiments. Overall, cattle grazing positively affected the caterpillars, increasing their numbers two-fold on average, regardless of grazing intensity. This effect was immediate, as the caterpillars rapidly responded to exclusion of cattle by moving away from non-grazed areas. Caterpillar growth rate was similar when feeding on grazed and non-grazed vegetation. Most caterpillars (over 80%) preferred sun over manipulated shaded microhabitats. Furthermore, we found that cattle usually do not ingest caterpillars while feeding. Cattle grazing likely benefited the caterpillars that develop under low temperatures by reducing plant cover, thus creating a warmer habitat. This study demonstrates how changes in vegetation structure caused by mammalian herbivores can rapidly and positively affect the abundance and distribution of herbivorous insects.

Polyphagy by omnivory: scavenging improves performance of a polyphagous caterpillar on marginal hosts.

Few species of insect herbivores are highly polyphagous, but those few species are disproportionately ecologically and economically important and include many of the most destructive crop pests. Common correlates of extreme polyphagy across insects include the related behaviors of cannibalism and omnivory, though any functional consequences of these behaviors on the host range are unknown. I hypothesized that omnivory may allow these insects to exploit marginal hosts successfully (an expansion of realized niche). Using the polyphagous pest caterpillar, Heliothis virescens, I tested the polyphagy by omnivory hypothesis using ten host plants of varying suitability and small quantities of insect carrion. Caterpillars which were allowed omnivory had increased performance on lower-quality hosts; this treatment raised survival, growth rate, and pupal mass over controls on a strictly plant diet. Omnivory allowed successful development on two plants that caterpillars could not exploit alone a potential niche expansion. This effect was limited, however: (1) on high-quality hosts, omnivory did not improve performance, and (2) omnivory on poor hosts did not increase growth rate or pupal mass to levels matching the most suitable hosts and it could not permit exploitation of a completely unpalatable plant. Omnivory may therefore be an important (and overlooked) factor in determining the success of generalist insect herbivores in a variety of ecological settings.

Bottom-Up Mechanisms Generate the Same Temporal Pattern of Attack by a Specialist and a Generalist Caterpillar on Short-Lived Plants.

The local population dynamics of insect herbivores in ephemeral patches of short-lived plants are poorly known. We investigated whether a specialist and a generalist caterpillar exhibit contrasting temporal patterns of attack during plant development and also assessed bottom-up forces related to plant ontogeny that govern such population trends. Immature stages of the polyphagous Trichoplusia ni (Hübner) and the oligophagous Plutella xylostella (L.) were sampled throughout the development of cabbage (Brassica oleracea L. var. capitata L.) crops. We measured protein and glucosinolate contents and insect performance with regard to plant age and leaf strata. The populations of both caterpillar species changed in close parallel throughout plant development, and a nonlinear temporal pattern of egg laying was reproduced in sequential population patterns of the larval stages until pupation. Reduced protein availability and insect performance coincided with a decline in egg laying and subsequent larval abundance in mature plants. By standardizing the plant size, we found that young and nutritious plants support proportionately more insects than large and mature plants. In our models of the population oscillations, the interaction between plant size and quality provided a strong causal explanation for the densities of both oligophagous and polyphagous caterpillars. Patches of fast-growing herbaceous plants are very common worldwide in the form of crop fields, and a generalized temporal pattern of attack may be widespread among caterpillars, regardless of their feeding specialization. Our results highlight the role of bottom-up forces in shaping the population dynamics of caterpillars in such systems.

Reduced consumption of protein-rich foods follows immune challenge in a polyphagous caterpillar.

Advances in ecological immunity have illustrated that, like vertebrates, insects exhibit adaptive immunity, including induced changes in feeding behavior that aid the immune system. In particular, recent studies have pointed to the importance of protein intake in mounting an immune response. In this study, we tested the hypothesis that the polyphagous caterpillar Grammia incorrupta (H. Edwards) (Family: Erebidae) would adaptively change its feeding behavior in response to immune challenge, predicting that caterpillars would increase their intake of dietary protein. We further predicted that this response would enhance the melanization response, a component of the immune system that acts against parasitoids. We challenged the immune system using either tachinid fly parasitoids or a bead injection technique that has been used in studies to simulate parasitism, and measured feeding before and after immune challenge on diets varying in their macronutrient content. To evaluate the effects of diet on melanization, we quantified melanization of beads following feeding assays. Contrary to our prediction, we found that parasitized or injected caterpillars given a choice between high- and low-protein foods reduced their intake of the high-protein food. Furthermore, in a no-choice experiment, caterpillars offered food with a protein concentration that is optimal for growth reduced feeding following immune challenge, whereas those offered a low-protein food did not. Although variation in protein intake did not change the caterpillars' melanization response, increased carbohydrate intake did increase melanization, suggesting a prophylactic role for carbohydrates. We discuss alternative mechanisms by which variation in protein intake could negatively or positively affect parasitized caterpillars, including nutritional interactions with the caterpillar's self-medication response.

Can caterpillar density or host-plant quality explain host-plant-related parasitism of a generalist forest caterpillar assemblage?

Herbivore-carnivore interactions are influenced by the plants on which herbivores feed. Accordingly, dietary generalist herbivores have been shown to experience differential risk of mortality from carnivores on different host-plant species. Here, we investigate whether caterpillar density and host-plant quality play a role in driving variation in generalist forest caterpillar mortality from insect parasitoids using a large-scale, multi-year observational study. A total of 4,500 polyphagous caterpillars were collected from eight host-tree species in Connecticut deciduous forests over 5 years, and frequencies of mortality from insect parasitoids (flies and wasps) were compared across the eight host-plant species for the entire generalist caterpillar assemblage (76 species). Separate comparisons were made using seven numerically dominant generalist species, allowing us to account for variation in caterpillar species-specific parasitism risk. We find significant variation in parasitism frequencies of generalist caterpillars across the eight host-plant species when accounting for variation in caterpillar density. We find no support for an influence of caterpillar density on parasitism and no clear evidence for an effect of host-plant quality on parasitism. Therefore, the results of this study discount the hypotheses that variation in caterpillar density and host-plant quality are responsible for variation in parasitism frequencies across host-plant species. Instead, our findings point to other plant-related characteristics, such as plant-derived parasitoid attractants, which may have robust, community-wide effects.

The interplay between toxin-releasing β-glucosidase and plant iridoid glycosides impairs larval development in a generalist caterpillar, Grammia incorrupta (Arctiidae)

Herbivores with polyphagous feeding habits must cope with a diet that varies in quality. One of the most important sources of this variation in host plant suitability is plant secondary chemistry. We examined how feeding on plants containing one such group of compounds, the iridoid glycosides, might affect the growth and enzymatic activity in a polyphagous caterpillar that feeds on over 80 plant species in 50 different families. Larvae of the polyphagous arctiid, Grammia incorrupta, were reared exclusively on one of two plant species, one of which contains iridoid glycosides (Plantago lanceolata, Plantaginaceae) while the other does not (Taraxacum officinale, Asteraceae). Larval weight was measured on the two host plants, and midgut homogenates of last instar larvae were then assayed for activity and kinetic properties of β-glucosidases, using both a standard substrate, 4-nitrophenyl-β-d-glucose (NPβGlc), and the iridoid glycoside aucubin, one of the two main iridoid glycosides in P. lanceolata. Larvae feeding on P. lanceolata weighed significantly less and developed more slowly compared to larvae on T. officinale. While the larval midgut β-glucosidase activity determined with NPβGlc was significantly decreased when fed on P. lanceolata, aucubin was substantially hydrolyzed and the larval β-glucosidase activity towards both substrates correlated negatively with larval weight. Our results demonstrate that host plants containing high concentrations of iridoid glycosides have a negative impact on larval development of this generalist insect herbivore. This is most likely due to the hydrolysis of plant glycosides in the larval midgut which results in the release of toxic aglycones. Linking the reduced larval weight to the toxin-releasing action of an iridoid glycoside cleaving β-glucosidase, our results thus support the detoxification limitation hypothesis, suggesting fitness costs for the larvae feeding solely on P. lanceolata. Thus, in addition to the adaptive regulation of midgut β-glucosidase activity, host plant switching as a behavioral adaptation might be a prerequisite for generalist herbivores that allows them to circumvent the negative effects of plant secondary compounds.

Dropping from Host Plants in Response to Predators by a Polyphagous Caterpillar

Dropping from Host Plants in Response to Predators by a Polyphagous Caterpillar

Cascading effects of host size and host plant species on parasitoid resource allocation

1. The bottom-up factors that determine parasitoid host use are an important area of research in insect ecology. Host size is likely to be a primary cue for foraging parasitoids due to its potential influence on offspring development time, the risk of multiparasitism, and host immunocompetence. Host size is mediated in part by host-plant traits that influence herbivore growth and potentially affect a herbivore's quality as a host for parasitoids. 2. Here, we tested how caterpillar host size and host plant species influence adult fly parasitoid size and whether host size influences wasp parasitoid sex allocation. We measured the hind tibia lengths and determined the sex of wasp and fly parasitoids reared from 11 common host species of polyphagous caterpillars (Limacodidae) that were in turn reared on foliage of seven different host plant species. 3. We also tested how host caterpillar species, host caterpillar size, and host and parasitoid phenology affect how the parasitoid community partitions host resources. We found evidence that parasitoids primarily partition their shared hosts based on size, but not by host species or phenology. One index of specialisation (d′) supports our observation that these parasitoids are quite generalised within the Limacodidae. In general, wasps were reared from caterpillars collected in early instars, while flies were reared from caterpillars collected in late instars. Furthermore, for at least one species of solitary wasp, host size influenced sex allocation of offspring by ovipositing females. 4. Host-plant quality indirectly affected the size attained by a tachinid fly parasitoid through its direct effects on the size and performance of the caterpillar host. The host plants that resulted in the highest caterpillar host performance in the absence of enemies also yielded the largest parasitoid flies, which suggests that host plant quality can cascade up to influence the third trophic level.

Effects of Ingested Secondary Metabolites on the Immune Response of a Polyphagous Caterpillar Grammia incorrupta

We considered the effects of plant secondary metabolites on the immune response, a key physiological defense of herbivores against pathogens and parasitoids. We tested the effect of host plant species and ingested iridoid glycosides on the immune response of the grazing, polyphagous caterpillar, Grammia incorrupta (Arctiidae). Individuals of G. incorrupta were fed either one of three plant diets with varying secondary metabolites, or an artificial diet with high or low concentrations of iridoid glycosides. An immune challenge was presented, followed by measurement of the encapsulation response. We failed to detect a significant difference in the immune response of G. incorrupta feeding on diets with varying concentrations of iridoid glycosides, or feeding on different host plants. However, the immune response was lower in caterpillars consuming the artificial diet compared to those consuming the plant diets. When caterpillar performance was measured, pupal weights were lower when caterpillars ingested high concentrations of iridoid glycosides due to a decrease in feeding efficiency. Overall, individuals of G. incorrupta that consumed different plant diets exhibited a high immune response with low variation. We conclude that the immune response of G. incorrupta is adapted to feeding on a variety of plants, which may contribute to the maintenance of this caterpillar's polyphagous habit.

Complex effects of parasitoids on pharmacophagy and diet choice of a polyphagous caterpillar

This study investigates complex effects of parasitoid infection on herbivore diet choice. Specifically, we examine how immunological resistance, parasitoid infection stage, and parasitoid taxonomic identity affect the pharmacophagous behavior of the polyphagous caterpillar, Grammia incorrupta (Arctiidae). Using a combination of lab and field experiments, we test the caterpillar's pharmacophagous response to pyrrolizidine alkaloids (PAs) over the course of parasitoid infection, as well as the effect of dietary PAs on the caterpillar's immunological response. Previous work from other systems gave the prediction that dietary PAs would be detrimental to the immune response and thus less acceptable to feeding early in the infection, when encapsulation of the parasitoid is most crucial. We found that the feeding acceptability of PAs was indeed low for caterpillars with early-stage parasitoid infections; however, this was not explained by PA interference with immune function. When allowed to choose among three host plant species, individuals harboring early-stage parasitoids increased their consumption of a nutritious plant containing antioxidants. This result was driven by wasp-parasitized caterpillars, whereas fly-parasitized caterpillars increased their consumption of plants containing iridoid glycosides. Individuals in the later time phase of infection exhibited an increase in PA intake that was consistent with previously reported self-medication behavior during late-stage parasitoid infection. This study reveals the depth of complexity and the dynamic nature of herbivore host plant choice, and underscores the importance of considering multitrophic interactions when studying insect diet choice.

Influence of iridoid glycoside containing host plants on midgut β-glucosidase activity in a polyphagous caterpillar, Spilosoma virginica Fabricius (Arctiidae)

Iridoid glycosides are secondary plant compounds that have deterrent, growth reducing or even toxic effects on non-adapted herbivorous insects. To investigate the effects of iridoid glycoside containing plants on the digestive metabolism of a generalist herbivore, larvae of Spilosoma virginica (Lepidoptera: Arctiidae) were reared on three plant species that differ in their secondary plant chemistry: Taraxacum officinale (no iridoid glycosides), Plantago major (low iridoid glycoside content), and P. lanceolata (high iridoid glycoside content). Midguts of fifth instar larvae were assayed for the activity and kinetic properties of β-glucosidase using different substrates. Compared to the larvae on T. officinale, the β-glucosidase activity of larvae feeding on P. lanceolata was significantly lower measured with 4-nitrophenyl-β-d-glucopyranoside. Using the iridoid glycoside aucubin as a substrate, we did not find differences in the β-glucosidase activity of the larvae reared on the three plants. Heat inactivation experiments revealed the existence of a heat-labile and a more heat-stable β-glucosidase with similar Michaelis constants for 4-nitrophenyl-β-d-glucopyranoside. We discuss possible mechanisms leading to the observed decrease of β-glucosidase activity for larvae reared on P. lanceolata and its relevance for generalist herbivores in adapting to iridoid glycoside containing plant species and their use as potential host plants.