Induced resistance to Mexican bean beetles in soybean: variation among genotypes and lack of correlation with constitutive resistance

Fourteen genotypes (varieties) of soybean (Glycine max) were screened for levels of induced resistance to Mexican bean beetle (Epilachna varivestis) damage, and a subset of 6 of those varieties was screened for levels of constitutive resistance to Mexican bean beetles. Experiments were carried out in the greenhouse, with damage imposed by Mexican bean beetle larvae, and levels of resistance measured by a choice test bioassay with adult beetles. We found significant variation among soybean genotypes in levels of both induced and constitutive resistance. We found no significant correlation between levels of induced and constitutive resistance measured in the same genotypes. We compare these results to past work on resistance in the soybean-Mexican bean beetle system, consider the implications of variation in both types of resistance for plant-herbivore interactions in agricultural and natural systems, and discuss the relationship between induced and constitutive resistance. Received: 30 November 1998 / Accepted: 25 June 1999     

Effects of grazing on above-ground biomass on a mountain snowbed, NW Finland

The density-dependent effect of induced plant resistance on herbivore populations depends on the relationship between the amount of herbivore damage and the level of induced resistance produced by the plant. This relationship should influence the interaction of induced resistance and herbivore population dynamics, and if the relationship varies among plant genotypes, it could be subject to natural selection by herbivores. In this study the relationship between percent leaf area damaged and level of induced resistance was characterized for four genotypes of soybeans grown in a greenhouse. Damage ranging from 8 to 92% of leaf area was imposed using Mexican bean beetle larvae, and induced resistance was measured by bioassay using Mexican bean beetle adults. The level of induced resistance was significantly affected by the amount of damage, and the level of induced resistance varied significantly among the four genotypes. There was also a marginally significant interaction of damage and plant genotype, suggesting that the form of density dependence varies among these four genotypes of soybeans. These results suggest that these genotypes of plants might affect herbivore populations differently. If this variation is heritable, the form of density-dependent effects of induced resistance has the potential to evolve in this system.     

The timing of induced resistance and induced susceptibility in the soybean-Mexican bean beetle system

Induced plant responses to herbivory have been demonstrated in many systems. It has been suggested that the timing of these responses may influence the impact of induced resistance on herbivore populations, and may affect the evolution of induced defenses. This study used a bioassay to characterize the time course of systemic induced responses to Mexican bean beetle herbivory in four genotypes of soybeans. The results suggest that the time course of induced responses in this system is more complex than most previous studies have indicated. Herbivory provoked both rapid induced resistance and subsequent induced susceptibility to beetle feeding. All four genotypes of soybean induced significant resistance to beetle damage (beetles preferred undamaged to damaged plants) by 3 days after damage. By 15 days after damage, this resistance had decayed (beetles showed no preference for undamaged over damaged plants), and by 20 days after damage, all four genotypes exhibited significant induced susceptibility (beetles preferred previously damaged plants over undamaged plants). The magnitude of induced resistance in each genotype correlated strongly with the magnitude of induced susceptibility in that genotype. Received: 28 September 1997 / Accepted: 1 December 1997     

Comparing the consequences of induced and constitutive plant resistance for herbivore population dynamics

Although it has been suggested that induced and constitutive plant resistance should have different effects on insect herbivore population dynamics, there is little experimental evidence that plant resistance can influence herbivore populations longer than one season. We used a density-manipulation experiment and model fitting to examine the effects of constitutive and induced resistance on herbivore dynamics over both the short and long term. We used likelihood methods to fit population dynamic models to recruitment data for populations of Mexican bean beetles on soybean varieties with no resistance, constitutive resistance, or induced resistance. We compared model configurations that fit parameters for resistance types separately to models that did not account for resistance type. Models representing the hypothesis that the three resistance types differed in their effects on beetle dynamics received the most support. Induced resistance resulted in lower population growth rates and stronger density dependence than no resistance. Constitutive resistance resulted in lower population growth rates and stronger density dependence than induced resistance. Constitutive resistance had a stronger effect on both short-term beetle recruitment and predicted beetle population dynamics than induced resistance. The results of this study suggest that induced and constitutive resistance can differ in their effects on herbivore populations even in a relatively complex system.     

Organellar and Cytosolic Localization of Four Phosphoribosyl Diphosphate Synthase Isozymes in Spinach

We report on the efficacy of proteinase inhibitors (PIs) from three host plants (chickpea [Cicer arietinum], pigeonpea [Cajanus cajan], and cotton [Gossypium arboreum]) and three non-host (groundnut [Arachis hypogea], winged bean [Psophocarpus tetragonolobus], and potato [Solanum tuberosum]) in retarding the growth of Helicoverpa armigera larvae, a devastating pest of important crop plants. Enzyme assays and electrophoretic analysis of interaction of H. armigera gut proteinases (HGPs) with PIs revealed that non-host PIs inhibited HGP activity efficiently whereas host PIs were ineffective. In the electrophoretic assay, trypsin inhibitor activity bands were detected in all of the host and non-host plants, but HGP inhibitor activity bands were present only in non-host plants (except cotton in the host plant group). H. armigera larvae reared on a diet containing non-host PIs showed growth retardation, a reduction in total and trypsin-like proteinase activity, and the production of inhibitor-insensitive proteinases. Electrophoretic analysis of PI-induced HGP showed differential regulation of proteinase isoforms. Interestingly, HGP activity induced in response to dietary potato PI-II was inhibited by winged bean PIs. The optimized combination of potato PI-II and winged bean PIs identified in the present study and their proposed successive use has potential in developing H. armigera-resistant transgenic plants.     

Gene expression patterns of Helicoverpa armigera gut proteases

Relative quantification of reported gut proteinase cDNAs from Helicoverpa armigera larvae fed on various host plants (cotton, chickpea, pigeonpea, tomato and okra), non-host plant PIs (winged bean, bitter gourd, ground nut, and capsicum) and during larval development has been carried out using semi-quantitative RT-PCR. Five trypsin-like and three chymotrypsin-like proteinases were categorized as insensitive or sensitive to most of the proteinase inhibitors (PIs) and insensitive/sensitive to specific PIs based on their expression analysis. These results were supported by amino acid sequence analysis, where diverged amino acids were observed in the regions, which are reported to be involved in typical trypsin-trypsin inhibitor interactions and critical for proteinase inhibitor resistance. Among exopeptidases (five aminopeptidase and three carboxypeptidase), HaAmi4 and HaAmi5 of aminopeptidase and HaCar1 of carboxypeptidase exhibited considerable differential expression. Elastase and cathepsin B-like proteinases were expressed at negligible levels. The proteases identified in the study would be ideal candidates for further interactions studies with protease inhibitors to understand the structural reasons of protease inhibitor insensitivity.     

A fungal root symbiont modifies plant resistance to an insect herbivore

Vesicular-arbuscular mycorrhizal (VAM) fungi are common root-colonizing symbionts that affect nutrient uptake by plants and can alter plant susceptibility to herbivores. I conducted a factorial experiment to test the hypotheses that colonization by VAM fungi (1) improves soybean (Glycine max) tolerance to grazing by folivorous Mexican bean beetle (Epilachna varivestis), and (2) indirectly affects herbivores by increasing host resistance. Soybean seedlings were inoculated with the VAM fungus Glomus etunicatum or VAM-free filtrate and fertilized with high-[P] or low-[P] fertilizer. After plants had grown for 7 weeks first-instar beetle larvae were placed on bagged leaves. Growth of soybean was little affected by grazing larvae, and no effects of treatments on tolerance of soybeans to herbivores were evident. Colonization by VAM fungus doubled the size of phosphorus-stressed plants but these plants were still half the size of plants given adequate phosphorus. High-[P] fertilizer increased levels of phosphorus and soluble carbohydrates, and decreased levels of soluble proteins in leaves of grazed plants. Colonization of grazed plants by VAM fungus had no significant effect on plant soluble carbohydrates, but increased concentration of phosphorus and decreased levels of proteins in phosphorus-stressed plants to concentrations similar to those of plants given adequate phosphorus. Mexican bean beetle mass at pupation, pupation rate, and survival to eclosion were greatest for beetles reared on phosphorus-stressed, VAM-colonized plants, refuting the hypothesis that VAM colonization improves host plant resistance. VAM colonization indirectly affected performance of Mexician bean beetle larvae by improving growth and nutrition of the host plant. Received: 28 February 1997 / Accepted: 23 June 1997     

Transformation of white poplar (Populus alba L.) with a novel Arabidopsis thaliana cysteine proteinase inhibitor and analysis of insect pest resistance

Transgenic white poplar (Populus alba L.) plants expressing a novel Arabidopsis thaliana cysteine proteinase inhibitor (Atcys) gene have been produced using Agrobacterium tumefaciens-mediated gene transfer. Internodal stem segments of cv. Villafranca were co-cultivated with the EHA105 pBI-Atcys A. tumefaciens strain. Sixteen putative transgenic plant lines were regenerated from different calli with a transformation efficiency of 11%. The integration and expression of the cysteine proteinase inhibitor (Atcys) gene into the plant genome was confirmed by Southern and northern blot analyses. Papain inhibitory activity was detected in poplar transgenic tissues by means of a specific in vitro assay. Such activity was sufficient to inhibit most of the digestive proteinase activity of chrysomelid beetle (Chrysomela populi L.) and confer resistance to C. populi larvae on selected transgenic plants. A close correspondence between the inhibition of papain and resistance to poplar leaf beetle was observed in all tested transgenic lines. Our results indicate that Atcys could be succesfully employed in breeding programmes aimed at the selection of new poplar genotypes resistant to major insect pests.     

Herbivory,induced resistance,and interplant signal transfer in Alnus glutinosa

Field experiments with manually defoliated black alders (Alnus glutinosa) showed that defoliation affected herbivory by the major alder antagonist, the leaf beetle Agelastica alni. Herbivore damage increased with increasing distance to the defoliated tree, suggesting induced resistance not only on the damaged tree, but also on the neighbouring trees. The beetles also avoided leaves from the nearest neighbours for both feeding and oviposition in a laboratory assay, so the alders showed interplant resistance transfer. Natural enemies did not appear to shape this pattern, because the number of entomophagous arthropods and predator–prey ratios even increased with increasing distance to the defoliated tree. The numbers of all specialist, but not the generalist, herbivore species paralleled the increase in the attack of the specialist A. alni, supporting the view that specialists are more affected by plant resistance than generalists.Mechanisms causing this pattern, found in the field, were studied in more detail using biochemical analyses and further bioassays. Responses of alder leaves to herbivory of A. alni were shown to include ethylene emission and the release of a blend of volatiles with mono-, sesqui- and homoterpenes. Changes in leaf chemistry after herbivory included increases in the activity of oxidative enzymes (polyphenoloxidase, PPO, lipoxygenase, LOX, and peroxidase, POD) and proteinase inhibitors (PIs), and an increase in the phenolic contents of the leaves. Quantification of the endogenous jasmonic acid (JA) showed the activation of the octadecanoid pathway following herbivory.The active components in mediating a possible interplant signal transfer via airborne volatiles may have included ethylene, β-ocimene, 4,8-dimethylnona-1,3,7-triene (DMNT), and 4,8,12-trimethyltrideca-1,3,7,11-tetraene (TMTT). The incubation with volatiles resulted in an increase in the activity of catalase (CAT) and PIs (after MeJA application) and in an increase in the content of phenolics and PI activity (after ethylene application). Further evidence that airborne interplant communication may be important in the response of alder trees to beetle attack came from container experiments. In airtight chambers, unattacked leaves significantly increased the activity of proteinase inhibitors when they were associated with leaves previously attacked by beetle larvae.In conclusion, field experiments, bioassays in the laboratory as well as biochemical analyses suggest the existence of interplant resistance transfer in A. glutinosa, with airborne volatiles as a possible mechanism. However, the relative importance of airborne and possible soil-borne signals as well as unknown effects of intensified nutrient absorption of defoliated trees, possibly reducing foliage quality of undamaged neighbours, remains to be shown.     

Resistance of sweetpotato genotypes to adult Diabrotica beetles

Production of sweetpotatoes, Ipomoea batatas (L.) Lam. (Convolvulaceae), is limited by several insect pests, including Diabrotica spp. (Coleoptera: Chrysomelidae), and new integrated pest management (IPM) techniques for this crop are needed. Host plant resistance is one attractive approach that fits well into IPM programs. A host plant resistance research program typically depends on reliable bioassay procedures to streamline evaluation of germplasm. Thus, a bioassay technique was developed for evaluating sweetpotato germplasm by using adults of the banded cucumber beetle, Diabrotica balteata LeConte, and spotted cucumber beetle, Diabrotica undecimpunctata howardi Barber. A single beetle was placed on a piece of sweetpotato peel (periderm and cortex with stele removed) that was embedded periderm-side up in plaster in a petri dish. Feeding and longevity of insects on 30 sweetpotato genotypes were evaluated in two experiments by using this procedure. Adult longevity ranged from 7 to 11 d for starved individuals to 211 d for beetles fed a dry artificial diet. Longevity of banded cucumber beetles that fed on sweetpotato peels ranged from 12 d for the most-resistant genotype to 123 d for SC1149-19, a susceptible control cultivar. Longevity of spotted cucumber beetles was slightly shorter than longevity of banded cucumber beetles. For the most resistant sweetpotato genotypes, both Diabrotica species exhibited a significant delay in initiation of feeding, and more beetles died on these genotypes before they had fed. Both antibiosis and nonpreference (antixenosis) are important mechanisms of resistance in sweetpotato genotypes. This bioassay was consistent with field results, indicating that this technique could be useful for evaluating resistance to Diabrotica spp. in sweetpotato genotypes.     

Chemical defense lowers plant competitiveness

Both plant competition and plant defense affect biodiversity and food web dynamics and are central themes in ecology research. The evolutionary pressures determining plant allocation toward defense or competition are not well understood. According to the growth–differentiation balance hypothesis (GDB), the relative importance of herbivory and competition have led to the evolution of plant allocation patterns, with herbivore pressure leading to increased differentiated tissues (defensive traits), and competition pressure leading to resource investment towards cellular division and elongation (growth-related traits). Here, we tested the GDB hypothesis by assessing the competitive response of lima bean (Phaseolus lunatus) plants with quantitatively different levels of cyanogenesis—a constitutive direct, nitrogen-based defense against herbivores. We used high (HC) and low cyanogenic (LC) genotypes in different competition treatments (intra-genotypic, inter-genotypic, interspecific), and in the presence or absence of insect herbivores (Mexican bean beetle, Epilachna varivestis) to quantify vegetative and generative plant parameters (above and belowground biomass as well as seed production). Highly defended HC-plants had significantly lower aboveground biomass and seed production than LC-plants when grown in the absence of herbivores implying significant intrinsic costs of plant cyanogenesis. However, the reduced performance of HC- compared to LC-plants was mitigated in the presence of herbivores. The two plant genotypes exhibited fundamentally different responses to various stresses (competition, herbivory). Our study supports the GDB hypothesis by demonstrating that competition and herbivory affect different plant genotypes differentially and contributes to understanding the causes of variation in defense within a single plant species.     

Negative interactions between chemical resistance and predators affect fitness in soybeans

Abstract 1. Plants may benefit from both chemical resistance traits and the presence of predators of herbivores. In past studies, the interaction between resistance and predators varies from complementary to antagonistic among different systems. However, this interaction has primarily been quantified by effects on predator abundance or vigor, not effects on plant fitness. 2. In this study, the combined effects of chemical resistance and predators on plant fitness were examined using soybeans (Glycine max), herbivorous Mexican bean beetles (Epilachna varivestis), and predaceous spined soldier bugs (Podisus maculiventris). Mexican bean beetles were reared in field cages in the presence or absence of spined soldier bugs on soybeans with or without strong constitutive chemical resistance. 3. Spined soldier bugs were more likely to feed on Mexican bean beetles that fed on susceptible than on resistant plants. 4. Susceptible plants with predators produced significantly more seeds than those without predators, while resistant plants did not produce significantly different numbers of seeds based on the presence or absence of predators. 5. Selection for the production of some types of chemical resistance in plants would thus be expected to be stronger with lower predation rates. 6. These results also suggest predator introductions would be more effective on plants without a strong constitutive chemical resistance to herbivores.     

Compensatory proteolytic responses to dietary proteinase inhibitors from Albizia lebbeck seeds in the Helicoverpa armigera larvae

Plant proteinase inhibitors (PIs) have been shown to reduce the growth rates in larvae of numerous insect species. On the other hand, insects can also regulate their proteinases against plant PIs. In the present study, we report the compensatory activities of Helicoverpa armigera (Hubner) (Lepidoptera: Noctuidae) gut proteinases against the PIs of Albizia lebbeck seeds. Total of ten proteinase inhibitor bands were detected in the seed extract of A. lebbeck. Bioassays were conducted by feeding H. armigera larvae on diet containing partially purified PIs from A. lebbeck seeds. Results show that larval growth and survival was significantly reduced by A. lebbeck PIs. We found that higher activity H. armigera gut proteinase (HGP) isoforms observed in the midgut of control larvae were inhibited in the midgut of larvae fed on test diet. Some HGP isoforms were induced in the larvae fed on PI containing test diet; however, these isoforms showed lower activity in the larvae fed on control diet. Aminopeptidase activities were significantly increased in the midgut of larvae fed on test diet. A population of susceptible and resistant enzymes was observed in the midgut of H. armigera, when fed on diet containing PIs from A. lebbeck seeds. Our initial observations indicate that H. armigera can regulate its digestive proteinase activity against non-host plant PIs, too. It is important to study the exact biochemical and molecular mechanisms underlying this phenomenon in order to develop PI-based insect control strategies.     

Salicylic acid inhibits jasmonic acid-induced resistance of Arabidopsis thaliana to Spodoptera exigua

The role of salicylic acid (SA) in plant responses to pathogens has been well documented, but its direct and indirect effects on plant responses to insects are not so well understood. We examined the effects of SA, alone and in combination with jasmonic acid (JA), on the performance of the generalist herbivore, Spodoptera exigua, in wild-type and mutant Arabidopsis thaliana genotypes that varied genetically in their ability to mount SA- and JA-mediated defence responses. In one experiment, growth of S. exigua larvae was highest on the Wassilewskija wild-type, intermediate on the Columbia wild-type and the JA-deficient fad mutant, and lowest on the nim1-1 and jar1-mutants, which are defective in the SA and JA pathways, respectively. Activity of guaiacol peroxidase, polyphenoloxidase, n-acetylglucosaminidase, and trypsin inhibitor varied by genotype but did not correlate with insect performance. SA treatment increased growth of S. exigua larvae by approximately 35% over all genotypes, but had no discernable effect on activities of the four defence proteins. In a second experiment, growth of S. exigua was highest across treatments on the cep1 mutant, a constitutive expressor of high SA levels and systemic acquired resistance, and lowest on the fad mutant, which is JA-deficient. JA treatment generally increased activity of all four defence proteins, increased total glucosinolate levels and reduced insect growth by approximately 25% over all genotypes. SA generally inhibited expression of JA-induced resistance to S. exigua when both hormones were applied simultaneously. Across genotypes and treatments, larval mass was negatively correlated with the activity of trypsin inhibitor and polyphenoloxidase and with total glucosinolate levels, and insect damage was negatively correlated with the activity of polyphenoloxidase. SA had little effect on the induction of defence protein activity by JA. However, SA attenuated the induction of glucosinolates by JA and therefore may explain better the interactive effects of SA and JA on insect performance. This study illustrates that direct and indirect cross-effects of SA on resistance to S. exigua can occur in A. thaliana. Effects of SA may be mediated through effects on plant defence chemistry or other aspects of the suitability of foliage for insect feeding and growth.     

Microarray analysis of the interaction between the aphid <Emphasis Type="Italic">Rhopalosiphum padi</Emphasis> and host plants reveals both differences and similarities between susceptible and partially resistant barley lines

The bird cherry-oat aphid (Rhopalosiphum padi L.) is an important pest on cereals causing plant growth reduction without specific leaf symptoms. Breeding of barley (Hordeum vulgare L.) for R. padi resistance shows that there are several resistance genes, reducing aphid growth. To identify candidate sequences for resistance-related genes, we performed microarray analysis of gene expression after aphid infestation in two susceptible and two partially resistant barley genotypes. One of the four lines is a descendant of two of the other genotypes. There were large differences in gene induction between the four lines, indicating substantial variation in response even between closely related genotypes. Genes induced in aphid-infested tissue were mainly related to defence, primary metabolism and signalling. Only 24 genes were induced in all lines, none of them related to oxidative stress or secondary metabolism. Few genes were down-regulated, with none being common to all four lines. There were differences in aphid-induced gene regulation between resistant and susceptible lines. Results from control plants without aphids also revealed differences in constitutive gene expression between the two types of lines. Candidate sequences for induced and constitutive resistance factors have been identified, among them a proteinase inhibitor, a serine/threonine kinase and several thionins. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Characterization of resistance to the bean weevil <Emphasis Type="Italic">Acanthoscelides obtectus</Emphasis> Say, 1831 (Coleoptera: Bruchidae) in common bean genotypes

The bean weevil Acanthoscelides obtectus (Say, 1831) (Coleoptera: Bruchidae) is one of the most serious pests of stored beans worldwide because of the damage it causes to grains within warehouses. The use of resistant genotypes may offer a control strategy for this pest. In the current study, we screened common bean genotypes of Andean American and Mesoamerican origin in laboratory and greenhouse bioassays to select the most promising beans for resistance to the bean weevil. In the laboratory, we evaluated number of eggs, period of development (egg-adult), number of emerged adults, dry weight of adults, and weight of consumed grains. In the greenhouse, number of pods per plant and number of grains per pod were evaluated. We also assessed the percentages of damaged pods per plant and damaged grains per pod. Combining the results obtained in the laboratory and greenhouse assays, the common bean genotypes Arc.1, Arc.2, Arc.1S, Arc.5S, and Arc.3S were identified as resistance expressing antibiosis against A. obtectus. The lowest percentages of damaged pods were found in the Arc.1 and Arc.1S genotypes, and their resistance to damage was apparently morphological (antixenotic) because they possessed structures that prevented contact between larvae and grains. The use of resistant genotypes in combination with other techniques may improve management of the weevil. Additionally, the resistant genotypes identified here can be used in breeding programs to develop common bean lines with resistance to A. obtectus.     

Insect pest damage increases faba bean (Vicia faba) yield components but only in the absence of insect pollination

Identifying and quantifying crop stressors interactions in agroecosystems is necessary to guide sustainable crop management strategies. Over the last 50 years, faba bean cropping area has been declining, partly due to yield instabilities associated with uneven insect pollination and herbivory. Yet, the effect of interactions between pollinators and a key pest, the broad bean beetle Bruchus rufimanus (florivorous and seed predating herbivore) on faba bean yield has not been investigated. Using a factorial cage experiment in the field, we investigated how interactions between two hypothesized stressors, lack of insect pollination by bumblebees and herbivory by the broad bean beetle, affect faba bean yield. Lack of bumblebee pollination reduced bean weight per plant by 15%. Effects of the broad bean beetle differed between the individual plant and the plant‐stand level (i.e., when averaging individual plant level responses at the cage level), likely due to high variation in the level of herbivory among individual plants. At the individual plant level, herbivory increased several yield components but only in the absence of pollinators, possibly due to plant overcompensation and/or pollination by the broad bean beetle. At the plant‐stand level, we found no effect of the broad bean beetle on yield. However, there was a tendency for heavier individual bean weight with bumblebee pollination, but only in the absence of broad bean beetle herbivory, possibly due to a negative effect of the broad bean beetle on the proportion of legitimate flower visits by bumblebees. This is the first experimental evidence of interactive effects between bumblebees and the broad bean beetle on faba bean yield. Our preliminary findings of negative and indirect associations between the broad bean beetle and individual bean weight call for a better acknowledgment of these interactions in the field in order to understand drivers of crop yield variability in faba bean.     

Tolerance as a mechanism of resistance to Thrips palmi in common beans

Tolerance as a mechanism of resistance to the melon thrips, Thrips palmi Karny (Thysanoptera: Thripidae), in common beans, Phaseolus vulgaris L., was evaluated under field and greenhouse conditions. Seven resistant (Brunca, BH‐5, BH‐60, BH‐130, BH‐144, EMP 486, and FEB 115) and five susceptible (PVA 773, EMP 514, BAT 477, APN 18, and RAZ 136) bean genotypes were assessed according to adult and larval populations, visual damage and reproductive adaptation scores, and yield components in field trials. From these genotypes, four resistant (Brunca, BH‐130, EMP 486, and FEB 115) and two susceptible (APN 18 and RAZ 136) genotypes were selected for quantification of proportional plant weight and height increase changes due to thrips infestation in greenhouse tests. Under medium to high thrips infestation in the field, most resistant genotypes tended to have higher reproductive adaptation and lower yield losses, though they did not always suffer less damage, as compared to susceptible genotypes. In the greenhouse, resistant genotypes showed less reduction in plant dry weight and height increase than did some susceptible ones under the same infestation pressure. Results from both field trials and greenhouse tests suggest the possible expression of tolerance as a mechanism of resistance to T. palmi in the resistant genotype EMP 486, and confirm the existence of antixenosis in FEB 115, whereas tolerance might be combined with other resistance mechanisms in Brunca.     

Herbivore performance and plant defense after sequential attacks by inducing and suppressing herbivores

It is well known that herbivore-induced plant defenses alter host plant quality and can affect the behavior and performance of later arriving herbivores.Effects of sequential attacks by herbivores that either suppress or induce plant defenses are less well studied.We sequentially infested leaves of tomato plants with a strain of the phytophagous spider mite Tetranychus urticae that induces plant defenses and the closely related Tetranychus evansi, which suppresses plant defenses.Plant quality was quantified through oviposifion of both spider mite species and by measuring proteinase inhibitor activity using plant material that had been sequentially attacked by both herbivore species.Spider-mite oviposifion data show that T.evansi could suppress an earlier induction of plant defenses by T.urticae,and T.urticae could induce defenses in plants previously attacked by T.evansi in 1 day.Longer attacks by the second species did not result in further changes in oviposifion.Proteinase inhibitor activity levels showed that T.evansi suppressed the high activity levels induced by T.urticae to constitutive levels in 1 day,and further suppressed activity to levels similar to those in plants attacked by T.evansi alone.Attacks by T.urticae induced proteinase inhibitor activity in plants previously attacked by T.evansi,eventually to similar levels as induced by T.urticae alone.Hence,plant quality and plant defenses were significantly affected by sequential attacks and the order of attack does not affect subsequent performance,but does affect proteinase inhibitor activity levels.Based on our results,we discuss the evolution of suppression of plant defenses.     

Plant genotype and induced responses affect resistance to herbivores on evening primrose (Oenothera biennis)

Abstract. 1. Although both genotype and induced responses affect a plant's resistance to herbivores, little is known about their relative and interactive effects. This study examined how plant genotype of a native plant (Oenothera biennis) and induced plant responses to herbivory affect resistance to, and interactions among, several herbivores. 2. In a field experiment, genetic and environmental variation among habitats led to variation in the amount of early season damage and plant quality. The pattern of variation in early season infestation by spittlebugs (Philaenus spumarius, a piercing–sucking herbivore) negatively correlated with oviposition preference by a later feeding specialist weevil (Tyloderma foveolatum, a leaf‐chewer). 3. To determine if plant genotype and induced responses to herbivory might be responsible for these field patterns, we performed no‐choice and choice bioassays using four genotypes of O. biennis that varied in resistance. Plants were induced by either spittlebugs or weevils and assays measured the responses of the same specialist weevil as well as a generalist caterpillar (Spodoptera exigua). 4. Resistance to adult weevils was largely unaffected by plant genotype, while they experienced induced resistance following damage by conspecific weevils in no‐choice assays. Caterpillars were more strongly affected by plant genotype than induced responses in both no‐choice and choice assays, but they also fed less and experienced higher mortality on plants previously damaged by weevils. In contrast to the pattern suggested by the field experiment, spittlebugs did consistently induce resistance against either weevils or caterpillars in the bioassay experiment. 5. These results support recent findings that show herbivore species can compete via induced plant responses. Additionally, a quantitative review of the literature demonstrates that plant genotype tends to be more important than interspecific competition among herbivores (plant‐mediated or otherwise) in affecting herbivore preference and performance.