Disease Interactions in a Shared Host Plant: Effects of Pre-Existing Viral Infection on Cucurbit Plant Defense Responses and Resistance to Bacterial Wilt Disease

Both biotic and abiotic stressors can elicit broad-spectrum plant resistance against subsequent pathogen challenges. However, we currently have little understanding of how such effects influence broader aspects of disease ecology and epidemiology in natural environments where plants interact with multiple antagonists simultaneously. In previous work, we have shown that healthy wild gourd plants (Cucurbita pepo ssp. texana) contract a fatal bacterial wilt infection (caused by Erwinia tracheiphila) at significantly higher rates than plants infected with Zucchini yellow mosaic virus (ZYMV). We recently reported evidence that this pattern is explained, at least in part, by reduced visitation of ZYMV-infected plants by the cucumber beetle vectors of E. tracheiphila. Here we examine whether ZYMV-infection may also directly elicit plant resistance to subsequent E. tracheiphila infection. In laboratory studies, we assayed the induction of key phytohormones (SA and JA) in single and mixed infections of these pathogens, as well as in response to the feeding of A. vittatum cucumber beetles on healthy and infected plants. We also tracked the incidence and progression of wilt disease symptoms in plants with prior ZYMV infections. Our results indicate that ZYMV-infection slightly delays the progression of wilt symptoms, but does not significantly reduce E. tracheiphila infection success. This observation supports the hypothesis that reduced rates of wilt disease in ZYMV-infected plants reflect reduced visitation by beetle vectors. We also documented consistently strong SA responses to ZYMV infection, but limited responses to E. tracheiphila in the absence of ZYMV, suggesting that the latter pathogen may effectively evade or suppress plant defenses, although we observed no evidence of antagonistic cross-talk between SA and JA signaling pathways. We did, however, document effects of E. tracheiphila on induced responses to herbivory that may influence host-plant quality for (and hence pathogen acquisition by) cucumber beetles.     

Resistance and tolerance to herbivory in Salix cordata are affected by different environmental factors

Abstract.  1. Effects of sand burial and nutrients on the ability of sand-dune willow ( Salix cordata ) to tolerate or resist herbivory by the beetle Altica subplicata were evaluated in field experiments.
2. To assess tolerance, all combinations of sand burial (none, 50%), nutrients (presence, absence), and beetles (presence, absence) were applied to caged plants and growth responses to herbivory were measured. Sand burial increased plant growth rate, but decreased S. cordata 's tolerance to herbivory. Although nutrients increased growth, tolerance to herbivory was unaffected.
3. To assess resistance, plants were exposed to all combinations of sand burial and nutrients, and then to natural beetle colonisation. The presence of nutrients, but not sand burial, significantly increased the percentage of plants with beetles, for both adults and larvae. This decreased resistance to beetles of plants grown with added nutrients occurred only in the absence of sand burial.
4. The performance and preference of beetles were examined in laboratory experiments. Larvae developed faster and had increased pupation success on plants with nutrients added. Beetles also showed a marginally significant feeding preference for leaves grown with added nutrients. Thus, S. cordata tolerance to herbivory was affected by sand burial, whereas resistance, preference, and performance were affected by nutrient level.     

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     

Heterogeneity of plant phenotypes caused by herbivore-specific induced responses influences the spatial distribution of herbivores

Abstract.  1. Herbivory can induce resistance in a plant and the induced phenotype may be disfavoured by subsequent herbivores. Yet, as the distance between plants in a population increases, limited mobility may make a herbivore more likely to feed and oviposit on host plants in its immediate surroundings.
2. The present study tested whether a herbivore's preference and distribution across plants with different induced phenotypes was influenced by the spatial distribution of plants. A fragmented population of Solanum dulcamara plants was created. This consisted of discrete, spatially separated patches with different histories of damage, either herbivory from adult flea beetles ( Psylliodes affinis ), tortoise beetles ( Plagiometriona clavata ), or mechanical damage. Each patch was separated by 7 m and consisted of 12 plants that were spaced 30 cm apart. Then a fixed number of adult tortoise beetles were introduced to each patch, and movement and oviposition within and between spatially separate homogeneous patches (receiving one type of damage) were compared with movement and oviposition within heterogeneous patches (containing all three types of damage) over the growing season.
3. Flea beetle and tortoise beetle herbivory consistently induced different phytochemical responses in S. dulcamara (polyphenol oxidase and peroxidase), and adult tortoise beetles avoided oviposition on the flea beetle induced plants within heterogeneous patches. However, between homogeneous patches, plant phenotype did not influence oviposition. Colonisation by naturally occurring flea beetle adults followed a similar pattern.
4. These results suggest that the heterogeneity of plant phenotypes can influence herbivore choice and distribution at small but not large spatial scales.     

Insect feeding on cucumber mediated by rhizobacteria-induced plant resistance

Select strains of plant growth-promoting rhizobacteria (PGPR) were evaluated in greenhouse experiments with cucumber for induction of resistance against cucumber beetle (Diabrotica undecimpunctata howardi Barber) feeding and the beetle-transmitted cucurbit wilt disease. When beetles were given a choice between PGPR-treated and nontreated cucumber, their feeding on stems and cotyledons and the severity of wilt symptoms were significantly lower on PGPR-treated plants. HPLC analysis demonstrated that cotyledons from PGPR-treated plants contained significantly lower concentrations of the cucumber beetle feeding stimulant cucurbitacin than nontreated plants. These results suggest that a mechanism for PGPR-induced resistance against cucumber beetle feeding may involve a change in the metabolic pathway for cucurbitacin synthesis.     

Latitudinal variation in resistance and tolerance to herbivory of a salt marsh shrub

Interactions between plants and herbivores often vary on a geographic scale. Although theory about plant defenses and tolerance is predicated on temporal or spatial variation in herbivore damage, no single study has compared the pattern of herbivory, plant defenses and tolerance to herbivory of a single species across a latitudinal gradient. In 2002–2005 we surveyed replicate salt marshes along the Atlantic coast of the United States from Florida to Maine. At each field site we scored leaves of Iva frutescens for herbivore damage. In laboratory experiments we measured constitutive resistance and induced resistance in I. frutescens from high and low latitude sites along the Atlantic Coast. In another common garden experiment we studied tolerance to herbivory of I. frutescens from various sites. Theory predicts that constitutive resistance should matter more when damage is high, and induced resistance when herbivory is high but variable. In the field, average levels of herbivore damage, and spatial and temporal variation in herbivore damage were all greater at low versus high latitudes, indicating that constitutive as well as induced resistance should be stronger at low latitudes. Consistent with this prediction, constitutive resistance to herbivory was stronger at low latitudes. Induced resistance to herbivores was also stronger at low latitudes: it was deployed faster and lasted longer. Theory also predicts that tolerance to herbivory should be greater where average herbivory damage is greater; however, tolerance to herbivory in Iva did not depend on geographic origin. Our results emphasize the value of considering multiple ways in which plants respond to herbivores when examining geographic variation in plant–herbivore interactions.     

Differences in growth and herbivory damage of native and invasive kudzu (Peuraria montana var. lobata) populations grown in the native range

The invasion success of exotic plants is often attributed to escape from natural enemies in their introduced ranges and subsequent evolutionary change in resource allocation from defense to growth and reproduction. We tested this idea by comparing resistance, tolerance, and growth between native (China) and invasive (US) populations of kudzu (Peuraria montana var. lobata) exposed to natural herbivores in the native range. The percentage of foliar damage was much higher in invasive populations than in native populations, indicating that plants from invasive populations had lower resistance to herbivory. Regression of total mass on percentage of foliar damage showed no significant differences in tolerance to herbivory between native and invasive populations. However, stem diameter and mass were significantly greater in invasive populations than in native populations. Our results may suggest geographic variation in herbivory damage and plant growth among kudzu native and invasive populations, but the role of herbivores influencing kudzu invasion requires further investigation.     

Influence of leaf herbivory,root herbivory,and pollination on plant performance in Cucurbita moschata

Abstract 1. Plants experience herbivory on many different tissues that can affect reproduction directly by damaging tissues and decreasing resource availability, or indirectly via interactions with other species such as pollinators. 2. This study investigated the combined effects of leaf herbivory, root herbivory, and pollination on subsequent damage, pollinator preference, and plant performance in a field experiment using butternut squash (Cucurbita moschata). Leaf and root herbivory were manipulated using adult and larval striped cucumber beetles (Acalymma vittatum F.), a cucurbit specialist. 3. Leaf herbivory reduced subsequent pistillate floral damage and powdery mildew (Sphaerotheca fuliginea) infection. In spite of these induced defences, the overall effect of leaf herbivory on plant reproduction was negative. Leaf herbivory reduced staminate flower production, fruit number, and seed weight. In contrast, root herbivory had a minimal impact on plant reproduction. 4. Neither leaf nor root herbivory altered pollinator visitation or floral traits, suggesting that reductions in plant performance from herbivory were as a result of direct rather than indirect effects. In addition, no measured aspect of reproduction was pollen limited. 5. Our study reveals that although leaf herbivory by the striped cucumber beetle can protect against subsequent damage, this protection was not enough to prevent the negative impacts on plant performance.     

Specificity of induced plant responses to specialist herbivores of the common milkweed Asclepias syriaca

Induced plant responses to herbivory appear to be universal, yet the degree to which they are specific to sets of herbivores is poorly understood. The generalist/specialist hypothesis predicts that generalist herbivores are more often negatively affected by host plant defenses, wheras specialists may be either unaffected by or attracted to these same "plant defenses". Therefore, specialists should be less predictable than generalists in their responses to induced plant resistance traits. To better understand the variation in plant responses to herbivore attack, and the impacts these responses have on specialist herbivores, we conducted a series of experiments examining pairwise interactinos between two specialaist herbivores of the common milkweed ( Asclepias syriaca ). We damaged plants mechnically, with swamp milkweed beetles ( Labidomera clivicollis ), or with monarchs ( Danaus plexippus ), and then asessed specificity of elicitation, both by measuring a putative defensive trait (latex volume) and by challenging plants with insects of both species in bioasays. Latex production increased by 34% and 13% following beetle and monarch herbivory, respectively, but only beetles significantly elevated latex production compared to undamaged controls. While beetle growth was negatively affected by latex across all experiments, beetles were not affected by previous damage caused by conspecifies or by monarchs. In contrast, monarchs feeding on previously damaged plants were 20% smaller, and their response was the same on plants damaged mechnically or by either herbivore. Therefore, these specialist herbivores exhibit both specificity of elicitation in plant responses and specificity of effects in response to prior damage.     

Effects of plant diversity and time of colonization on an herbivore-plant interaction

Summary Experimental field plantings showed that plant diversity strongly affected the population dynamics of a specialist herbivore, the striped cucumber beetle, Acalymma vittata (Fab.) (Coleoptera: Chrysomelidae). Population densities over time were characterized by two peaks in numbers (from colonization and reproduction, respectively) and were consistently higher in cucumber monocultures (Cucumis sativus L.) than in polycultures of cucumbers, corn (Zea mays L.), and broccoli (Brassica oleracea L.). Greater abundances in monocultures appear to result from two factors: (1) per individual reproductive rates were greater in monocultures than in polycultures, and (2) mark-recapture studies confirmed that beetles stay in monocultures for a longer period of time than in polycultures. Differences in predation did not appear to contribute to the overall differences in herbivore abundances.The primary impact of A. vittata on its host plant, C. sativus, is the dissemination of bacterial wilt disease, Erwinia tracheiphila (E.F.Sm.). Greater numbers of beetles led to greater plant mortality in monocultures. It is suggested that factors other than numbers of beetles (e.g., shading, allelopathy, microclimate) are more important in influencing plant reproduction, since cucumber plants in monocultures had greater yields than did plants grown in polycultures. However, time of beetle colonization strongly affected plant parameters, indicating that the length of time during which herbivores are interacting with plants is of critical importance to plant survivorship, and thus reproduction.     

The evolution of plant response to herbivory: simultaneously considering resistance and tolerance in Brassica rapa

Although the evolution of plant response to herbivory can involve either resistance (a decrease in susceptibility to herbivore damage) or tolerance (a decrease in the per unit effect of herbivory on plant fitness), until recently few studies have explicitly incorporated both of these characters. Moreover, theory suggests these characters do not evolve independently, and also that the pattern of natural selection acting on resistance and tolerance depends on their costs and benefits. In a genotypic selection analysis on an experimental population of Brassica rapa (Brassicaceae) I found a complex set of correlational selection gradients acting on resistance and tolerance of damage by flea beetles (Phyllotreta cruciferae: Chrysomelidae) and weevils (Ceutorhynchus assimilis: Curculionidae), as well as directional and stabilizing selection on resistance to attack by weevils. Evolution of response to flea beetle attack is constrained by a strong allocation cost of tolerance, and this allocation cost may be caused by a complex correlation among weevil resistance, weevil tolerance, flea beetle resistance, and flea beetle tolerance. Thus, one important conclusion of this study is that ecological costs may involve complex correlations among multiple characters, and for this reason these costs may not be detectable by simple pairwise correlations between characters. The evolution of response to weevil attack is probably constrained by a series of correlations between weevil resistance, weevil tolerance, and fitness in the absence of weevil damage, and possibly by a cost of tolerance of weevil damage. However, the nature of these constraints is complicated by apparent overcompensation for weevil damage. Because damage by both flea beetles and weevils had non-linear effects on plant fitness, standard measures of tolerance were not appropriate. Thus, a second important contribution of this study is the use of the area under the curve defined by the regression of fitness on damage and damage-squared as a measure of tolerance. This revised version was published online in July 2006 with corrections to the Cover Date.     

TWO HERBIVORES AND CONSTRAINTS ON SELECTION FOR RESISTANCE IN BRASSICA RAPA

Although most plants experience herbivory by several insect species, there has been little empirical work directed toward understanding plant responses to these simultaneous selection pressures. In an experiment in which herbivory by flea beetles (Phyllotreta cruciferae) and diamondback moths (Plutella xylostella) was manipulated in a factorial design, I found that selection for resistance to these herbivores is not independent in Brassica rapa. Specifically, the effect of flea beetle damage on B. rapa fitness depends on the amount of diamondback moth damage a plant experiences: damage by these herbivores has a nonadditive effect on plant fitness. When diamondbacks are abundant, plants that sustain high levels of damage by flea beetles are favored by natural selection, but when diamondbacks are rare, a low level of damage by flea beetles is favored. However, resistance to the later-feeding diamondback moth is not affected by the presence or absence of damage by early-feeding flea beetles. Thus, there are no plant-mediated ecological interactions between these herbivores that affect the outcome of selection for resistance. Because these herbivores do not independently affect plant fitness, neither is likely to develop a pairwise coevolutionary relationship with its host. Instead, coevolution is diffuse.     

Tolerance to deer herbivory and resistance to insect herbivores in the common evening primrose (Oenothera biennis)

The evolution of plant defence in response to herbivory will depend on the fitness effects of damage, availability of genetic variation and potential ecological and genetic constraints on defence. Here, we examine the potential for evolution of tolerance to deer herbivory in Oenothera biennis while simultaneously considering resistance to natural insect herbivores. We examined (i) the effects of deer damage on fitness, (ii) the presence of genetic variation in tolerance and resistance, (iii) selection on tolerance, (iv) genetic correlations with resistance that could constrain evolution of tolerance and (v) plant traits that might predict defence. In a field experiment, we simulated deer damage occurring early and late in the season, recorded arthropod abundances, flowering phenology and measured growth rate and lifetime reproduction. Our study showed that deer herbivory has a negative effect on fitness, with effects being more pronounced for late‐season damage. Selection acted to increase tolerance to deer damage, yet there was low and nonsignificant genetic variation in this trait. In contrast, there was substantial genetic variation in resistance to insect herbivores. Resistance was genetically uncorrelated with tolerance, whereas positive genetic correlations in resistance to insect herbivores suggest there exists diffuse selection on resistance traits. In addition, growth rate and flowering time did not predict variation in tolerance, but flowering phenology was genetically correlated with resistance. Our results suggest that deer damage has the potential to exert selection because browsing reduces plant fitness, but limited standing genetic variation in tolerance is expected to constrain adaptive evolution in O. biennis.     

Benefits for plants in ant-plant protective mutualisms: a meta-analysis

Costs and benefits for partners in mutualistic interactions can vary greatly, but surprisingly little is known about the factors that drive this variation across systems. We conducted a meta-analysis of ant-plant protective mutualisms to quantify the effects of ant defenders on plant reproductive output, to evaluate if reproductive effects were predicted from reductions in herbivory and to identify characteristics of the plants, ants and environment that explained variation in ant protection. We also compared our approach with two other recent meta-analyses on ant-plant mutualisms, emphasizing differences in our methodology (using a weighted linear mixed effects model) and our focus on plant reproduction rather than herbivore damage. Based on 59 ant and plant species pairs, ant presence increased plant reproductive output by 49% and reduced herbivory by 62%. The effects on herbivory and reproduction within systems were positively correlated, but the slope of this relationship (0.75) indicated that tolerance to foliar herbivory may be a common plant response to absence of ant guards. Furthermore, the relationship between foliar damage and reproduction varied substantially among systems, suggesting that herbivore damage is not a reliable surrogate for fitness consequences of ant protection. Studies that experimentally excluded ants reported a smaller effect of ant protection on plant reproduction than studies that relied upon natural variation in ant presence, suggesting that study methods can affect results in these systems. Of the ecological variables included in our analysis, only plant life history (i.e., annual or perennial) explained variation in the protective benefit of mutualistic ants: presence of ants benefitted reproduction of perennials significantly more than that of annuals. These results contrast with other quantitative reviews of these relationships that did not include plant life history as an explanatory factor and raise several questions to guide future research on ant-plant protection mutualisms.     

Effects of above- and belowground herbivory on growth,pollination, and reproduction in cucumber

Plants experience unique challenges due to simultaneous life in two spheres, above- and belowground. Interactions with other organisms on one side of the soil surface may have impacts that extend across this boundary. Although our understanding of plant–herbivore interactions is derived largely from studies of leaf herbivory, belowground root herbivores may affect plant fitness directly or by altering interactions with other organisms, such as pollinators. In this study, we investigated the effects of leaf herbivory, root herbivory, and pollination on plant growth, subsequent leaf herbivory, flower production, pollinator attraction, and reproduction in cucumber (Cucumis sativus). We manipulated leaf and root herbivory with striped cucumber beetle (Acalymma vittatum) adults and larvae, respectively, and manipulated pollination with supplemental pollen. Both enhanced leaf and root herbivory reduced plant growth, and leaf herbivory reduced subsequent leaf damage. Plants with enhanced root herbivory produced 35% fewer female flowers, while leaf herbivory had no effect on flower production. While leaf herbivory reduced the time that honey bees spent probing flowers by 29%, probing times on root-damaged plants were over twice as long as those on control plants. Root herbivory increased pollen limitation for seed production in spite of increased honey bee preference for plants with root damage. Leaf damage and hand-pollination treatments had no effect on fruit production, but plants with enhanced root damage produced 38% fewer fruits that were 25% lighter than those on control plants. Despite the positive effect of belowground damage on honey bee visitation, root herbivory had a stronger negative effect on plant reproduction than leaf herbivory. These results demonstrate that the often-overlooked effects of belowground herbivores may have profound effects on plant performance.     

Ontogenetic changes in tolerance to herbivory in Arabidopsis

Tolerance to herbivory—the ability of plants to maintain fitness despite herbivore damage—is expected to change during the life cycle of plants because the physiological mechanisms underlying tolerance to herbivory are linked to growth, and resource allocation to growth changes throughout ontogeny. We used the model plant Arabidopsis thaliana to test two hypotheses: that tolerance increases as plants grow, and that tolerance decreases at the onset of reproduction. We chose three accessions previously reported to vary for resistance to herbivory in order to explore whether tolerance and resistance are inversely related. Cabbage looper (Trichoplusia ni) larvae were allowed to feed on plants at either the four-leaf, six-leaf, or 1st-flower developmental stage until 50% of the leaf area was removed. Overall, we found a trend for increased tolerance with ontogenetic stage, but there were important differences among accessions in their response to herbivory at different stages. Tolerance did not decrease with the onset of flowering, nor did we find any correlation between resistance and tolerance levels. Three main plant traits correlated strongly with tolerance: stem mass, an earlier onset of reproduction and a longer fruiting period. This study suggests there may be considerable variation in ontogenetic patterns of tolerance in natural populations of A. thaliana, and warrants further investigations with more accessions or natural populations, and detailed measurements of traits purported to contribute to tolerance in our quest to understand the mechanisms of tolerance to herbivory.     

Decreased resistance and increased tolerance to native herbivores of the invasive plant Sapium sebiferum

Release from natural enemies may favor invasive plants evolving traits associated with reduced herbivore‐resistance and faster‐growth in introduced ranges. Given a genetic trade‐off between resistance and tolerance, invasive plants could also become more tolerant to herbivory than conspecifics in the native range. We conducted a field common garden study in the native range of Sapium sebiferum using seeds from native Chinese populations and invasive North American populations to compare their growth and herbivory resistance. We also performed a cage‐pot experiment to compare their resistance and tolerance to Bikasha collaris beetles that are specialist feeders on S. sebiferum trees in China. Results of the common garden study showed that Sapium seedlings of invasive populations relative to native populations were more frequently attacked by native herbivores. Growth and leaf damage were significantly higher for invasive populations than for native populations. Growth of invasive populations was not significantly affected by insecticide spray, but insecticide spray benefited that of native populations. In the bioassay trial, beetles preferentially consumed leaf tissue of invasive populations compared to native populations when beetles had a choice between them. Regression of percent leaf damage on biomass showed that invasive populations tolerated herbivory more effectively than native populations. Our results suggest that S. sebiferum from the introduced range had lower resistance but higher tolerance to specialist herbivores. Both defense strategies could have evolved as a response to the escape from natural enemies in the introduced range.     

No evidence for evolutionarily decreased tolerance and increased fitness in invasive Chromolaena odorata: implications for invasiveness and biological control

Tolerance, the degree to which plant fitness is affected by herbivory, is associated with invasiveness and biological control of introduced plant species. It is important to know the evolutionary changes in tolerance of invasive species after introduction in order to understand the mechanisms of biological invasions and assess the feasibility of biological control. While many studies have explored the evolutionary changes in resistance of invasive species, little has been done to address tolerance. We hypothesized that compared with plants from native populations, plants from invasive populations may increase growth and decrease tolerance to herbivory in response to enemy release in introduced ranges. To test this hypothesis, we compared the differences in growth and tolerance to simulated herbivory between plants from invasive and native populations of Chromolaena odorata, a noxious invader of the tropics and subtropics, at two nutrient levels. Surprisingly, flower number, total biomass (except at high nutrient), and relative increase in height were not significantly different between ranges. Also, plants from invasive populations did not decrease tolerance to herbivory at both nutrient levels. The invader from both ranges compensated fully in reproduction after 50?% of total leaf area had been damaged, and achieved substantial regrowth after complete shoot damage. This strong tolerance to damage was associated with increased resource allocation to reproductive structures and with mobilization of storage reserves in roots. The innately strong tolerance may facilitate invasion success of C. odorata and decrease the efficacy of leaf-feeding biocontrol agents. Our study highlights the need for further research on biogeographical differences in tolerance and their role in the invasiveness of exotic plants and biological control.     

Genetic variation in resistance and tolerance to insect herbivory in Salix cordata

1. This study investigated whether sand-dune willow Salix cordata , exhibits genetic variation in resistance and tolerance to herbivory.
2. A field experiment using cuttings from nine willow clones demonstrated genetic variation in resistance to the specialist herbivore Altica subplicata , as measured by beetle densities. Willow clones differed significantly in both total biomass and leaf trichome densities, and herbivore densities were marginally correlated with both of these parameters.
3. Tolerance to herbivory was measured in a greenhouse experiment by comparing growth response of plants experiencing 50% artificial defoliation and plants experiencing no defoliation. Clones showed significant differences in tolerance to herbivory for some growth measures (changes in height and number of leaves), but not for other growth measures (stem diameter growth and final biomass).
4. Despite the significant genetic variation in both resistance and tolerance, no trade-off was found between resistance and tolerance to herbivory.     

Latitudinal variation in resistance and tolerance to herbivory in the perennial herb Lythrum salicaria is related to intensity of herbivory and plant phenology

Both the length of the growing season and the intensity of herbivory often vary along climatic gradients, which may result in divergent selection on plant phenology, and on resistance and tolerance to herbivory. In Sweden, the length of the growing season and the number of insect herbivore species feeding on the perennial herb Lythrum salicaria decrease from south to north. Previous common‐garden experiments have shown that northern L. salicaria populations develop aboveground shoots earlier in the summer and finish growth before southern populations do. We tested the hypotheses that resistance and tolerance to damage vary with latitude in L. salicaria and are positively related to the intensity of herbivory in natural populations. We quantified resistance and tolerance of populations sampled along a latitudinal gradient by scoring damage from natural herbivores and fitness in a common‐garden experiment in the field and by documenting oviposition and feeding preference by specialist leaf beetles in a glasshouse experiment. Plant resistance decreased with latitude of origin, whereas plant tolerance increased. Oviposition and feeding preference in the glasshouse and leaf damage in the common‐garden experiment were negatively related to damage in the source populations. The latitudinal variation in resistance was thus consistent with reduced selection from herbivores towards the northern range margin of L. salicaria. Variation in tolerance may be related to differences in the timing of damage in relation to the seasonal pattern of plant growth, as northern genotypes have developed further than southern have when herbivores emerge in early summer.