Prior Herbivory Research Articles

Species‐specific herbivore–herbivore interactions and plant responses to sequential attack by multiple herbivores on a perennial woody plant

AbstractWhen plants are sequentially attacked by multiple herbivores, herbivore identity and host specialization can greatly influence the patterns of herbivore–herbivore and plant–herbivore interactions. However, how prior herbivory and the resulting induced plant responses potentially affect subsequent herbivores deserves further investigation. In this study, we conducted a common‐garden experiment that manipulated sequential herbivory by the specialist caterpillar Gadirtha fusca Pogue (Lepidoptera: Nolidae) and the generalist caterpillar Spodoptera litura (Fabricius) (Lepidoptera: Noctuidae) on Chinese tallow, Triadica sebifera (L.) Small (Euphorbiaceae). We tested how prior exposure to herbivores with different levels of host specialization affected the performance of subsequently arriving con‐ and heterospecifics, as well as plant growth and defense responses under subsequent herbivory. We found that prior exposure to the specialist G. fusca facilitated the performance of subsequent conspecifics, resulting in a significant decrease in the growth (height and stem diameter at ground level) of tallow plants. However, prior exposure to the generalist S. litura did not affect the feeding of subsequent con‐ or heterospecifics or the growth of tallow plants. Sequential herbivory by specialist and generalist conspecifics resulted in lower levels of tannins and flavonoids, respectively, in leaves of tallow plants, whereas sequential herbivory by the two species did not affect the levels of tannins or flavonoids, compared to a single damage event. We conclude that herbivore species‐specific plant responses appear to be more important than herbivore identity or specialization in determining herbivore–herbivore interactions and plant responses to sequential herbivore attack.

Stem borer herbivory dependent on interactions of sugarcane variety, associated traits, and presence of prior borer damage.

Herbivory risk is mediated by plant traits related to nutrition and defense that can vary within a species by genotype and age. Prior herbivore damage accrued by a plant can also interact with these traits to alter future herbivory potential by changing plant quantity or quality. Sugarcane (Saccharum spp.) is a perennial crop where aboveground biomass is harvested annually and with varieties differing in nutrition and defenses, making it conducive to evaluating varietal resistance mechanisms. Using data from 16 sugarcane varieties and 28 years, we assessed damage from the primary pest in Louisiana, the sugarcane borer (Diatraea saccharalis, SCB), relative to variety, crop year (ratoon), plant traits, and incidence of prior herbivory. SCB damage differed among varieties but not crop year, mostly following previously established classifications of SCB resistance, and correlated with select nutritional and defense traits. Within a crop year, the probability of SCB damage increased with prior conspecific damage on the same stalk. However, the strength of this prior damage effect did not match known resistance patterns but still differed with variety. Interactions of plant variety, traits, and prior pest damage but not age impacted sugarcane borer risk. Borer damage was associated with nutritional traits of fiber and sugar content, but not consistently with defensive traits like high stalk wax or hair density, indicating there may be additional resistance traits or indirect impacts of these traits on predators. Published 2023. This article is a U.S. Government work and is in the public domain in the USA.

Impact of early insect herbivory on the invasive oak lace bug (Corythucha arcuata Say, 1832) in different oak species

Insect herbivores co-occurring on the same host plant interact in various ways. In particular, early-season insect herbivory triggers a wide range of plant responses that can determine the performance of herbivores colonizing the plant later in the course of the season. But the strength and direction of such effects are debated, and virtually unknown in the case of novel interactions involving exotic insects in their introduction range. We conducted an observational field study in SW France, a region recently invaded by the Oak Lace Bug (OLB, Corythucha arcuata Say). We measured early chewing damage and subsequent OLB damage in four oak species (Quercus robur, Q. pubescens, Q. cerris and Q. ilex). We set up a complementary non-choice experiment in the laboratory, feeding OLB with leaves with or without prior herbivory. The four oak species differed in their sensitivity to OLB damage, Q. ilex being broadly resistant. Prior herbivory promoted OLB damage in the laboratory experiment, but not in the field. However, prior herbivory did not alter the rank of oak resistance to the OLB. Our results suggest possible synergistic effects between spring defoliators and the OLB. This study brings insight into herbivore-herbivore interactions and their possible implications for forest management.

Arbuscular mycorrhizal fungi mediate how plant herbivory history influences herbivore performance

Abstract Herbivore performance is influenced by plant host quality and chemical defences, both of which are influenced by belowground symbionts of plants and herbivory history. The previous herbivory may suppress or facilitate subsequent herbivore performance through changes in plant chemistry and tissue quality. Likewise, arbuscular mycorrhizal fungi (AM fungi) can alter both plant quality and plant responses to aboveground herbivory, as mycorrhizal‐mediated alterations of the nutritional budget may alter leaf nutritional quality, chemical profiles, and available leaf surface area. Despite evidence that both herbivory history and AM fungi can independently affect herbivore performance, few studies directly test both at the same time. Here, the role of AM fungi and herbivory history on the performance of a pest insect (Spodoptera frugiperda) on a common forb (Solidago altissima) grown with different AM fungal additions (no AM fungi, single species, multispecies) was directly tested. Evidence that the effect of herbivory history on herbivore performance depended on the AM fungal environment of the plant was found. Specifically, the composition of the AM fungal additions altered the directionality of herbivore performance on plants with and without prior herbivory. Without added AM fungi, herbivores demonstrated a classically suppressed growth rate on grazed plants. With a single species AM fungi added, herbivores performed the same on host plants, regardless of herbivory history. With diverse AM fungi added, herbivore growth increased on grazed plants. Mycorrhizal‐mediated changes in plant chemical profiles did not explain herbivore performance. These results emphasise the need for direct tests of mycorrhizal diversity on plant‐insect interactions.

Variation in both host defense and prior herbivory can alter plant-vector-virus interactions

BackgroundWhile virus-vector-host interactions have been a major focus of both basic and applied ecological research, little is known about how different levels of plant defense interact with prior herbivory to affect these relationships. We used genetically-modified strains of tomato (Solanum lycopersicum) varying in the jasmonic acid (JA) plant defense pathways to explore how plant defense and prior herbivory affects a plant virus (tomato yellow leaf curl virus, ‘TYLCV’), its vector (the whitefly Bemisia tabaci MED), and the host.ResultsVirus-free MED preferred low-JA over high-JA plants and had lower fitness on high-JA plants. Viruliferous MED preferred low-JA plants but their survival was unaffected by JA levels. While virus-free MED did not lower plant JA levels, viruliferous MED decreased both JA levels and the expression of JA-related genes. Infestation by viruliferous MED reduced plant JA levels. In preference tests, neither virus-free nor viruliferous MED discriminated among JA-varying plants previously exposed to virus-free MED. However, both virus-free and viruliferous MED preferred low-JA plant genotypes when choosing between plants that had both been previously exposed to viruliferous MED. The enhanced preference for low-JA genotypes appears linked to the volatile compound neophytadiene, which was found only in whitefly-infested plants and at concentrations inversely related to plant JA levels.ConclusionsOur findings illustrate how plant defense can interact with prior herbivory to affect both a plant virus and its whitefly vector, and confirm the induction of neophytadiene by MED. The apparent attraction of MED to neophytadiene may prove useful in pest detection and management.

Insect Community Structure Covaries with Host Plant Chemistry but is Not Affected by Prior Herbivory

Insect Community Structure Covaries with Host Plant Chemistry but is Not Affected by Prior Herbivory

Insect community structure covaries with host plant chemistry but is not affected by prior herbivory.

By feeding on plant tissue, insect herbivores can change several characteristics of their hosts. These changes have the potential to alter the quality of the plant for other herbivore species, potentially altering the structure of the community of species attacking the plant at a later point in time. We tested whether herbivory early in the season changes host plant performance, polyphenol chemistry, and the community structure of sessile herbivores later in the season. We experimentally manipulated densities of early-season moth caterpillars on a set of young oak trees and measured tree growth, reproduction, leaf chemistry, and the abundance and community composition of leafmining and galling species later in the season. The experimental manipulations of early-season herbivores did not affect late-season leaf chemistry or tree performance. Early-season herbivores had a weak negative effect on the abundance of gallers and a positive, tree-dependent effect on the overall diversity of late-season sessile herbivores. The chemical composition of leaves covaried with the species composition of the late-season leafmining and galling community. Both the chemical composition of the host tree and the late-season insect community structure were strongly affected by the growth location of the tree. Our results suggest that plant-mediated indirect effects between herbivores might play a limited role in this system, whereas the underlying variation in plant chemistry is an important factor structuring the associated insect community. Our results emphasize that factors other than prior herbivory can be important determinants of plant chemistry and the community composition of herbivores.

Impacts of feral horses and deer on an endangered woodland of Kosciuszko National Park

SummaryIn recent years, the impacts of rapidly increasing populations of feral horses and deer on the vegetation and stability of soils have become highly visible and widespread in Kosciuszko National Park. We investigated these impacts in the White Cypress Pine (Callitris glaucophylla Joy Thomps. & L.A.S. Johnson) – White Box (Eucalyptus albens Benth) woodlands of the lower Snowy River valley. This woodland is a component of the White Box‐Yellow Box‐Blakely's Red Gum Grassy Woodland and Derived Native Grasslands complex that is nationally listed as a critically endangered ecological community. To investigate the severity of the impacts of feral horse (Equus caballus) and deer (Dama dama and Rusa unicolor) in the valley in 2013 and 2017/18, we surveyed fenced exclosures and paired grazed plots that were first established and surveyed in 1984 and re‐surveyed in 1987. Using LFA and VAST methodologies (not used in 1987), we compared the relative response of environmental variables in plots inside and outside the exclosures in an attempt to ascertain recent herbivore impacts. While there was no evidence of horses or deer from dung surveys in 1987, in 2018, 84% of the dung was from horses, 13% from deer, 1% from rabbits and 2% from macropods. Total herbivore dung density increased fourfold since the 1987 survey. On the understanding that all plots had the same starting condition in 1984 with respect to prior herbivory, we deduce that horses and deer are having significant ecological impacts. There was a far greater cover of understorey plants and the midstorey was denser and taller inside the exclosures. Outside the exclosures, the vegetation cover was far more sparse and soil erosion was active and extensive. The total number of invertebrates captured in small pitfall traps was nearly twice as many within the exclosures compared to the grazed plots. The dense even‐aged regrowth overstorey stands of White Cypress Pine, inside and outside the exclosures, have changed little in 34 years.

Effects of Intraspecific Genetic Variation and Prior Herbivory in an Old-Field Plant on the Abundance of the Specialist Aphid Uroleucon nigrotuberculatum (Hemiptera: Aphididae).

Intraspecific genetic variation in plants can contribute to the diversity and abundance of associated insects, though many questions remain about why some genotypes support more insects than others. Since plant secondary metabolites, which may be induced after insect attack, may potentially vary among genotypes, these compounds provide a possible explanation for insect abundance variation in plants with substantial genetic variation. In this study, we examined four genotypes of the old-field plant species Solidago altissima (L.; Asterales: Asteraceae) and asked if the abundance of the specialist aphid Uroleucon nigrotuberculatum (Olive; Hemiptera: Aphididae) was affected by genotype and previous foliage damage by a specialist beetle. We hypothesized that different genotypes and prior herbivory would result in different quantities of terpenes produced by S. altissima, and that terpenes would affect aphid abundance. We found evidence of foliar terpene induction in a greenhouse environment, and significant differences in terpene production among genotypes in a field setting, though prior damage had little effect on aphid abundance in the field. There were significant effects of genotypes on aphid abundance, as well as genotype effects on terpenes and foliar nutrients (leaf N and C:N). Noteworthy was a change in the allocation of particular terpenes among genotypes that related to aphid abundance. Our analyses demonstrated that phytochemicals, and especially terpenes, related to aphid abundance. This study adds to a previous finding that variation in leaf terpenes in S. altissima provides a partial explanation for variable abundance among genotypes of a specialist aphid, and suggests that differences in the allocation of compounds is important.

Parasite Removal, but Not Herbivory, Deters Future Parasite Attachment on Tomato.

Plants face many antagonistic interactions that occur sequentially. Often, plants employ defense strategies in response to the initial damage that are highly specific and can affect interactions with subsequent antagonists. In addition to herbivores and pathogens, plants face attacks by parasitic plants, but we know little about how prior herbivory compared to prior parasite attachment affects subsequent host interactions. If host plants can respond adaptively to these different damage types, we predict that prior parasitism would have a greater deterrent effect on subsequent parasites than would prior herbivory. To test the effects of prior parasitism and prior herbivory on subsequent parasitic dodder (Cuscuta spp.) preference, we conducted two separate greenhouse studies with tomato hosts (Solanum lycopersicum). In the first experiment, we tested the effects of previous dodder attachment on subsequent dodder preference on tomato hosts using three treatments: control plants that had no previous dodder attachment; dodder-removed plants that had an initial dodder seedling attached, removed and left in the same pot to simulate parasite death; and dodder-continuous plants with an initial dodder seedling that remained attached. In the second experiment, we tested the effects of previous caterpillar damage (Spodoptera exigua) and mechanical damage on future dodder attachment on tomato hosts. Dodder attached most slowly to tomato hosts that had dodder plants previously attached and then removed, compared to control plants or plants with continuous dodder attachment. In contrast, herbivory did not affect subsequent dodder attachment rate. These results indicate that dodder preference depended on the identity and the outcome of the initial attack, suggesting that early-season interactions have the potential for profound impacts on subsequent community dynamics.

Plant inbreeding and prior herbivory influence the attraction of caterpillars (Manduca sexta) to odors of the host plant Solanum carolinense (Solanaceae)

The mediation of plant-insect interactions by plant odors has been studied extensively, but most previous work has focused on documenting the role of constitutive and herbivore- or pathogen-induced plant volatiles as foraging cues for insect herbivores and their natural enemies. Relatively little work has explored genotypic variation in plant-odor profiles within species, and few studies have addressed the perception and use of olfactory cues by lepidopteran larvae or other herbivores during feeding. We examined the effects of plant breeding (inbred vs. outbred individuals) and plant exposure to prior herbivory on the preferences of caterpillars (Manduca sexta) for odors of Solanum carolinense in leaf-disc and whole-plant choice assays. Second- and third-instar larvae of M. sexta clearly and consistently preferred undamaged over herbivore-damaged plants of both breeding types and also consistently preferred inbred over outbred plants that had the same damage status. Similar preferences were observed even when plants were covered with bridal-veil cloth to mask visual cues, demonstrating that olfactory cues influence larval preferences. The observed preferences are consistent with our previous findings regarding the constitutive and induced volatile profiles of inbred and outbred horsenettle plants and their effects on plant-herbivore interactions. They furthermore correspond to differences in host-plant quality predicted by previous work and, thus, suggest that naive larvae of M. sexta can accurately assess aspects of host-plant quality via olfactory cues perceived at a distance.

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.

Herbivore- and Elicitor-Induced Resistance in Rice to the Rice Water Weevil (Lissorhoptrus oryzophilus Kuschel) in the Laboratory and Field

Feeding by herbivores can change plants in ways that make them more resistant to subsequent herbivory. Such induced responses are better-studied in a number of model dicots than in rice and other cereals. In a series of greenhouse and field experiments, we assessed the effects of prior herbivory by the fall armyworm, Spodoptera frugiperda (J.E. Smith) and of exogenous applications of jasmonic acid (JA) on the resistance of rice plants to the rice water weevil, Lissorhoptrus oryzophilus (Kuschel), the major pest of rice in the United States. Prior feeding by S. frugiperda and treatment of plants with exogenous JA resulted in increases in the resistance of plants to the weevil. Increases in resistance were manifested as reduced numbers of eggs and first-instars associated with armyworm-injured or JA-treated plants relative to control plants. In field experiments, there was a transient but significant reduction in the number of immature L. oryzophilus on JA-treated plants relative to untreated plants. To our knowledge, this is the first example of direct induced resistance in rice demonstrated in small-plot field experiments. We discuss the potential for the use of elicitor induced resistance in rice.

Potential for the use of elicitors of plant resistance in arthropod management programs.

Plants protect themselves from arthropod herbivores both directly, by expressing biochemical and morphological traits that interfere with herbivore development or behavior, and indirectly, by facilitating the action of natural enemies of herbivores. These direct and indirect resistance mechanisms are not always expressed at maximal levels by plants, but rather can be induced to higher levels by a variety of stimuli, most notably prior herbivory. The recent discovery of chemical elicitors of induced responses has led to interest in manipulating the inducible responses of plants for crop protection. Applications of elicitors of induced responses made at appropriate times during the growing season of a crop have the potential of activating both direct and indirect mechanisms of plant resistance and thereby simultaneously augmenting host-plant resistance and biological control. This strategy may serve as an important component of a multifaceted, ecologically-based pest management program and is unlikely to precipitate the rapid evolution of countermeasures by target pests. However, this strategy will not be appropriate in all crops or against all arthropod pests. The conditions under which the use of an elicitor is likely to be successful are discussed, and examples of the successful use of elicitors are reviewed.

Influence of semilooper damage on cotton host-plant resistance to Helicoverpa armigera (Hub)

The semilooper, Anomis flava Fab is a foliage feeding lepidopteran insect, often found occurring in the cotton ecosystem during the first 60–75 days of sowing. Its occurrence overlaps or just precedes the incidence of Helicoverpa armigera. This study reports the impact of mechanical wounding or of prior herbivory by semilooper larvae on cotton host-plant resistance to H. armigera. It is proposed that cotton plant damaged by semilooper or by mechanical means is less preferred for the growth of H. armigera. This is probably due to a reduction in the nutritive quality, induction of allelochemicals, such as chlorogenic acid and of oxidative enzymes, such as lipoxygenases and peroxidases in the damaged plants.

Beetle grazing reduces natural infection of Rumex obtusifolius by fungal pathogens.

In previously reported laboratory experiments, infection of Rumex obtusifolius by the rust fungus Uromyces rumicis was decreased on leaves which had prior herbivory by the beetle Gastrophysa viridula. In this paper we investigate whether this interaction is robust for natural infection by a variety of fungi in field experiments carried out in spring and autumn with plants given different levels of nitrogen fertilization. Grazing by G. viridula led to a decrease in lesion density of Ramularia rubella and Venturia rumicis in the spring and V. rumicis and U. rumicis in the autumn experiment. For V. rumicis and U. rumicis significant reductions in lesion density occurred on the undamaged leaves of damaged plants, compared with similar leaves on undamaged plants, suggesting systemic induced resistance. This induced resistance was usually independent of the amount of nitrogen fertilization, although the inhibitory effect of grazing on R. rubella in the spring and V. rumicis in the autumn experiment was enhanced by increasing nitrogen fertilization and was inhibited by increasing nitrogen fertilization for V. rumicis in the spring. In both experiments, the lesion density of V. rumicis was greater on leaves on which R. rubella was also present, and the presence of U. rumicis in the autumn experiment was linked to a similar but greater effect on V. rumicis lesion density. We found no evidence of induced resistance by fungi against fungi in these experiments. We highlight the complex interactions between inhibitory and facilitatory processes acting on leaf fungal infection. These results are compared with the proposed molecular mechanisms of induced resistance(s) and we consider the benefits of closer integration between molecular and ecological investigations of induced resistances that occur in the field.

Effects of Endophytic Fungus, Nutrient Level, and Plant Damage on Performance of Fall Armyworm (Lepidoptera: Noctuidae)

We tested the effects of soil nutrient level, endophyte infection status, and prior herbivory on the performance (survival, growth, and assimilation) of fall armyworm, Spodoptera frugiperda (J.E. Smith), Using a 3-factor design, we fed larvae all possible combinations of the following treatments: endophyte-infected or uninfected tall fescue, Festuca arundinacea von Schreber, fescue receiving high or low nutrient levels, and fescue that was either previously damaged or not previously damaged with a fabric tracing wheel. Low nutrient treatments showed a consistent negative effect on 8-d larval mass, pupal mass, development time, assimilation efficiency, and survival of fall armyworm larvae, Previous damage negatively affected larval mass, development time, and assimilation efficiency, but did not affect pupal mass or survival. Infection status also gave mixed results; infection negatively affected pupal mass, but accelerated development time. positively affected larval mass and did not affect survival or assimilation. In general, interactions among nutrient level, wounding damage, and infection status did not affect insect performance. However, we did find that the influence of endophyte on insects depended upon nutrient level supplied to the plants. The adverse effect of endophyte on pupal mass was most pronounced when plants received low fertilizer applications; under high fertilizer applications the effect of endophyte on pupal mass was negligible, Our results show that the interaction among endophytic fungi, their host grasses. and insect herbivores is complex and can be influenced by enviromnental factors, such as nutrient availability.

Influence of Interspecific and Intraspecific Host Plant Variation on the Susceptibility of Heliothines to a Baculovirus

In this paper we report on the effect of host plant variation on the susceptibility of Helicoverpa zea and Heliothis virescens larvae to the Helicoverpa zea nucleopolyhedrovirus (HzNPV). Larval H. zea and H. virescens treated with HzNPV on foliage from various hosts showed that H. virescens were significantly more susceptible to HzNPV on cotton and Carolina geranium than H. zea, whereas H. zea were significantly more susceptible to HzNPV on velvetleaf than H. virescens. Plant phenology had an effect on larval susceptibility to HzNPV. Vegetative or reproductive tissues of cotton, soybean, tomato, crimson clover, Carolina geranium, and velvetleaf were treated with HzNPV and fed to second-instar H. zea or H. virescens. Host phenology significantly affected the mortality of HzNPV-treated H. zea on all hosts except tomato. HzNPV-treated H. zea fed on vegetative tissues of crimson clover, Carolina geranium, velvetleaf, and soybean had significantly higher mortality than those fed on reproductive tissues. On the contrary, larval susceptibility to HzNPV on cotton was greater on reproductive than vegetative tissues. H. virescens susceptibility to HzNPV was greater on vegetative than reproductive tissue for crimson clover. In addition, we examined the effect of prior herbivory on larval susceptibility to the HzNPV. Cotton, soybean, tomato, and velvetleaf plants were either untreated or wounded with three fourth-instar H. zea or H. virescens. Neonates were reared on the respective treatments and then treated with HzNPV as second instars. Non-HzNPV-treated H. zea larvae fed on wounded foliage had up to 93.1% reduced weight gain compared with those fed on unwounded foliage. The weight reduction was greatest on cotton (93.1%), followed by velvetleaf (80.6%), tomato (78.4%), and soybean (54.1%). Prior herbivory on all plants significantly affected the larval growth of nontreated H. zea. In nontreated H. virescens, the reduction in weight was greatest on velvetleaf (63.1%), followed by soybean (29.2%) and cotton (21.8%). However, the effect of prior herbivory on cotton did not significantly affect the larval growth of nontreated H. virescens. Only virustreated H. zea that fed on wounded virus-treated tomato foliage showed significantly enhanced larval mortality (47.7%) compared with larvae on unwounded foliage. For HzNPV-treated H. virescens, only prior herbivory on cotton significantly enhanced larval mortality (39.4%). r 1998 Academic Press

Disentangling Effects of Induced Plant Defenses and Food Quantity on Herbivores by Fitting Nonlinear Models

Plants can respond to herbivore damage through both broad-scale (systemic) and localized induced responses. While many studies have quantified the impact of systemic responses on herbivores, measuring the impact of localized changes is difficult because plant tissues that have suffered direct damage may represent both a lower quality and a lower quantity of food. This article uses nonlinear models to disentangle the confounding effects of prior herbivory on food quantity and quality. The first (null) model assumes that herbivore performance is determined only by the quantity of food available to an average herbivore. Modified models allow two distinct effects of damage-induced defenses: an increase in the amount of food each herbivore is required to consume in order to achieve maximum performance and a reduction in the maximum performance even when herbivores are fed ad lib. Maximum likelihood methods were used to fit the models to data from field experiments in which Colorado potato beetle (Leptinotarsa decemlineata) larvae were reared on three varieties of potatoes that had been damaged to varying degrees by adult beetles. Prior damage reduced the mean mass of beetles at pupation, and this effect was due to both a decrease in food quantity and induced changes in food quality. In contrast, beetle survival was affected in some cases by reduced food quantity but showed no responses that could be attributed to induced defenses. I discuss this result in the context of previous studies of induced (mostly systemic) responses in the potato-potato beetle system, and I suggest that detailed studies of particular chemical responses and the proposed method of combining bioassays with quantitative models should be used as complementary approaches in future studies of herbivore-induced defenses in plants.

Oxidative responses in soybean foliage to herbivory by bean leaf beetle and three-cornered alfalfa hopper

Variation in induced responses in soybean is shown to be dependent, in part, upon herbivore species. Herbivory by the phloem-feeding three-cornered alfalfa hopper caused increases in the activities of several oxidative enzymes including lipoxygenases, peroxidases, ascorbate oxidase, and polyphenol oxidase. Bean leaf beetle defoliation caused increased lipoxygenase activity, but had little effect upon peroxidase, polyphenol oxidase, ascorbate oxidase, or trypsin inhibitor levels in either field or greenhouse studies. In one field experiment, prior herbivory by the bean leaf beetle subsequently reduced the suitability of foliage to the corn earwormHelicoverpa zea. The contribution of these findings to emerging theories of insect-plant interactions is discussed.