PAIRWISE VERSUS DIFFUSE NATURAL SELECTION AND THE MULTIPLE HERBIVORES OF SCARLET GILIA,IPOMOPSIS AGGREGATA

Recent theoretical studies have argued that plant-herbivore coevolution proceeds in a diffuse rather than a pairwise manner in multispecies interactions when at least one of two conditions are met: (1) genetic correlations exist between plant resistances to different herbivore species; and (2) ecological interactions between herbivores sharing a host plant cause nonadditive impacts of herbivory on plant fitness. We present results from manipulative field experiments investigating the single and interactive fitness effects of three types of herbivory on scarlet gilia (Ipomopsis aggregata) over two years of study. We utilize these data to test whether selection imposed by herbivore attack on date of first flowering is pairwise (independent) or diffuse (dependent) in nature. Our results reveal complex patterns of the fitness effect of herbivores. Simulated early season browsing had a strong negative fitness effect on plants and also reduced subsequent insect attack. Surprisingly, this ecological interaction did not translate into significant interactions between clipping and insect manipulations on plant fitness. However, we detected a significant interaction between seed fly and caterpillar herbivory on plant fitness, with the negative effect of either insect being greatest when occurring alone. These results suggest that herbivore-imposed selection may have pairwise and diffuse components. In our selection analysis of flowering phenology, we discovered significant pairwise linear selection imposed by clipping, diffuse linear selection imposed by insects, and diffuse nonlinear selection imposed by clipping and insect attack acting simultaneously. Our results reveal that the evolution of flowering phenology in scarlet gilia may be in response to diffuse and pairwise natural selection imposed by multiple herbivores. We discuss the evolution of resistance characters in light of diffuse versus pairwise forms of linear and nonlinear selection and stress the complexity of selection imposed by suites of interacting species.     

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.     

Increased moth herbivory associated with environmental stress of pinyon pine at local and regional levels

 Using 6 years of observational and experimental data, we examined the hypothesis that water and nutrient stress increase the susceptibility of pinyon pine (Pinus edulis) to the stem- and cone-boring moth (Dioryctria albovittella). At two geographic levels, a local scale of 550 km² and a regional scale of 10,000 km², moth herbivory was strongly correlated with an edaphic stress gradient. At a local scale, from the cinder soils of Sunset Crater to nearby sandy-loam soils, nine of ten soil macro- and micronutrients, and soil water content were lowest in cinder-dominated soils. Herbivore damage was six times greater on trees growing in the most water and nutrient deficient site at Sunset Crater compared to sites with well-developed soils. Percentage silt-clay content of soil, which was highly positively correlated with soil nutrient and soil moisture at a local scale, accounted for 56% of the variation in herbivory at a regional scale among 22 sites. Within and across sites, increased stem resin flow was positively associated with reduced moth attack. On the basis of moth distribution across a stress gradient, we predicted that pinyons growing in highly stressful environments would show increased resistance to herbivores if supplemented with water and/or nutrients. We conducted a 6-year experiment at a high-stress site where individual trees received water only, fertilizer only, and water + fertilizer. Relative to control trees, stem growth and resin flow increased in all three treatments, but only significantly in the water + fertilizer treatment. Although there was no significant difference in herbivore damage among these three treatments, there was an overall reduction in herbivore damage on all treatment trees combined, compared to control trees. This experiment suggests that release from stress leads to increased resistance to insect attack and is consistent with our observational data. While other studies have predicted that short-term stress will result in herbivore outbreaks, our studies extend this prediction to chronically stressed host populations. Finally, while flush-feeders are not predicted to respond positively to stressed host plants, we found a positive association between herbivore attack and stressed pinyon populations. Received: 25 December 1995 / Accepted: 15 August 1996     

Temporal consistency in herbivore responses to glucosinolate polymorphism in populations of wild cabbage (Brassica oleracea)

Natural populations of wild cabbage (Brassica oleracea) show significant qualitative diversity in heritable aliphatic glucosinolates, a class of secondary metabolites involved in defence against herbivore attack. One candidate mechanism for the maintenance of this diversity is that differential responses among herbivore species result in a net fitness balance across plant chemotypes. Such top-down differential selection would be promoted by consistent responses of herbivores to glucosinolates, temporal variation in herbivore abundance, and fitness impacts of herbivore attack on plants varying in glucosinolate profile. A 1-year survey across 12 wild cabbage populations demonstrated differential responses of herbivores to glucosinolates. We extended this survey to investigate the temporal consistency of these responses, and the extent of variation in abundance of key herbivores. Within plant populations, the aphid Brevicoryne brassicae consistently preferred plants producing the glucosinolate progoitrin. Among populations, increasing frequencies of sinigrin production correlated positively with herbivory by whitefly Aleyrodes proletella and negatively with herbivory by snails. Two Pieris butterfly species showed no consistent response to glucosinolates among years. Rates of herbivory varied significantly among years within populations, but the frequency of herbivory at the population scale varied only for B. brassicae. B. brassicae emerges as a strong candidate herbivore to impose differential selection on glucosinolates, as it satisfies the key assumptions of consistent preferences and heterogeneity in abundance. We show that variation in plant secondary metabolites structures the local herbivore community and that, for some key species, this structuring is consistent over time. We discuss the implications of these patterns for the maintenance of diversity in plant defence chemistry.     

Herbivore effects on developmental instability and fecundity of holm oaks

Plants are able to compensate for loss of tissue due to herbivores at a variety of spatial and temporal scales, masking detrimental effects of herbivory on plant fitness at these scales. The stressing effect of herbivory could also produce instability in the development of plant modules, and measures of such instability may reflect the fitness consequences of herbivory if instability is related to components of plant fitness. We analyse the relationships between herbivory, developmental instability and production of female flowers and fruits of holm oak Quercus ilex trees by means of herbivore removal experiments. Removal of leaf herbivores reduced herbivory rates at the tree level, but had no effect on mean production of female flowers or mature fruits, whereas herbivory tended to enhance flower production and had no effect on fruit abortion at the shoot level. Differences in herbivory levels between shoots of the same branch did not affect the size and fluctuating asymmetry of intact leaves. These results indicate compensation for herbivory at the tree level and over-compensation at the shoot level in terms of allocation of resources to female flower production. Removal of insect herbivores produced an increase in the mean developmental instability of leaves at the tree level in the year following the insecticide treatment, and there was a direct relationship between herbivory rates in the current year and leaf fluctuating asymmetry the following year irrespective of herbivore removal treatment. Finally, the production of pistillate flowers and fruits by trees was inversely related to the mean fluctuating asymmetry of leaves growing the same year. Leaf fluctuating asymmetry was thus an estimator of the stressing effects of herbivory on adult trees, an effect that was delayed to the following year. As leaf fluctuating asymmetry was also related to tree fecundity, asymmetry levels provided a sensitive measure of plant performance under conditions of compensatory responses to herbivory.     

GENETIC VARIATION IN A FUNGAL PATHOGEN: RESPONSE TO HOST DEFENSIVE CHEMICALS

In response to infection by shoot infecting pathogens, Scots pine releases cortical resin into affected tissues. The resin contains a mixture of monoterpene compounds (α-pinene, β-pinene, 3-carene, β-myrcene, limonene and β-phellandrene) that retard the growth of a range of pathogens. The proportion of each monoterpene in the resin shows substantial variation among trees within a population. Thus pathogens on different trees encounter quite different monoterpene environments. To investigate the evolutionary response of pathogens to the chemically heterogeneous environment provided by Scots pine, isolates of the ascomycete canker pathogen Crumenulopsis sororia were collected from trees within a range of natural Scots pine populations. Growth rates of these isolates were measured in the presence and absence of five host monoterpenes. Substantial heritable variation for growth rate in the presence and absence of monoterpenes, and for monoterpene tolerance was recorded, suggesting the potential for the evolution of chemically specialized pathogen subpopulations on different trees within a wood. However, genetic correlations between growth rates in different monoterpene environments and between tolerance of different monoterpenes were either positive or non-significant, and there was no evidence of “tradeoffs” in performance under different monoterpene regimes. The results suggest that, on its own, the presence of monoterpene variability within Scots pine will not lead to disruptive selection on the C. sororia population. The relationship between defensive chemical diversity and pest resistance is discussed in the light of these results.     

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.     

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.     

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.     

Chronic herbivory negatively impacts cone and seed production,seed quality and seedling growth of susceptible pinyon pines

Although herbivory often reduces the reproduction of attacked trees, few studies have examined how naturally occurring insect-resistant and susceptible trees differ in their reproduction, nor have these effects been experimentally examined through long-term herbivore removals. In addition, few studies have examined the effects of herbivory on the quality of seeds produced and the implications of reduced seed quality on seedling establishment. We evaluated the impact of chronic herbivory by the stem-boring moth, Dioryctria albovittella, on cone and seed production of the pinyon pine (Pinus edulis) during two mast years. Three patterns emerged. First, moth herbivory was associated with reductions in cone production, viable seed production and seed mass. Specifically, pinyons susceptible to moth attack had 93–95% lower cone production, and surviving cones produced 31–37% fewer viable seeds, resulting in a 96–97% reduction in whole tree viable seed production. In addition, surviving seeds from susceptible trees had 18% lower mass than resistant trees. Second, long-term experimental removal of the herbivore resulted in increased rates of cone and seed production and quality, indicating that moth herbivory was the driver of these reductions. Third, seed size was positively associated with seed germination and seedling biomass and height, suggesting that trees suffering chronic herbivory produce poorer quality offspring. Thus, the resistance traits of pinyons can affect the quality of offspring, which in turn may affect subsequent seedling establishment and population dynamics.     

Antiherbivore defenses alter natural selection on plant reproductive traits

While many studies demonstrate that herbivores alter selection on plant reproductive traits, little is known about whether antiherbivore defenses affect selection on these traits. We hypothesized that antiherbivore defenses could alter selection on reproductive traits by altering trait expression through allocation trade‐offs, or by altering interactions with mutualists and/or antagonists. To test our hypothesis, we used white clover, Trifolium repens, which has a Mendelian polymorphism for the production of hydrogen cyanide—a potent antiherbivore defense. We conducted a common garden experiment with 185 clonal families of T. repens that included cyanogenic and acyanogenic genotypes. We quantified resistance to herbivores, and selection on six floral traits and phenology via male and female fitness. Cyanogenesis reduced herbivory but did not alter the expression of reproductive traits through allocation trade‐offs. However, the presence of cyanogenic defenses altered natural selection on petal morphology and the number of flowers within inflorescences via female fitness. Herbivory influenced selection on flowers and phenology via female fitness independently of cyanogenesis. Our results demonstrate that both herbivory and antiherbivore defenses alter natural selection on plant reproductive traits. We discuss the significance of these results for understanding how antiherbivore defenses interact with herbivores and pollinators to shape floral evolution.     

Effects of two types of herbivores on the population dynamics of a perennial herb

The joint effects of multiple herbivores on their shared host plant have received increasing interest recently. The influence of herbivores on population dynamics of their host plants, especially the relative roles of different types of damage, is, however, still poorly understood. Here, we present a modelling approach, including both deterministic and stochastic matrix modelling, to be used in estimating fitness effects of multiple herbivores on perennial plants. We examined the effects and relative roles of two specialist herbivores, a pre-dispersal seed predator, Euphranta connexa, and a leaf-feeding moth, Abrostola asclepiadis, on the population dynamics and long-term fitness of their shared host plant, a long-lived perennial herb Vincetoxicum hirundinaria (Asclepiadaceae). We collected demographic data during 3 years and combined these data with the effects of natural levels of herbivory measured from the same individuals. We found that both seed predation and leaf herbivory reduced population growth of V. hirundinaria, but only very high damage levels changed the growth trend of the vigorously growing study populations from positive to negative. Demographic modelling indicated that seed predation had a greater impact on plant population growth than leaf herbivory. The effect of leaf herbivory was weaker and diminished with increasing level of seed predation. Evaluation of individual fitness components, however, suggested that leaf herbivory contributed more strongly to host plant fitness than seed predation. Our results emphasize that understanding the effects of a particular herbivore on plant population dynamics requires also knowledge on other herbivores present in the system, because the effect of a particular type of herbivory on plant population dynamics is likely to vary according to the intensity of other types of herbivory. Furthermore, evaluating herbivore impact from using individual fitness components does not necessarily reflect the long-term effects on total plant fitness.     

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.     

Transgenerational consequences of plant responses to herbivory: an adaptive maternal effect?

Herbivory has many effects on plants, ranging from shifts in primary processes such as photosynthesis, growth, and phenology to effects on defense against subsequent herbivores and other species interactions. In this study, I investigated the effects of herbivory on seed and seedling characteristics of several families of wild radish (Raphanus raphanistrum) to test the hypothesis that herbivory may affect the quality of offspring and the resistance of offspring to plant parasites. Transgenerational effects of herbivory may represent adaptive maternal effects or factors that constrain or amplify natural selection on progeny. Caterpillar (Pieris rapae) herbivory to greenhouse-grown plants caused plants in some families to produce smaller seeds and those in other families to produce larger seeds compared with undamaged controls. Seed mass was positively associated with probability of emergence in the field. The number of setose trichomes, a putative plant defense, was higher in the progeny of damaged plants in some families and lower in the progeny of damaged plants in other families. In a field experiment, plant families varied in their resistance to several herbivores and pathogens as well as in growth rate and time to flowering. Seeds from damaged parent plants were more likely to become infested with a plant virus. Although herbivory on maternal plants did not directly affect interactions of offspring with other plant parasites, seed mass influenced plant resistance to several attackers. Thus, herbivory affected seed characters, which mediated interactions between plants and their parasites. Finally, irrespective of seed mass, herbivory on maternal plants influenced components of progeny fitness, which was dependent on plant family. Natural selection may act on plant responses to herbivory that affect seedling-parasite interactions and, ultimately, fitness.     

Foliage value,apparency and defence investment in birch seedlings and trees

Summary Two factors determining plant anti-herbivore defence investment fitness loss due to herbivory and the probability of herbivory occurring in the field were quantified for birch seedlings and trees. Fitness loss due to defoliation (assumed to be related to loss of growth increment compared to controls) appeared to be greater in seedlings compared to trees, but the result was equivocal. In contrast, seedling foliage at the field site — a typical habitat for birch — suffered much less natural defoliation than tree foliage, suggesting that seedlings are markedly less apparent to most birch herbivores than trees. This low apparency should result in lower investment in anti-herbivore defences by seedlings compared to trees — and being a strong effect, should outweigh the possibly greater growth loss suffered by seedlings, which in isolation would tend to increase their optimum defence investment compared to trees. This prediction was tested using palatability trials with a wide range of common birch herbivores and by direct quantification of anti-herbivore defences. Problems and assumptions inherent in these approaches are discussed, but it seems that birch seedlings are genuinely unapparent to herbivores, and consequently do not need the degree of defence investment required by trees.     

Non-host volatiles mediate associational resistance to the pine processionary moth

An increasing body of evidence indicates that the association between different plant species may lead to a reduction in insect herbivory, i.e. associational resistance. This might be due to a top–down regulation of herbivores by increased numbers of natural enemies or to a disruptive bottom–up influence of lower host plant accessibility. In particular, the richer plant communities release more diverse plant odours that may disturb olfactory-guided host choice and mating behaviour of insect herbivores, i.e. the “semiochemical diversity hypothesis”. However, this hypothesis has been rarely tested experimentally in natural habitats, notably forest ecosystems. We tested the effects of non-host volatiles (NHV) on mate and host location by the pine processionary moth (PPM) at the scale of individual pine trees with branches of non-host tree (birch) at their base. Pheromone trap catches and the numbers of larval nests were both reduced by non-host presence under treated pine trees, confirming an associational resistance mediated by NHV. In both males and females, the antenna could detect several birch volatiles, including methyl salicylate (MeSa). MeSa inhibited the attraction of the PPM male to pheromone traps, as did bark and leaf chips from birch trees. Our test of three doses of MeSa at the habitat scale (50 m forest edges) showed that the reduction in the numbers of male PPM captured in traps and in larval nests was MeSa dose-dependent. These results show that odours released by deciduous non-host trees can reduce herbivory by a forest defoliator in conifers, providing support to the “semiochemical diversity hypothesis” as a mechanism of associational resistance.     

Comparative analysis of quantitative trait loci controlling glucosinolates,myrosinase and insect resistance in Arabidopsis thaliana

Evolutionary interactions among insect herbivores and plant chemical defenses have generated systems where plant compounds have opposing fitness consequences for host plants, depending on attack by various insect herbivores. This interplay complicates understanding of fitness costs and benefits of plant chemical defenses. We are studying the role of the glucosinolate-myrosinase chemical defense system in protecting Arabidopsis thaliana from specialist and generalist insect herbivory. We used two Arabidopsis recombinant inbred populations in which we had previously mapped QTL controlling variation in the glucosinolate-myrosinase system. In this study we mapped QTL controlling resistance to specialist (Plutella xylostella) and generalist (Trichoplusia ni) herbivores. We identified a number of QTL that are specific to one herbivore or the other, as well as a single QTL that controls resistance to both insects. Comparison of QTL for herbivory, glucosinolates, and myrosinase showed that T. ni herbivory is strongly deterred by higher glucosinolate levels, faster breakdown rates, and specific chemical structures. In contrast, P. xylostella herbivory is uncorrelated with variation in the glucosinolate-myrosinase system. This agrees with evolutionary theory stating that specialist insects may overcome host plant chemical defenses, whereas generalists will be sensitive to these same defenses.     

Monoterpene hydrocarbon contents of the resin from seeds of silver fir (Abies alba Mill.)

Summary Using gas-chromatographic methods, the variability of the contents of monoterpene hydrocarbons (-pinene, -pinene, ₃carene and limonene) in the resin of silver fir seeds (Abies alba Mill.) was studied. Resin cavities were characterized according to their position on the seed surface. It was estabilished that the terpene content of the resin of cavities localized on the abaxialadaxial surfaces of the seed differs significantly, creating a gradient of resin composition around the circumference of the seed. The differences between various resin cavities of single seeds were greater than the differences between different seeds of a single cone and between seeds of various cones on a single tree. An accurate definition of the resin cavity location on the seeds appears to be a fundamental condition for the collection of a sample representing the resin composition of individual trees. Resin biosynthesis in the course of organogenesis and the control of terpene contents in the resin of various locations on the seed and the cone are discussed.     

Floral trait expression and plant fitness in response to below- and aboveground plant–animal interactions

Although plant–animal interactions like pollination and herbivory are obviously interdependent, ecological investigations focus mainly on one kind of interaction ignoring the possible significance of the others. Plants with flowers offer an extraordinary possibility to study such mutualistic and antagonistic interactions since it is possible to measure changes in floral traits and fitness components in response to different organisms or combinations of them. In a three factorial common garden experiment we investigated single and combined effects of root herbivores, leaf herbivores and decomposers on flowering traits and plant fitness of Sinapis arvensis. Leaf herbivory negatively affected flowering traits indicating that it could significantly affect plant attractiveness to pollinators. Decomposers increased total plant biomass and seed mass indicating that plants use the nutrients liberated by decomposers to increase seed production. We suggest that S. arvensis faced no strong selection pressure from pollen limitation, for two reasons. First, reduced nutrient availability through leaf herbivory affected primarily floral traits that could be important for pollinator attraction. Second, improved nutrient supply through decomposer activity was invested in seed production and not in floral traits. This study indicates the importance of considering multiple plant–animal interactions simultaneously to understand selection pressures underlying plant traits and fitness.     

Cost of trichome production and resistance to a specialist insect herbivore in <Emphasis Type="Italic">Arabidopsis lyrata</Emphasis>

Theory predicts that trade-offs between resistance to herbivory and other traits positively affecting fitness can maintain genetic variation in resistance within plant populations. In the perennial herb Arabidopsis lyrata, trichome production is a resistance trait that exhibits both qualitative and quantitative variation. Using a paternal half-sib design, we conducted two greenhouse experiments to ask whether trichomes confer resistance to oviposition and leaf herbivory by the specialist moth Plutella xylostella, and to examine potential genetic constraints on evolution of increased resistance and trichome density. In addition, we examined whether trichome production is induced by insect herbivory. We found strong positive genetic and phenotypic correlations between leaf trichome density and resistance to leaf herbivory, demonstrating that the production of leaf trichomes increases resistance to leaf damage by P. xylostella. Also resistance to oviposition tended to increase with increasing leaf trichome density, but genetic and phenotypic correlations were not statistically significant. Trichome density and resistance to leaf herbivory were negatively correlated genetically with plant size in the absence of herbivores, but not in the presence of herbivores. There was no evidence of increased trichome production after leaf damage by P. xylostella. The results suggest that trichome production and resistance to leaf herbivory are associated with a cost and that the direction of selection on resistance and trichome density depends on the intensity of herbivory.