Genotype and environment determine allocation to and costs of resistance in quaking aspen

Although genetic variability and resource availability both influence plant chemical composition, little is known about how these factors interact to modulate costs of resistance, expressed as negative correlations between growth and defense. We evaluated genotype × environment effects on foliar chemistry and growth of quaking aspen (Populus tremuloides) by growing multiple aspen genotypes under variable conditions of light and soil nutrient availability in a common garden. Foliage was analyzed for levels of nitrogen, phenolic glycosides and condensed tannins. Bioassays of leaf quality were conducted with fourth-stadium gypsy moth (Lymantria dispar) larvae. Results revealed strong effects of plant genotype, light availability and nutrient availability; the importance of each factor depended upon compound type. For example, tannin concentrations differed little among genotypes and across nutrient regimes under low light conditions, but markedly so under high light conditions. Phenolic glycoside concentrations, in contrast, were largely determined by genotype. Variation in phenolic glycoside concentrations among genotypes was the most important factor affecting gypsy moth performance. Gypsy moth biomass and development time were negatively and positively correlated, respectively, with phenolic glycoside levels. Allocation to phenolic glycosides appeared to be costly in terms of growth, but only under resource-limiting conditions. Context-dependent trade-offs help to explain why costs of allocation to resistance are often difficult to demonstrate.     

Effects of elicitation treatment and genotypic variation on induced resistance in Populus: impacts on gypsy moth (Lepidoptera: Lymantriidae) development and feeding behavior

We examined the effects of various wounding treatments and genotypic variation on induced resistance in Populus (Salicales: Salicaceae) against herbivory by the gypsy moth, Lymantria dispar L. (Lepidoptera: Lymantriidae). Second-instar larvae grew and consumed less on leaves from induced than non-induced trees. Likewise, larvae preferred leaf disks from non-induced trees. Among induction treatments, gypsy moth feeding had the strongest and most consistent effect in behavioral choice tests. Mechanical wounding of leaves and mechanical wounding plus application of gypsy moth regurgitant had intermediate effects, while application of jasmonic acid had the weakest overall effect. Under no-choice conditions, there were no consistent trends across clones in the ability of various treatments to elicit plant responses affecting the herbivore. Levels of constitutive and inducible resistance to herbivory varied significantly among 12 Populus clones. Larvae grew up to 30-fold more, and consumed up to 250-fold more on the most suitable than the least suitable clone. Prior feeding by gypsy moths reduced larval feeding up to 71.4% on the most highly inducible clone, but it had little or no effect for the least inducible clones. There was no evidence for a relationship between levels of inducible and constitutive resistance, or between inducible resistance and phylogenetic relatedness among clones. We discuss implications for the ecology and evolution of plant-insect interactions and the management of insect pests. Received: 12 October 1998 / Accepted: 22 March 1999     

Effects of leaf maturity and wind stress on the nutrition of the generalist caterpillar Lymantria dispar feeding on poplar

The growth rates of insect herbivores commonly decrease when they feed on mature leaves due to the combined effects of several nutritional and physiological mechanisms. Environmental stresses during leaf development may also decrease herbivore performance. The present study tests two main hypotheses to help clarify the importance of these factors for the nutrition and growth of an insect herbivore: (i) decreases in nutrient levels, consumption rates and nutrient assimilation efficiencies impact negatively on herbivores feeding on mature leaves and (ii) wind stress has a negative impact on herbivores feeding on mature leaves. The results show that mature poplar (Populus alba × Populus tremula) leaves have decreased levels of protein and increased levels of fibre, and that growth rates of gypsy moth (Lymantria dispar L.) are decreased on mature leaves in association with decreased consumption rates. However, in contrast to the first hypothesis, protein and carbohydrate are assimilated efficiently (74–82% and 84–87%, respectively) from immature and mature poplar leaves. The larvae are able to chew mature leaves as efficiently as immature leaves, potentially maximizing nutrient extraction. By contrast to the second hypothesis, wind‐stressed leaves have no significant detrimental effects on nutrient assimilation efficiencies, and the lower growth rates of L. dispar larvae feeding on mature wind‐stressed leaves can be explained by lower consumption rates. Therefore, the availability of nutrients to herbivores feeding on mature tree leaves is not necessarily impacted by lower assimilation efficiencies, even when leaves develop under wind stress. These results help explain some of the large variation between the nutritional qualities of trees for forest Lepidoptera.     

Interactions between a fungal pathogen, foliar properties, and generalist herbivores

American [Castanea dentata (Marsh) Borkh.] × Chinese [Castanea mollissima Blume] chestnut (Fagac, ae) hybrids are a novel system in which to study influences of phytopathogenic fungi and woody plant hybridization on herbivore susceptibility, as the hybrids are well characterized with regard to resistance to the chestnut blight fungus [Cryphonectria parasita (Murr) Barr (Endothia) Diaporthales: Valsaceae] and variability is present. We chose two groups of resistance‐rated backcross chestnut that shared an F1 parent and had different American parents. Foliage from both backcross groups and the parent trees was sampled on three dates for use in feeding assays with gypsy moth larvae [Lymantria dispar (L.) [Lepidoptera: Lymantriidae], adult Japanese beetles [Popillia japonica Newman (Coleoptera: Scarabaeidae)], and fall webworm larvae [Hyphantria cunea Drury (Lepidoptera: Arctiidae)], respectively. Foliar analyses were performed concurrently and included carbohydrate, tannin, and nitrogen content, toughness, and density. Blight resistance had almost no effect on herbivore performance or foliar chemistry. When the parent trees and backcross groups were compared, however, significant differences in gypsy moth performance and Japanese beetle consumption were evident. There were no differences in fall webworm consumption. Most foliar characteristics measured differed among chestnut genotypes at some point in the season, and all varied seasonally. No clear pattern emerges with respect to the relationship among blight resistance, herbivore susceptibility, foliar properties, and plant genotype, and more research is needed to separate these effects.     

Relative Importance of Environmental Stress and Herbivory in Reducing Litter Fall in a Semiarid Woodland

We examined the impact of soil stress (low water and nutrient availabilities) and two keystone insect herbivores on pinyon pine (Pinus edulis) needle litterfall. We compared trees growing on two distinct soil types: volcanic cinders, which exhibit pronounced water and nutrient limitation, and sandy-loam soils, which have higher water-storage capacity and nutrient availability. Using two long-term herbivore removal experiments (15 and 18 years, respectively), we also examined the effects of the pinyon needle scale (Matsucoccus acalyptus, which attacks juvenile trees) and the stem-boring moth (Dioryctria albovittella, which attacks mature trees) on pinyon litterfall. These herbivores reach high densities on cinder soils but are absent or occur at much lower levels on sandy-loam soils. Four years of litterfall measurements showed four major patterns. First, independent of herbivory, needle litterfall was 20% lower under trees on high-stress cinder soils than on sandy-loam soils. Second, in agreement with the negative impact of scales on tree growth (that is, a 30% decline in stem growth), trees with scale infestations had 25% lower litterfall rates than trees resistant to scale; however, 15 years of scale-insect removal did not significantly increase needle litterfall. This implies possible intrinsic differences in litter production between scale-resistant and scale-susceptible trees. Third, in contrast with significant negative effects of moth herbivory on tree growth (that is, a 27% decline in stem growth), moth herbivory had no effect on needle litterfall. This, along with increased stem density in moth-susceptible trees, may be evidence of compensatory production. Fourth, there were strong year by soil type and year by scale herbivory interactions, such that in some years the effect on litterfall can be obscured or reversed by some other factor. In summary, soil stress has a strong and predictable effect on needle litterfall, whereas the relationship between insect herbivory and needle litterfall is weaker and depends on the individual herbivore. These effects, however, are mediated by other environmental factors that have considerable annual variation.     

Effect of herbivory on growth and biomass allocation of Brazilian peppertree (Sapindales: Anacardiaceae) seedlings in the laboratory

Brazilian peppertree (Schinus terebinthifolius Raddi), native to South America, is invading many ecosystems in south and central Florida. The defoliating tortricid moth Episimus unguiculus Clarke was selected as a potential biocontrol agent of Brazilian peppertree in Florida. The objective of this study was to examine the effect of different levels of herbivore damage on growth and biomass allocation of Brazilian peppertree seedlings in the laboratory. Three treatments were established: (1) no herbivory (control), (2) low herbivory (~4 larvae/plant), and (3) high herbivory (~12 larvae/plant). High levels of herbivory significantly reduced the number of leaflets, plant height, foliar biomass, foliar relative growth rate (RGR) and shoot: root ratio of Brazilian peppertree seedlings. Moreover, plants were not able to recover from herbivory after 2 months. The performance of Brazilian peppertree subjected to low herbivory levels did not differ from the control plants (no herbivory). The potential effectiveness of the biocontrol agent E. unguiculus to suppress this noxious weed is examined.     

Herbivore‐induced poplar cytochrome P450 enzymes of the CYP71 family convert aldoximes to nitriles which repel a generalist caterpillar

Numerous plant species emit volatile nitriles upon herbivory, but the biosynthesis as well as the relevance of these nitrogenous compounds in plant–insect interactions remains unknown. Populus trichocarpa has been shown to produce a complex blend of nitrogenous volatiles, including aldoximes and nitriles, after herbivore attack. The aldoximes were previously reported to be derived from amino acids by the action of cytochrome P450 enzymes of the CYP79 family. Here we show that nitriles are derived from aldoximes by another type of P450 enzyme in P. trichocarpa. First, feeding of deuterium‐labeled phenylacetaldoxime to poplar leaves resulted in incorporation of the label into benzyl cyanide, demonstrating that poplar volatile nitriles are derived from aldoximes. Then two P450 enzymes, CYP71B40v3 and CYP71B41v2, were characterized that produce aliphatic and aromatic nitriles from their respective aldoxime precursors. Both possess typical P450 sequence motifs but do not require added NADPH or cytochrome P450 reductase for catalysis. Since both enzymes are expressed after feeding by gypsy moth caterpillars, they are likely to be involved in herbivore‐induced volatile nitrile emission in P. trichocarpa. Olfactometer experiments showed that these volatile nitriles have a strong repellent activity against gypsy moth caterpillars, suggesting they play a role in induced direct defense against poplar herbivores.     

Facilitation between invasive herbivores: hemlock woolly adelgid increases gypsy moth preference for and performance on eastern hemlock

1. Interactions between invertebrate herbivores with different feeding modes are common on long-lived woody plants. In cases where one herbivore facilitates the success of another, the consequences for their shared host plant may be severe. Eastern hemlock (Tsuga canadensis), a canopy-dominant conifer native to the eastern U.S., is currently threatened with extirpation by the invasive stylet-feeding hemlock woolly adelgid (Adelges tsugae). The effect of adelgid on invasive hemlock-feeding folivores remains unknown. 2. This study evaluated the impact of feeding by hemlock woolly adelgid on gypsy moth (Lymantria dispar) larval preference for, and performance on, eastern hemlock. To assess preference, 245 field-grown hemlocks were surveyed for gypsy moth herbivory damage and laboratory paired-choice bioassays were conducted. To assess performance, gypsy moth larvae were reared to pupation on adelgid-infested or uninfested hemlock foliage, and pupal weight, proportional weight gain, and larval period were analysed. 3. Adelgid-infested hemlocks experienced more gypsy moth herbivory than did uninfested control trees, and laboratory tests confirmed that gypsy moth larvae preferentially feed on adelgid-infested hemlock foliage. Gypsy moth larvae reared to pupation on adelgid-infested foliage gained more weight than larvae reared on uninfested control foliage. 4. These results suggest that the synergistic effect of adelgid and gypsy moth poses an additional threat to eastern hemlock that may increase extirpation risk and ecological impact throughout most of its range.     

Aboveground resource allocation in response to root herbivory as affected by the arbuscular mycorrhizal symbiosis

Aims

Arbuscular mycorrhizal (AM) fungi associate with the majority of terrestrial plants, influencing their growth, nutrient uptake and defence chemistry. Consequently, AM fungi can significantly impact plant-herbivore interactions, yet surprisingly few studies have investigated how AM fungi affect plant responses to root herbivores. This study aimed to investigate how AM fungi affect plant tolerance mechanisms to belowground herbivory.

Methods

We examined how AM fungi affect plant (Saccharum spp. hybrid) growth, nutrient dynamics and secondary chemistry (phenolics) in response to attack from a root-feeding insect (Dermolepida albohirtum).

Results

Root herbivory reduced root mass by almost 27%. In response, plants augmented investment in aboveground biomass by 25%, as well as increasing carbon concentrations. The AM fungi increased aboveground biomass, phosphorus and carbon. Meanwhile, root herbivory increased foliar phenolics by 31% in mycorrhizal plants, and increased arbuscular colonisation of roots by 75% overall. AM fungi also decreased herbivore performance, potentially via increasing root silicon concentrations.

Conclusions

Our results suggest that AM fungi may be able to augment plant tolerance to root herbivory via resource allocation aboveground and, at the same time, enhance plant root resistance by increasing root silicon. The ability of AM fungi to facilitate resource allocation aboveground in this way may be a more widespread strategy for plants to cope with belowground herbivory.

     

Compound effects of induced plant responses on insect herbivores and parasitoids: implications for tritrophic interactions

1. Induced plant responses can affect herbivores either directly, by reducing herbivore development, or indirectly, by affecting the performance of natural enemies. Both the direct and indirect impacts of induction on herbivore and parasitoid success were evaluated in a common experimental system, using clonal poplar trees Populus nigra (Salicales: Salicaceae), the gypsy moth Lymantria dispar (L.) (Lepidoptera: Lymantriidae), and the gregarious parasitoid Glyptapanteles flavicoxis (Marsh) (Hymenoptera: Braconidae). 2. Female parasitoids were attracted to leaf odours from both damaged and undamaged trees, however herbivore‐damaged leaves were three times more attractive to wasps than undamaged leaves. Parasitoids were also attracted to herbivore larvae reared on foliage and to larval frass, but they were not attracted to larvae reared on artificial diet. 3. Prior gypsy moth feeding elicited a systemic plant response that retarded the growth rate, feeding, and survival of gypsy moth larvae, however induction also reduced the developmental success of the parasitoid. 4. The mean number of parasitoid progeny emerging from hosts fed foliage from induced trees was 40% less than from uninduced trees. In addition, the proportion of parasitised larvae that survived long enough to issue any parasitoids was lower on foliage from induced trees. 5. A conceptual and analytical model is provided to describe the net impacts of induced plant responses on parasitoids, and implications for tritrophic interactions and biological control of insect pests are discussed.     

Effects of elevated ultraviolet-B radiation on a plant–herbivore interaction

Enhanced ultraviolet-B (UV-B) radiation may have multiple effects on both plants and animals and affect plant–herbivore interactions directly and indirectly by inducing changes in host plant quality. In this study, we examined combined effects of UV-B and herbivory on the defence of the mountain birch (Betula pubescens ssp. czerepanovii) and also the effects of enhanced UV-B radiation on a geometrid with an outbreak cycle: the autumnal moth (Epirrita autumnata). We established an experiment mimicking ozone depletion of 30% (a relevant level when simulating ozone depletion above Northern Lapland). Both arctic species responded only slightly to the enhanced level of UV-B radiation, which may indicate that these species are already adapted to a broader range of UV-B radiation. UV-B exposure slightly induced the accumulation of myricetin glycosides but had no significant effect on the contents of quercetin or kaempferol derivatives. Mountain birch seedlings responded more efficiently to herbivory wounding than to enhanced UV-B exposure. Herbivory induced the activities of foliar oxidases that had earlier been shown to impair both feeding and growth of moth larvae. In contrast, the contents of foliar phenolics did not show the same response in different clones, except for a decrease in the contents of tannin precursors. The induction of foliar phenoloxidase activities is a specific defence response of mountain birches against insect herbivory. To conclude, our results do not support the hypothesis that the outbreak cycle of the autumnal moth can be explained by the cycles of solar activity and UV-B.     

Aboveground phytochemical responses to belowground herbivory in poplar trees and the consequence for leaf herbivore preference

Belowground (BG) herbivory can influence aboveground (AG) herbivore performance and food preference via changes in plant chemistry. Most evidence for this phenomenon derives from studies in herbaceous plants but studies in woody plants are scarce. Here we investigated whether and how BG herbivory on black poplar (Populus nigra) trees by Melolontha melolontha larvae influences the feeding preference of Lymantria dispar (gypsy moth) caterpillars. In a food choice assay, caterpillars preferred to feed on leaves from trees that had experienced attack by BG herbivores. Therefore, we investigated the effect of BG herbivory on the phytochemical composition of P. nigra trees alone and in combination with AG feeding by L. dispar caterpillars. BG herbivory did not increase systemic AG tree defences like volatile organic compounds, protease inhibitors and salicinoids. Jasmonates and salicylic acid were also not induced by BG herbivory in leaves but abscisic acid concentrations drastically increased together with proline and few other amino acids. Leaf coating experiments with amino acids suggest that proline might be responsible for the caterpillar feeding preference via presumptive phagostimulatory properties. This study shows that BG herbivory in poplar can modify the feeding preference of AG herbivores via phytochemical changes as a consequence of root‐to‐shoot signaling.     

Potential interactions between invasive woody shrubs and the gypsy moth (Lymantria dispar), an invasive insect herbivore

As the range of the invasive and highly polyphagous gypsy moth (Lymantria dispar) expands, it increasingly overlaps with forest areas that have been subject to invasion by non-native shrubs. We explored the potential for interactions between these co-occurring invasions through a gypsy moth feeding trial using the following three highly invasive, exotic shrubs: honeysuckle (Lonicera maackii), privet (Ligustrum sinense) and burning bush (Euonymus alatus). We compared these with two native shrubs: spicebush (Lindera benzoin) and pawpaw (Asimina triloba). We fed gypsy moth caterpillars foliage exclusively from one of the five shrubs and measured their relative consumptive rate (RCR), relative growth rate (RGR), and development time (DT). The RCR of gypsy moth was strongly influenced by the species of foliage (F = 31.9; P < 0.0001) with little or no consumption of honeysuckle and privet. Caterpillar RGR was influenced by the shrub species (F = 66.2; P < 0.0001), and those caterpillars fed spicebush, honeysuckle or privet lost weight over the course of the assay. Caterpillar DT was also significantly (F = 11.79, P < 0.0001) influenced by the shrub species and those fed honeysuckle, privet and spicebush died prior to molting. Overall, our data suggest that honeysuckle, privet, and spicebush could benefit (indirectly) from the invasion of gypsy moth, while burning bush and pawpaw could be negatively impacted due to direct effects (herbivory). Similarly, invading gypsy moth populations could be sustained on a shrub layer of burning bush and pawpaw in the event of canopy defoliation. Further field and laboratory analysis is needed to clarify herbivore resistance of invasive shrubs, and to investigate the potential interactions among co-occurring insect and plant invasions.     

Soil carbon and nitrogen mineralization following deposition of insect frass and greenfall from forests under elevated CO2 and O3

Elevated CO₂ and O₃ alter tree quality and the quality of herbivore inputs, such as frass, to forest soil. Altered quality or quantity of herbivore inputs to the forest floor can have large impacts on belowground processes. We collected green leaves and frass from whitemarked tussock moth caterpillars from aspen-birch stands at the Aspen Free Air CO₂ Enrichment (FACE) site near Rhinelander, WI, USA. Small or large quantities of frass, greenfall, or a 1:1 ratio of frass and greenfall were added to microcosms for each FACE treatment (control, +CO₂, +O₃, +CO₂+O₃). We measured initial frass and greenfall quality, and recorded microbial respiration, and nitrate leaching over 40 days. Elevated carbon dioxide (eCO₂) and tropospheric ozone (eO₃) significantly altered the carbon, nitrogen, and condensed tannin content of insect frass and green leaves. Although FACE treatments affected input quality, they had minimal effect on microbial respiration and no effect on nitrogen leaching. In contrast, input quantity substantially influenced microbial respiration and nitrate leaching. Respiratory carbon loss and nitrate immobilization were nearly double in microcosms receiving large amounts of herbivore inputs than those receiving no herbivore inputs. Small amounts of herbivore inputs, however, did not significantly alter microbial respiration or immobilization, suggesting that effects of herbivore inputs on soil processes will be detected only at moderate to high herbivory/input levels. These results suggest that subtle changes in frass and greenfall quality may not affect soil nutrient cycling. In contrast, environmental change induced increases in insect population size or frass and greenfall inputs to the soil may substantially impact nutrient cycling.     

Physiological and growth responses of Centaurea maculosa (Asteraceae) to root herbivory under varying levels of interspecific plant competition and soil nitrogen availability

Summary Centaurea maculosa seedlings were grown in pots to study the effects of root herbivory by Agapeta zoegana L. (Lep.: Cochylidae) and Cyphocleonus achates Fahr. (Col.: Curculionidae), grass competition and nitrogen shortage (each present or absent), using a full factorial design. The aims of the study were to analyse the impact of root herbivory on plant growth, resource allocation and physiological processes, and to test if these plant responses to herbivory were influenced by plant competition and nitrogen availability. The two root herbivores differed markedly in their impact on plant growth. While feeding by the moth A. zoegana in the root cortex had no effect on shoot and root mass, feeding by the weevil C. achates in the central vascular tissue greatly reduced shoot mass, but not root mass, leading to a reduced shoot/root ratio. The absence of significant effects of the two herbivores on root biomass, despite considerable consumption, indicates that compensatory root growth occurred. Competition with grass affected plant growth more than herbivory and nutrient status, resulting in reduced shoot and root growth, and number of leaves. Nitrogen shortage did not affect plant growth directly but greatly influenced the compensatory capacity of Centaurea maculosa to root herbivory. Under high nitrogen conditions, shoot biomass of plants infested by the weevil was reduced by 30% compared with uninfested plants. However, under poor nitrogen conditions a 63% reduction was observed compared with corresponding controls. Root herbivory was the most important stress factor affecting plant physiology. Besides a relative increase in biomass allocation to the roots, infested plants also showed a significant increase in nitrogen concentration in the roots and a concomitant reduction in leaf nitrogen concentration, reflecting a redirection of the nitrogen to the stronger sink. The level of fructans was greatly reduced in the roots after herbivore feeding. This is thought to be a consequence of their mobilisation to support compensatory root growth. A preliminary model linking the effects of these root herbivores to the physiological processes of C. maculosa is presented.     

Atmospheric change alters performance of an invasive forest insect

Atmospheric change and species invasions are arguably two of the most important factors affecting the long‐term sustainability of natural ecosystems. We examined the independent and interactive effects of atmospheric carbon dioxide (CO₂) and tropospheric ozone (O₃) on the foliar quality of two host species and performance of an invasive folivorous insect. Trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera) were grown at the Aspen FACE research site in northern Wisconsin, USA, under all combinations of ambient and elevated CO₂ and O₃. We measured the effects of elevated CO₂ and O₃ on aspen and birch phytochemistry and on the survivorship, development time, growth, and fecundity of the gypsy moth (Lymantria dispar). Elevated CO₂ had little effect on, whereas elevated O₃ altered, the composite phytochemical profiles of aspen and birch. Nutritional quality in aspen and birch leaves was marginally affected by elevated CO₂ and reduced by elevated O₃. Both gases increased concentrations of phenolic and structural compounds in aspen and birch. Elevated CO₂ offset reduced foliar quality under elevated O₃, but only in aspen, and to a greater extent later than earlier in spring. Elevated CO₂ generally had beneficial effects on, while elevated O₃ detrimentally affected, gypsy moth performance. Elevated CO₂ ameliorated most of the reductions in gypsy moth performance under elevated O₃. Our findings suggest that atmospheric change can alter foliar quality in gypsy moth hosts sufficiently to influence gypsy moth performance, but that these responses will depend on interactions among CO₂, O₃, and tree species. Our findings also contrast with those of earlier studies at Aspen FACE, indicating that foliar quality responses to environmental change are likely influenced by tree stand age and longevity of exposure to pollutants to the extent that they affect plant‐herbivore interactions differently over decadal time spans.     

Metabolism of poplar salicinoids by the generalist herbivore Lymantria dispar (Lepidoptera)

The survival of insect herbivores on chemically defended plants may often depend on their ability to metabolize these defense compounds. However, only little knowledge is available on how insects actually process most plant defense compounds. We investigated the metabolism of salicinoids, a major group of phenolic glycosides in poplar and willow species, by a generalist herbivore, the gypsy moth (Lymantria dispar). Seven salicinoid metabolites identified in gypsy moth caterpillar feces were mostly conjugates with glucose, cysteine or glycine. Two of the glucosides were phosphorylated, a feature not previously reported for insect metabolites of plant defense compounds. The origins of these metabolites were traced to specific moieties of three major poplar salicinoids ingested, salicin, salicortin and tremulacin. Based on the observed metabolite patterns we were able to deduce the initial steps of salicinoid breakdown in L. dispar guts, which involves cleavage of ester bonds. The conjugated molecules were effectively eliminated within 24 h after ingestion. Some of the initial breakdown products (salicin and catechol) demonstrated negative effects on insect growth and survival in bioassays on artificial diets. Gypsy moth caterpillars with prior feeding experience on salicinoid-containing poplar foliage converted salicinoids to the identified metabolites more efficiently than caterpillars pre-fed an artificial diet. The majority of the metabolites we identified were also produced by other common poplar-feeding insects. The conversion of plant defenses like salicinoids to a variety of water-soluble sugar, phosphate and amino acid conjugates and their subsequent excretion fits the general detoxification strategy found in insect herbivores and other animals.     

Additive effects of aboveground and belowground herbivores on the dominance of spotted knapweed (Centaurea stoebe)

Spotted knapweed (Centaurea stoebe) is found in over 3 million ha of rangeland and forests across North America, and evidence supporting the use of biological control as a regional method to reduce infestations and their associated impacts remains inconclusive. Several species of insects have been reported to reduce plant densities in some areas; however, rigorous studies that test combinations of these species and the influence of resource availability are lacking. We examined the singular and combined effects of herbivory by a root weevil (Cyphocleonus achates) and a flower head weevil (Larinus minutus) on the growth and flower production of C. stoebe. We also manipulated soil resource fertility as an additional factor that could explain the outcomes of contradictory biological control herbivore effects on C. stoebe. In a greenhouse study, herbivory by C. achates decreased flower production for plants across all resource environments. In a caged common garden study, the negative effects of herbivory also did not interact with soil nutrient status. However, the presence of plant competition further decreased knapweed growth, and the negative effects of concurrent herbivory by C. achates and L. minutus on plant biomass and flower production were additive. Derived within the context of variable levels of soil nutrient availability and competing vegetation, these results support the cumulative stress hypothesis and the contention that combined above- and belowground herbivory can reduce spotted knapweed densities and reduce the ecological and economic impacts of this species in rangelands of western North America.     

Herbivore‐induced plant responses in Brassica oleracea prevail over effects of constitutive resistance and result in enhanced herbivore attack

1. Plant responses to herbivore attack may have community‐wide effects on the composition of the plant‐associated insect community. Thereby, plant responses to an early‐season herbivore may have profound consequences for the amount and type of future attack. 2. Here we studied the effect of early‐season herbivory by caterpillars of Pieris rapae on the composition of the insect herbivore community on domesticated Brassica oleracea plants. We compared the effect of herbivory on two cultivars that differ in the degree of susceptibility to herbivores to analyse whether induced plant responses supersede differences caused by constitutive resistance. 3. Early‐season herbivory affected the herbivore community, having contrasting effects on different herbivore species, while these effects were similar on the two cultivars. Generalist insect herbivores avoided plants that had been induced, whereas these plants were colonised preferentially by specialist herbivores belonging to both leaf‐chewing and sap‐sucking guilds. 4. Our results show that community‐wide effects of early‐season herbivory may prevail over effects of constitutive plant resistance. Induced responses triggered by prior herbivory may lead to an increase in susceptibility to the dominant specialists in the herbivorous insect community. The outcome of the balance between contrasting responses of herbivorous community members to induced plants therefore determines whether induced plant responses result in enhanced plant resistance.