Variation in arbuscular mycorrhizal fungi colonization modifies the expression of tolerance to above-ground defoliation

1.  Plant association with arbuscular mycorrhizal fungi (AMF) has been considered a factor increasing plant tolerance to herbivory. However, this positive effect could decrease with colonization density if the benefit : cost ratio of the AMF–plant association changes. We measured plant performance and tolerance to defoliation across a gradient of commercial AMF ( Glomus sp.) inoculum concentration.
2.  Six genetic families of Datura stramonium were grown under greenhouse conditions and subjected to five increasing levels of AMF inoculum concentration and to defoliation treatments, i.e. the presence/absence of 50% artificial damage, following a full-factorial design.
3.  AMF colonization increased linearly with inoculum concentration while foliar area, root mass, flowering phenology and seed production expressed nonlinear functions. Plant genetic variation in the benefit function of AMF colonization was also detected. We show a negative interaction between AMF concentration and plant tolerance to defoliation.
4.   Synthesis . The negative correlation between plant tolerance and AMF concentration suggests that defoliation can reduce AMF benefits and that natural variations in AMF can limit the evolution of optimum levels of tolerance. Moreover, genetic variation in the shape of the reaction norms to AMF in the presence/absence of defoliation suggests that plants may evolve in response to variation in densities of AMF and herbivores.     

Genetic Constraints and Selection Acting on Tolerance to Herbivory in the Common Morning Glory Ipomoea purpurea

Tolerance to herbivory minimizes the effects of herbivory on plant fitness. In the presence of herbivores, maximal levels of tolerance may be expected to evolve. In many plant species, however, tolerance is found at an intermediate level. Tolerance may be prevented from evolving to a maximal level by genetic constraints or stabilizing selection. We report on a field study of Ipomoea purpurea, the common morning glory, in which we measured three types of costs that may be associated with tolerance and the pattern of selection acting on tolerance to two types of herbivore damage: apical meristem damage and folivory. We used genetic correlations to test for the presence of three types of costs: a trade-off between tolerance and fitness in the absence of herbivore damage, a trade-off between tolerance and resistance, and genetic covariances among tolerance to different types of damage. We found no evidence that tolerance to apical meristem damage or tolerance to folivory was correlated with resistance, although these two types of tolerance were positively correlated with one another. Tolerance to both types of damage involved costs of lower fitness in the absence of herbivory. Selection acting on tolerance to either type of herbivory was not detected at natural levels of herbivory. Selection is expected to act against tolerance at reduced levels of herbivory and favor tolerance at elevated levels of herbivory. In addition, significant correlational selection gradients indicate that the pattern of selection acting on tolerance depends on values of resistance. Although we found no evidence for stabilizing selection, fluctuating selection resulting from fluctuating herbivore loads may be responsible for maintaining tolerance at an intermediate level.     

Intraspecific competition facilitates the evolution of tolerance to insect damage in the perennial plant Solanum carolinense

Tolerance to herbivory (the degree to which plants maintain fitness after damage) is a key component of plant defense, so understanding how natural selection and evolutionary constraints act on tolerance traits is important to general theories of plant–herbivore interactions. These factors may be affected by plant competition, which often interacts with damage to influence trait expression and fitness. However, few studies have manipulated competitor density to examine the evolutionary effects of competition on tolerance. In this study, we tested whether intraspecific competition affects four aspects of the evolution of tolerance to herbivory in the perennial plant Solanum carolinense: phenotypic expression, expression of genetic variation, the adaptive value of tolerance, and costs of tolerance. We manipulated insect damage and intraspecific competition for clonal lines of S. carolinense in a greenhouse experiment, and measured tolerance in terms of sexual and asexual fitness components. Compared to plants growing at low density, plants growing at high density had greater expression of and genetic variation in tolerance, and experienced greater fitness benefits from tolerance when damaged. Tolerance was not costly for plants growing at either density, and only plants growing at low density benefited from tolerance when undamaged, perhaps due to greater intrinsic growth rates of more tolerant genotypes. These results suggest that competition is likely to facilitate the evolution of tolerance in S. carolinense, and perhaps in other plants that regularly experience competition, while spatio-temporal variation in density may maintain genetic variation in tolerance.     

Population variation in the cost and benefit of tolerance and resistance against herbivory in Datura stramonium

In this study we examine the hypothesis that divergent natural selection produces genetic differentiation among populations in plant defensive strategies (tolerance and resistance) generating adaptive variation in defensive traits against herbivory. Controlled genetic material (paternal half-sib families) from two populations of the annual Datura stramonium genetically differentiated in tolerance and resistance to herbivory were used. This set of paternal half-sib families was planted at both sites of origin and the pattern of genotypic selection acting on tolerance and resistance was determined, as well as the presence and variation in the magnitude of allocational costs of tolerance. Selection analyses support the adaptive differentiation hypothesis. Tolerance was favored at the site with higher average level of tolerance, and resistance was favored at the site with higher average level of resistance. The presence of significant environmentally dependent costs of tolerance was in agreement with site variation in the adaptive value of tolerance. Our results support the expectation that environmentally dependent costs of plant defensive strategies can generate differences among populations in the evolutionary trajectory of defensive traits and promote the existence of a selection mosaic. The pattern of contrasting selection on tolerance suggests that, in some populations of D. stramonium, tolerance may alter the strength of reciprocal coevolution between plant resistance and natural enemies.     

Evolutionary genetics of resistance and tolerance to natural herbivory in Arabidopsis thaliana

Resistance and tolerance are widely viewed as two alternative adaptive responses to herbivory. However, the traits underlying resistance and tolerance remain largely unknown, as does the genetic architecture of herbivory responses and the prevalence of genetic trade-offs. To address these issues, we measured resistance and tolerance to natural apical meristem damage (AMD) by rabbits in a large field experiment with recombinant inbred lines (RILs) of Arabidopsis thaliana (developed from a cross between the Columbia x Landsberg erecta ecotypes). We also measured phenological and morphological traits hypothesized to underlie resistance and tolerance to AMD. Recombinant inbred lines differed significantly in resistance (the proportion of replicates within an RIL that resisted herbivory), and early flowering plants with tall apical inflorescences were more likely to experience damage. Tolerance (the difference in fitness between the damaged and undamaged states), also differed significantly among RILs, with some lines overcompensating for damage and producing more fruit in the damaged than undamaged state. Plastic increases in basal branch number, basal branch height, and senescence date in response to damage were all associated with greater tolerance. There was no evidence for a genetic trade-off between resistance and tolerance, an observation consistent with the underlying differences in associated morphological and phenological characters. Selection gradient analysis detected no evidence for direct selection on either resistance or tolerance in this experiment. However, a statistical model indicates that the pattern of selection on resistance depends strongly on the mean level of tolerance, and selection on tolerance depends strongly on the mean level of resistance. These observations are consistent with the hypothesis that selection may act to maintain resistance and tolerance at intermediate levels in spatially or temporally varying environments or those with varying herbivore populations.     

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.     

Invasive and native populations of common ragweed exhibit strong tolerance to foliar damage

Tolerance and resistance are defence strategies evolved by plants to cope with damage due to herbivores. The introduction of exotic species to a new biogeographical range may alter the plant–herbivore interactions and induce selection pressures for new plant defence strategies with a modified resource allocation. To detect evolution in tolerance to herbivory in common ragweed, we compared 3 native (North America) and 3 introduced (France) populations, grown in a common garden environment. We explored the effect of leaf herbivory on plant vegetative and reproductive traits. Plants were defoliated by hand, simulating different degrees of insect grazing by removing 0%, 50% or 90% of each leaf blade. Total and shoot dry biomasses were not affected by increasing defoliation, whereas root dry biomass and root:shoot ratio decreased significantly for native and introduced populations. Furthermore, defoliation treatments did not affect any of the plant reproductive traits measured. Hence, common ragweed displayed an efficient reallocation of resources in shoot biomass at the expense of roots following defoliation, which allows the species to tolerate herbivory without obvious costs for fitness. We did not detect any difference in herbivory tolerance between introduced and native populations, but significant differences were found in reproduction with invasive populations producing more seeds than native populations. As a result, tolerance to herbivory has been maintained in the introduced plant populations. We discuss some implications of these preliminary results for biological control strategies dedicated to common ragweed.     

Resistance and tolerance to herbivory changes with inbreeding and ontogeny in a wild gourd (Cucurbitaceae)

Herbivory is a ubiquitous component of terrestrial communities that reduces plant growth and reproduction. Consequently, a goal of evolutionary ecology is to identify the causes and consequences of variation in herbivory within plant populations. This three-year study examined the effects of inbreeding on the resistance of wild gourd plants (Cucurbita pepo subsp. texana) to herbivory by cucumber beetles and the impact of the timing of herbivory on reproduction. We grew families of inbred and outbred gourds and recorded beetle damage at three developmental stages, incidence of beetle-vectored wilt disease, survival, and reproduction. While total beetle damage significantly depressed flower and fruit production, damage until mid-July did not depress any measure of reproduction, indicating that these gourds are tolerant of moderate levels of herbivory for most of the growing season. However, beetle damage accumulating after mid-July significantly depressed reproduction, indicating that plants have reduced tolerance during peak reproduction. Early damage, however, did increase the probability of contracting a deadly wilt disease that is vectored by the beetles, suggesting that tolerance and resistance are not alternative defense strategies. Inbreeding significantly reduced resistance to herbivory and, independently of beetle damage, reproductive output. Finally, we found additive genetic variation for both resistance and tolerance that varies with ontogeny.     

Tolerance to apical and leaf damage of Raphanus raphanistrum in different competitive regimes

Tolerance to herbivory is an adaptation that promotes regrowth and maintains fitness in plants after herbivore damage. Here, we hypothesized that the effect of competition on tolerance can be different for different genotypes within a species and we tested how tolerance is affected by competitive regime and damage type. We inflicted apical or leaf damage in siblings of 29 families of an annual plant Raphanus raphanistrum (Brassicaceae) grown at high or low competition. There was a negative correlation of family tolerance levels between competition treatments: plant families with high tolerance to apical damage in the low competition treatment had low tolerance to apical damage in the high competition treatment and vice versa. We found no costs of tolerance, in terms of a trade‐off between tolerance to apical and leaf damage or between tolerance and competitive ability, or an allocation cost in terms of reduced fitness of highly tolerant families in the undamaged state. High tolerance bound to a specific competitive regime may entail a cost in terms of low tolerance if competitive regime changes. This could act as a factor maintaining genetic variation for tolerance.     

COSTS OF INDUCED RESPONSES AND TOLERANCE TO HERBIVORY IN MALE AND FEMALE FITNESS COMPONENTS OF WILD RADISH

Theory predicts that plant defensive traits are costly due to trade-offs between allocation to defense and growth and reproduction. Most previous studies of costs of plant defense focused on female fitness costs of constitutively expressed defenses. Consideration of alternative plant strategies, such as induced defenses and tolerance to herbivory, and multiple types of costs, including allocation to male reproductive function, may increase our ability to detect costs of plant defense against herbivores. In this study we measured male and female reproductive costs associated with induced responses and tolerance to herbivory in annual wild radish plants (Raphanus raphanistrum). We induced resistance in the plants by subjecting them to herbivory by Pieris rapae caterpillars. We also induced resistance in plants without leaf tissue removal using a natural chemical elicitor, jasmonic acid; in addition, we removed leaf tissue without inducing plant responses using manual clipping. Induced responses included increased concentrations of indole glucosinolates, which are putative defense compounds. Induced responses, in the absence of leaf tissue removal, reduced plant fitness when five fitness components were considered together; costs of induction were individually detected for time to first flower and number of pollen grains produced per flower. In this system, induced responses appear to impose a cost, although this cost may not have been detected had we only quantified the traditionally measured fitness components, growth and seed production. In the absence of induced responses, 50% leaf tissue removal, reduced plant fitness in three out of the five fitness components measured. Induced responses to herbivory and leaf tissue removal had additive effects on plant fitness. Although plant sibships varied greatly (49–136%) in their level of tolerance to herbivory, costs of tolerance were not detected, as we did not find a negative association between the ability to compensate for damage and plant fitness in the absence of damage. We suggest that consideration of alternative plant defense strategies and multiple costs will result in a broader understanding of the evolutionary ecology of plant defense.     

Evolutionary ecology of Datura stramonium: equal plant fitness benefits of growth and resistance against herbivory

This study evaluated how natural selection act upon two proposed alternatives of defence (growth and resistance) against natural enemies in a common garden experiment using genetic material (full-sibs) from three populations of the annual plant Datura stramonium. Genetic and phenotypic correlations were used to search for a negative association between both alternatives of defence. Finally, the presence/absence of natural enemies was manipulated to evaluate the selective value of growth as a response against herbivory. Results indicated the presence of genetic variation for growth and resistance (1--relative damage), whereas only population differentiation for resistance was detected. No correlation between growth and resistance was detected either at the phenotypic or the genetic level. Selection analysis revealed the presence of equal fitness benefits of growth and resistance among populations. The presence/absence of natural herbivores revealed that herbivory did not alter the pattern of selection on growth. The results indicate that both strategies of defence can evolve simultaneously within populations of D. stramonium.     

The effect of inbreeding on defence against multiple enemies in Datura stramonium

The ability of plants to respond to natural enemies might depend on the availability of genetic variation for the optimal phenotypic expression of defence. Selfing can affect the distribution of genetic variability of plant fitness, resistance and tolerance to herbivores and pathogens. The hypothesis of inbreeding depression influencing plant defence predicts that inbreeding would reduce resistance and tolerance to damage by natural enemies relative to outcrossing. In a field experiment entailing experimentally produced inbred and outcrossed progenies, we assessed the effects of one generation of selfing on Datura stramonium resistance and tolerance to three types of natural enemies, herbivores, weevils and a virus. We also examined the effect of damage on relative growth rate (RGR), flower, fruit, and seed production in inbred and outcrossed plants. Inbreeding significantly reduced plant defence to natural enemies with an increase of 4% in herbivore damage and 8% in viral infection. These results indicate inbreeding depression in total resistance. Herbivory increased 10% inbreeding depression in seed number, but viral damage caused inbred and outcrossed plants to have similar seed production. Inbreeding and outcrossing effects on fitness components were highly variable among families, implying that different types or numbers of recessive deleterious alleles segregate following inbreeding in D. stramonium. Although inbreeding did not equally alter all the interactions, our findings indicate that inbreeding reduced plant defence to herbivores and pathogens in D. stramonium.     

Effect of water availability on tolerance of leaf damage in tall morning glory,Ipomoea purpurea

Resource availability may limit plant tolerance of herbivory. To predict the effect of differential resource availability on plant tolerance, the limiting resource model (LRM) considers which resource limits plant fitness and which resource is mostly affected by herbivore damage. We tested the effect of experimental drought on tolerance of leaf damage in Ipomoea purpurea, which is naturally exposed to both leaf damage and summer drought. To seek mechanistic explanations, we also measured several morphological, allocation and gas exchange traits. In this case, LRM predicts that tolerance would be the same in both water treatments. Plants were assigned to a combination of two water treatments (control and low water) and two damage treatments (50% defoliation and undamaged). Plants showed tolerance of leaf damage, i.e., a similar number of fruits were produced by damaged and undamaged plants, only in control water. Whereas experimental drought affected all plant traits, leaf damage caused plants to show a greater leaf trichome density and reduced shoot biomass, but only in low water. It is suggested that the reduced fitness (number of fruits) of damaged plants in low water was mediated by the differential reduction of shoot biomass, because the number of fruits per shoot biomass was similar in damaged and undamaged plants. Alternative but less likely explanations include the opposing direction of functional responses to drought and defoliation, and resource costs of the damage-induced leaf trichome density. Our results somewhat challenge the LRM predictions, but further research including field experiments is needed to validate some of the preliminary conclusions drawn.     

Environmental dependency in the expression of costs of tolerance to deer herbivory

Plant tolerance to natural enemy damage is a defense strategy that minimizes the effects of damage on fitness. Despite the apparent benefits of tolerance, many populations exhibit intermediate levels of tolerance, indicating that constraints on the evolution of tolerance are likely. In a field experiment with the ivyleaf morning glory, costs of tolerance to deer herbivory in the form of negative genetic correlations between deer tolerance and fitness in the absence of damage were detected. However, these costs were detected only in the presence of insect herbivores. Such environmental dependency in the expression of costs of tolerance may facilitate the maintenance of tolerance at intermediate levels.     

EXPERIMENTAL EVOLUTION OF RESISTANCE IN BRASSICA RAPA: CORRELATED RESPONSE OF TOLERANCE IN LINES SELECTED FOR GLUCOSINOLATE CONTENT

The evolutionary response of plant populations to selection for increased defense may be constrained by costs of defense. The purpose of this study was to investigate such constraints on the evolution of defense due to a cost of defense manifested as a trade-off between defense and tolerance. Variation in the response to artificial damage (tolerance) among lines of Brassica rapa that had been artificially selected for foliar glucosinolate content (defense) was examined. Leaf area was removed from replicates of three selection lines (high glucosinolates, control, and low glucosinolates) at three damage levels (0%, 20%, and 60% damage). An external cost of defense would result in a statistically significant selection line by damage treatment interaction, with those selected for high defense expressing less tolerance than those selected for low defense. Damage treatment had a significant overall effect on estimated total fitness, with fitness declining with increasing damage level. Further, selection line also had a significant overall effect on estimated total fitness, with low-defense selection lines having higher fitness compared to both control and high-defense selection lines. More importantly, a cost of defense in terms of tolerance was demonstrated by a significant selection line-by-damage treatment interaction. This interaction was in the direction to demonstrate a genetic trade-off between defense and tolerance, with low-defense selection lines decreasing estimated total fitness in response to damage less than both control and high-defense selection lines. Variation in tolerance among selection lines was due to the greater ability of low-defense lines to maintain fruit and seed production despite the presence of damage. In terms of tolerance, this cost of glucosinolate production in B. rapa could constrain the evolution of increased defense and, in so doing, maintain individuals within the population that are poorly defended yet tolerant.     

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.     

Ontogenetic patterns in the mechanisms of tolerance to herbivory in Plantago

Background and Aims

Herbivory and plant defence differ markedly among seedlings and juvenile and mature plants in most species. While ontogenetic patterns of chemical resistance have been the focus of much research, comparatively little is known about how tolerance to damage changes across ontogeny. Due to dramatic shifts in plant size, resource acquisition, stored reserves and growth, it was predicted that tolerance and related underlying mechanisms would differ among ontogenetic stages.

Methods

Ontogenetic patterns in the mechanisms of tolerance were investigated in Plantago lanceolata and P. major (Plantaginaceae) using the genetic sib-ship approach. Pot-grown plants were subjected to 50 % defoliation at the seedling, juvenile and mature stages and either harvested in the short-term to look at plasticity in growth and photosynthesis in response to damage or allowed to grow through seed maturation to measure phenology, shoot compensation and reproductive fitness.

Key Results

Tolerance to defoliation was high in P. lanceolata, but low in P. major, and did not vary among ontogenetic stages in either species. Mechanisms underlying tolerance did vary across ontogeny. In P. lanceolata, tolerance was significantly related to flowering (juveniles) and pre-damage shoot biomass (mature plants). In P. major, tolerance was significantly related to pre-damage root biomass (seedlings) and induction of non-photochemical quenching, a photosynthetic parameter (juveniles).

Conclusions

Biomass partitioning was very plastic in response to damage and showed associations with tolerance in both species, indicating a strong role in plant defence. In contrast, photosynthesis and phenology showed weaker responses to damage and were related to tolerance only in certain ontogenetic stages. This study highlights the pivotal role of ontogeny in plant defence and herbivory. Additional studies in more species are needed to determine how seedlings tolerate herbivory in general and whether mechanisms vary across ontogeny in consistent patterns.     

Tolerance responses to herbivory: implications for future management strategies in potato

Tolerance is a plant response to herbivory that allows plants to sustain defoliation without any fitness consequences. In agriculture, the maximum amount of defoliation that crops can experience without yield loss is defined as the damage threshold. Damage thresholds can be used to develop tolerance‐based management strategies, which would offer guidelines for farmers for when pest control should be enacted to minimise pesticide use while maintaining yield. Damage thresholds for many varieties of potatoes, Solanum tuberosum, have been documented, but not consolidated into a general framework. We aggregated data from published studies and quantified the effects of variety, maturity group, life stage and defoliation mechanisms on damage thresholds to inform a tolerance‐based management strategy for potato. Late maturing varieties did not have different damage thresholds than early maturing varieties. Damage thresholds were greater than 60% defoliation at emergence and post‐bloom and less than 40% defoliation pre‐bloom and during bloom. If the stems were damaged, emergence and post‐bloom damage thresholds were reduced from 60% defoliation without stem damage to 35% and 52% defoliation with stem damage. Likewise, the damage thresholds during bloom were reduced to 20% defoliation when stems were damaged. We propose using action thresholds based on defoliation as a baseline to develop a tolerance‐based management strategy for potatoes.     

Resource allocation in an annual herb: Effects of light,mycorrhizal fungi,and defoliation

Concurrent interactions and the availability of resources (e.g., light) affect the cost/benefit balance during mutualistic and antagonistic interactions, as well as plant resource allocation patterns. Mycorrhizal interactions and herbivory concur in most plants, where mycorrhizae can enhance the uptake of soil nutrients by plants as well as consuming a large fraction of the plant's carbon, and defoliation usually reduces light interception and photosynthesis, thereby causing direct losses to the hosts of mycorrhizal fungi. Both types of interactions affect the carbon budget of their host plants and thus we predict that the relative costs of herbivory and mycorrhizal colonization will increase when photosynthesis is reduced, for instance in light limited environments. We conducted a greenhouse experiment using Datura stramonium to investigate the effects of defoliation and mycorrhizal inoculation on the resource allocation patterns in two different light environments. Defoliated plants overcompensated in terms of leaf mass in both light environments, but total seed mass per fruit was negatively affected by defoliation in both light environments. Mycorrhizal inoculation had a positive effect on vegetative growth and the leaf nitrogen content, but defoliation negates the benefit of mycorrhizal interactions in terms of the leaf nitrogen content. In general, D. stramonium compensated for the relative costs of concurrent mycorrhizal interactions and defoliation; plants that lacked both interactions exhibited the same performance as plants with both types of interactions.     

Ecological genetics of an induced plant defense against herbivores: additive genetic variance and costs of phenotypic plasticity

Abstract.— Adaptive phenotypic plasticity in chemical defense is thought to play a major role in plant-herbivore interactions. We investigated genetic variation for inducibility of defensive traits in wild radish plants and asked if the evolution of induction is constrained by costs of phenotypic plasticity. In a greenhouse experiment using paternal half-sibling families, we show additive genetic variation for plasticity in glucosinolate concentration. Genetic variation for glucosinolates was not detected in undamaged plants, but was significant following herbivory by a specialist herbivore, Pieris rapae . On average, damaged plants had 55% higher concentrations of glucosinolates compared to controls. In addition, we found significant narrow-sense heritabilities for leaf size, trichome number, flowering phenology, and lifetime fruit production. In a second experiment, we found evidence of genetic variation in induced plant resistance to P. rapae . Although overall there was little evidence for genetic correlations between the defensive and life-history traits we measured, we show that more plastic families had lower fitness than less plastic families in the absence of herbivory (i.e., evidence for genetic costs of plasticity). Thus, there is genetic variation for induction of defense in wild radish, and the evolution of inducibility may be constrained by costs of plasticity.