Developmental plasticity in birch leaves: defoliation causes a shift from glandular to nonglandular trichomes

The structures on leaf surfaces, e.g. trichomes, can act as effective antiherbivory mechanisms as chemical repellents. Structural defences usually represent constitutive resistance, but there are also a few cases of inducible morphological defences. We tested whether defoliation may induce changes in trichome production in white birch (Betula pubescens). The studied birches were either 0, 50 or 100% defoliated during the previous or current summer, and we measured the alterations in the production of glandular vs. nonglandular leaf trichomes, developmental instability (fluctuating asymmetry, FA) and leaf and shoot growth. We detected a clear shift from glandular to nonglandular leaf trichomes following previous‐year defoliation but not after current‐year defoliation. Furthermore, the density of nonglandular trichomes around the mid‐vein of leaves increased following previous‐year defoliation but decreased after current‐year defoliation. While leaf and shoot growth showed a distinct decrease in response to defoliation, FA turned out to be less sensitive. Consequently, previous‐year defoliation can induce the production of nonglandular trichomes in birch leaves. Because this response was accompanied by a reduction in glandular trichomes, the present results may suggest a trade‐off between the different trichome types of birch leaves.     

Rapid induced resistance of silver birch affects both innate immunity and performance of gypsy moths: the role of plant chemical defenses

In this study we tested the effects of rapid induced resistance of the silver birch, Betula pendula, on the performance and immune defense of the gypsy moth, Lymantria dispar. We also measured the effects of defoliation on the concentrations of plant secondary metabolites, particularly on phenolics and terpenoids. It was found that severe natural defoliation (by moth larvae) of silver birch led to an increase in lipophilic flavonoids on the leaf surface. The concentration of some simple phenolics and monoterpenes (linalool and geraniol) also increased, while that of several glycosides of quercetin decreased. The female pupal weights and survival rates of moths decreased, and larval development time increased, when the insects fed on defoliated trees. However, the feeding of caterpillars with the leaves of defoliated trees led to an increase in lysozyme-like activity in their hemolymph, with an increase in their ability to encapsulate potential parasites. Our data show that the silver birch deploys a rapid chemical defense against gypsy moth larvae. We suggest that lipophilic flavonoids are important compounds in the direct silver birch defense against L. dispar caterpillars. The increased strength of immune defense of insects exposed to trees that had deployed a rapid induced resistance may be an adaptation of the herbivores to resist the rising density of parasites when host population density is high.     

Delayed induced changes in the biochemical composition of host plant leaves during an insect outbreak

In birch, Betula pubescens, herbivore-induced delayed induced resistance (DIR) of defoliated trees may cause a strong reduction in the potential fecundity of a geometrid folivore Epirrita autumnata. In this study, we examined the biochemical basis of DIR in birch leaves during a natural outbreak of E. autumnata. A set of experimental trees was defoliated at four sites by wild larvae in the peak year of the outbreak, whereas control trees were protected from defoliation by spraying with an insecticide. The biochemical composition of leaves was analysed in the following year and, although the DIR response was weak during this outbreak, causing less than a 20% reduction in the potential fecundity of E. autumnata, some consistent relationships between defoliation, biochemistry and pupal mass of E. autumnata suggested a general biochemical basis for the defoliation-induced responses in birch leaves. Total concentrations of nitrogen, sugars and acetone-insoluble residue (e.g. cell wall polysaccharides, cell-wall-bound phenolics, protein, starch, lignin and hemicellulose) were consistently lower, and total concentrations of phenolics, especially of gallotannins and soluble proanthocyanidins, were higher in the leaves of trees defoliated in the previous year than in those protected from defoliation. The capacity of tannins to precipitate proteins correlated with contents of gallotannins, and was highest in defoliated trees. The pupal mass of E. autumnata showed a strong, positive correlation with concentrations of nitrogen and sugars, and a negative correlation with the acetone-insoluble residue and gallotannins in foliage. Correlations with other measured biochemical traits were weak. The correlation coefficients between biochemical traits and pupal mass consistently had similar signs for both defoliated and insecticide–sprayed trees, suggesting that variation in leaf quality due to defoliation in the previous year was based on similar biochemical traits as variation for other reasons. We suggest that DIR is associated with reduced growth activity of leaves, and may be seen as a delay in the biochemical maturation of leaves in defoliated trees. This explains the high concentration of gallotannins in defoliated trees, a characteristic feature of young leaves. However, the lower content of nitrogen and the higher content of soluble proanthocyanidins in defoliated trees are traits usually characterising mature, not young, leaves, indicating defoliation-induced changes in chemistry in addition to modified leaf age. Our results emphasise the importance of understanding the natural changes in chemistry during leaf maturation when interpreting defoliation-induced changes in leaf biochemistry. Received: 26 January 1998 / Accepted: 10 April 1998     

Nonglandular leaf trichomes as short-term inducible defense of the grey alder,Alnus incana (L.), against the chrysomelid beetle,Agelastica alni L.

Summary The grey alder compensates leaf area losses due to insect grazing by continuously producing new leaves throughout the vegetative period. Different degrees of defoliation were attained experimentally by a controlled release of the oligophagous beetle Agelastica alni on arbitrarily selected trees from a homogenous population of young alders. The reduction in leaf area per tree significantly influenced the density of leaf trichomes, assessed 10–30 days later, on newly sprouting leaves only. Cross-correlations between leaf area reduction and trichome density were strongest for leaves which completed unfolding 14–21 days after damage. Dualchoice assays suggested a negative influence of trichomes on oviposition rate of A. alni. Removal of trichomes by shaving demonstrated the highly significant effect of trichomes on feeding behavior of adults and larvae in dual-choice assays. The role of the induced increase in trichome density as a possible short-term defense reaction against herbivorous insects is discussed.     

Glandular trichomes as a barrier against atmospheric oxidative stress: Relationships with ozone uptake,leaf damage,and emission of LOX products across a diverse set of species

There is a spectacular variability in trichome types and densities and trichome metabolites across species, but the functional implications of this variability in protecting from atmospheric oxidative stresses remain poorly understood. The aim of this study was to evaluate the possible protective role of glandular and non‐glandular trichomes against ozone stress. We investigated the interspecific variation in types and density of trichomes and how these traits were associated with elevated ozone impacts on visible leaf damage, net assimilation rate, stomatal conductance, chlorophyll fluorescence, and emissions of lipoxygenase pathway products in 24 species with widely varying trichome characteristics and taxonomy. Both peltate and capitate glandular trichomes played a critical role in reducing leaf ozone uptake, but no impact of non‐glandular trichomes was observed. Across species, the visible ozone damage varied 10.1‐fold, reduction in net assimilation rate 3.3‐fold, and release of lipoxygenase compounds 14.4‐fold, and species with lower glandular trichome density were more sensitive to ozone stress and more vulnerable to ozone damage compared to species with high glandular trichome density. These results demonstrate that leaf surface glandular trichomes constitute a major factor in reducing ozone toxicity and function as a chemical barrier that neutralizes the ozone before it enters the leaf.     

Clumped distribution of oak leaf miners between and within plants

Leaf miners typically show non-random distributions both between and within plants. We tested the hypothesis that leaf miners on two oak species were clumped on individual host trees and individual branches and addressed whether clumping was influenced by aspects of plant quality and how clumping and/or interactions with other oak herbivores affected leaf-miner survivorship. Null models were used to test whether oak herbivores and different herbivore guilds co-occur at the plant scale. Twenty individual Quercus geminata plants and 20 Quercus laevis plants were followed over the season for the appearance of leaf miners and other herbivores, and foliar nitrogen, tannin concentration, leaf toughness and leaf water content were evaluated monthly for each individual tree. The survivorship of the most common leaf miners was evaluated by following the fate of marked mines in several combinations that involved intra- and inter-specific associations. We observed that all leaf miners studied were clumped at the plant and branch scale, and the abundance of most leaf-miner species was influenced by plant quality traits. Mines that occurred singly on leaves exhibited significantly higher survivorship than double and triple mines and leaves that contained a mine or a leaf gall and a mine and damage by chewers exhibited lowest survivorship. Although leaf miners were clumped at individual host trees, null model analyses indicated that oak herbivores do not co-occur significantly less than expected by chance and there was no evidence for biological mechanisms such as inter-specific competition determining community structure at the plant scale. Thus, despite co-occurrence resulting in reduced survivorship at the leaf scale, such competition was not strong enough to structure separation of these oak herbivore communities.     

Artificial Defoliation Induces Trichome Production in the Tropical Shrub Cnidoscolus aconitifolius (Euphorbiaceae)1

Induction of plant defenses and their spatial variability are key subjects in the field of ecology and evolution of defensive traits in plants. Nevertheless, induction has been more commonly studied under controlled environments, ignoring other factors that might influence this process in natural settings. The main goal of this study was to determine if artificial defoliation induces trichome production in three natural populations of the tropical shrub Cnidoscolus aconitifolius. First, we performed trichome counts for each population before imposing artificial defoliation to assess differences in trichome loads between populations. Trichome densities (trichomes/cm²) were quantified for leaf blades, petioles, and flower stalks. To determine if defoliation induced trichome production, three defoliation treatments (0% leaves defoliated or controls, 50% of total leaves defoliated, and 100% defoliation) were applied once at the beginning of the reproductive season. Trichome counts were performed on each structure every ～20 d during a 3‐mo period after the application of treatments. Trichome counts showed significant differences in trichome densities between populations for all three structures. In turn, artificial defoliation increased trichome density. Significant differences among treatments were found for trichome densities on leaf blades and petioles. In both these structures, the 100 percent defoliation treatment differed significantly from control plants, presenting higher trichome densities. In addition, the treatment × population interaction was not significant for leaf blades and petioles, indicating that induction is a generalized response in this species, at least at the study sites. These results indicate that trichomes in C. aconitifolius are inducible due to defoliation.     

Effects of simulated winter browsing on mountain birch foliar chemistry and on the performance of insect herbivores

Winter browsing by mammalian herbivores is known to induce a variety of morphological and physiological changes in plants. Browsing has been suggested to decrease the carbohydrate reserves in woody plants, which might lead to reduced tannin production in leaves during the following summer, and consequently, to increased herbivore damage on leaves. We conducted a clipping experiment with mature mountain birch trees and measured the effects of clipping on birch growth, leaf chemistry and toughness, as well as on the performance of insect herbivores. Leaves grew larger and heavier per unit area in the clipped ramets and had a higher content of proteins than leaves in the control trees. Clipping treatment did not affect the total content of sugars in the leaves (mg g^?1), suggesting that a moderate level of clipping did not significantly reduce the carbohydrate pools of fully‐grown mountain birch trees. Furthermore, the contents of proanthocyanidins (condensed tannins) and gallotannins were slightly higher in the leaves of clipped ramets, contrary to the hypothesis of reduced tannin production. The effects of clipping treatment on leaf and shoot growth and on foliar chemistry were mainly restricted to the clipped ramets, without spreading to untreated ramets within the same tree individual. The effects of clipping on leaf characters varied during the growing season; for instance, leaf toughness in clipped ramets was higher than toughness in control trees and ramets only when leaves were mature. Accordingly, clipping had inconsistent effects on insect herbivores feeding at different times of the growing season. The generally small impact of clipping on herbivore performance suggests that the low intensity of natural browsing at the study area, simulated by our clipping treatment, does not have strong consequences for the population dynamics of insect herbivores on mountain birch via enhanced population growth caused by browsing‐induced changes in food quality.     

Influence of plant ontogeny on compensation to leaf damage

Ontogenetic changes in architecture, carbohydrate reserves, and resource allocation can constrain the ability of plants to compensate for herbivore damage. To evaluate ontogenetic changes in compensation, saplings and reproductive individuals of the tropical tree Casearia nitida were subjected to three levels of defoliation (0, 25, and 75% leaf area removed) and regrowth was quantified. The impact of defoliation on fruit production was evaluated in reproductive trees. In addition, the influence of defoliation on carbohydrate reserves and on the production of phenolic compounds was assessed. Plants at both stages were able to compensate for 25% leaf area loss, but only saplings were able to compensate at the 75% defoliation level. Negative impacts of defoliation on reproductive trees were also suggested by their tendency to produce fewer fruits when defoliated. The concentration of nonstructural carbohydrates decreased with damage in saplings but not in reproductive trees, suggesting an ontogenetic stage-dependent impact of defoliation on carbohydrate reserves. The concentration of phenolic compounds in leaves decreased with increasing leaf damage in both ontogenetic stages. This suggests a resource based trade-off between defense and compensation. The results from this study suggest that ontogeny needs to be considered when assessing plant responses to herbivore damage.     

Allometric analysis of the induced flavonols on the leaf surface of wild tobacco (Nicotiana attenuata)

Trichomes excrete secondary metabolites that may alter the chemical composition of the leaf surface, reducing damage caused by herbivores, pathogens and abiotic stresses. We examined the surface exudates produced by Nicotiana attenuata Torr. Ex Wats., a plant known to contain and secrete a number of secondary metabolites that are toxic or a deterrent to herbivorous insects. Extractions specific to the leaf surface, the trichomes, and the laminar components demonstrated the localization of particular compounds. Diterpene glycosides occurred exclusively in leaf mesophyll, whereas nicotine was found in both the trichomes and mesophyll. Neither rutin nor nicotine was found on the leaf surface. Quercetin and 7 methylated derivatives were found in the glandular trichomes and appeared to be excreted onto the leaf surface. We examined the elicitation of these flavonols on the leaf surface with a surface-area allometric analysis, which measures changes in metabolites independent of the effects of leaf expansion. The flavonols responded differently to wounding, methyl jasmonate (MeJA), herbivore attack and UV-C radiation, and the response patterns corresponded to their compound-specific allometries. Finding greater amounts of quercetin on younger leaves and reduced amounts after herbivore feeding and MeJA treatment, we hypothesized that quercetin may function as an attractant, helping the insects locate a preferred feeding site. Consistent with this hypothesis, mirids (Tupiocoris notatus) were found more often on mature leaves sprayed with quercetin at a concentration typical of young leaves than on unsupplemented mature leaves. The composition of metabolites on the leaf surface of N. attenuata changes throughout leaf development and in response to herbivore attack or environmental stress, and these changes are mediated in part by responses of the glandular trichomes.     

Leaf surface traits: overlooked determinants of birch resistance to herbivores and foliar micro-fungi?

Previous studies on the mechanisms of birch resistance to herbivores and foliar micro-fungi (both pathogenic and endophytic) have focused mainly on the role of internal leaf chemistry. In the present study, we examined genetic correlations between leaf surface traits (glandular trichome density and total concentrations of surface flavonoid aglycones) and occurrence of three species of foliar micro-fungi, one pathogenic rust (Melampsoridium betulinum) and two endophytic fungi (Fusicladium sp. and Melanconium sp.), and performance of autumnal moth larvae (Epirrita autumnata) in two birch species, Betula pubescens ssp. czerepanovii and B. pendula. The performance of autumnal moth larvae on B. pubescens ssp. czerepanovii was negatively correlated with density of glandular trichomes (RGR: r=–0.855; pupal mass: r=–0.709). In addition, rust infection was negatively correlated with trichome density in B. pendula (r=–0.675) and with epicuticular flavonoid aglycones in B. pubescens ssp. czerepanovii (r=–0.855). The frequency of the endophytic fungus Fusicladium sp., was related to epicuticular flavonoid aglycones (r=–0.782), while another endophytic fungus, Melanconium sp., showed no associations with any of the studied variables in B. pubescens ssp. czerepanovii. Our results indicate that leaf surface traits may be at least as important determinants of herbivore performance and micro-fungi abundance in birch as leaf internal chemistry.     

Anti-herbivore strategies in mountain birch at the tree-line

Birch forest on the slope of the Jesnalvaara fell forms a gradient ranging from 7–8 m high trees (at 75 m a.s.l.) to birch shrubs less than 1 m high at the top of the fell (330 m a.s.l.). Upper parts of the forest (above 220 m) were defoliated by Oporinia autumnata (Lep., Geometridae) about ten years ago and, with the exception of the top, only a fraction of birches have recovered. Climatic conditions explain the limitation of the damaged area – Oporinia eggs were killed during cold winters. Predators may also limit the damage. Defensive mechanisms in the birch leaves – especially a low nitrogen content and high concentrations of phenols inhibiting trypsin – can slow down an increase in herbivore density by reducing their reproductive capacity. Birch leaves from the lower limit of the damaged area are least suitable for the growth of herbivores. They have the highest concentration of phenols, too. Birches at the foot of Jesnalvaara are able to increase leaf phenols after mechanical damage of nearby leaves. This is enough to retard the growth of several herbivore species. Ability to recover after defoliation is an important part of the anti-herbivore strategy of the birch. This ability is higher in shrub birches than in tall trees and better after a warm than a cold summer. Birch leaves in the year following defoliation are less usable for herbivores and cause increased mortality and lowered reproduction. These properties are the weakest in birches at the foot of Jesnalvaara where the probability of being defoliated is also lowest.     

Geographic association and temporal variation of chemical and physical defense and leaf damage in Datura stramonium

The evolution of plant defense traits has traditionally been explained trough the “coevolutionary arms race” between plants and herbivores. According to this, specialist herbivores have evolved to cope effectively with the defensive traits of their host plants and may even use them as a cue for host location. We analyzed the geographic association between leaf trichomes, two tropane alkaloids (putative resistance traits), and leaf damage by herbivores in 28 populations of Datura stramonium in central Mexico. Since the specialist leaf beetles Epitrix parvula and Lema trilineata are the main herbivores of D. stramonium in central Mexico, we predicted a positive association between plant defense and leaf damage across populations. Also, if physical environmental conditions (temperature or precipitation) constrain the expression of plant defense, then the geographic variation in leaf damage should be explained partially by the interaction between defensive traits and environmental factors. Furthermore, we studied the temporal and spatial variation in leaf trichome density and leaf damage in five selected populations of D. stramonium sampled in two periods (1997 vs. 2007). We found a positive association between leaf trichomes density and atropine concentration with leaf damage across populations. The interaction between defensive traits and water availability in each locality had a significant effect on the geographic variation in leaf damage. Differences among populations in leaf trichome density are maintained over time. Our results indicate that local plant–herbivore interaction plays an important role in shaping the geographic and temporal variation in plant defense in D. stramonium.     

Induced plant defenses breached? Phytochemical induction protects an herbivore from disease

Although wound-induced responses in plants are widespread, neither the ecological nor the evolutionary significance of phytochemical induction is clear. Several studies have shown, for example, that induced responses can act against both plant pathogens and herbivores simultaneously. We present the first evidence that phytochemical induction can inhibit a pathogen of the herbivore responsible for the defoliation. In 1990, we generated leaf damage by enclosing gypsy moth larvae on branches of red oak trees. We then inoculated a second cohort of larvae with a nuclear polyhedrosis virus (LdNPV) on foliage from the damaged branches. Larvae were less susceptible to virus consumed on foliage from branches with increasing levels of defoliation, and with higher concentrations of gallotannin. Defoliation itself was not related to any of our chemistry measures. Field sampling supported the results of our experiments: death from virus among feral larvae collected from unmanipulated trees was also negatively correlated with defoliation. In 1991, defoliation and gallotannin were again found to inhibit the virus. In addition, gallotannin concentrations were found to be positively correlated with defoliation the previous year. Compared with previous results that demonstrated a delecterious effect of induction on gypsy moth pupal weight and fecundity, the inhibition of the virus should confer an advantage to the gypsy moth. Since leaf damage levels increase as gypsy moth density increases, and since leaf damage inhibits the gypsy moth virus, there is the potential for positive feedback in the system. If phytochemical induction in red oak can inhibit an animal pathogen such as LdNPV, it suggests to us that induction in red oak is a generalized response to tissue damage rather than an adaptive defense against herbivores.     

Sucking herbivore assemblage composition on greenhouse <Emphasis Type="Italic">Ficus</Emphasis> correlates with host plant leaf architecture

Understanding arthropod herbivore selectivity trends towards host plant attributes is essential for predicting plant-associated herbivore assemblage structure. Little is known about such interactions between spontaneous herbivore species and cultivated plants under specific conditions of botanical garden greenhouses. In this study, the taxonomic and functional composition of sucking arthropod herbivore assemblages were correlated with leaf anatomical and surface features of 33 host species of Ficus L. (Moraceae) across four distantly located greenhouse complexes of botanical gardens. The analyses revealed that the species number and abundance of scale insects and their individual families, sessile phloem feeders, total phloem feeders and total herbivores were significantly positively correlated with the thickness of leaf lamina, epidermis and mesophyll, stomatal length and width, and the presence of abaxial multiple epidermis and weak ferruginous non-glandular trichomes. Significant negative correlations were revealed between the same herbivore parameters and the density of glandular trichomes. Heliomorphic leaves supported higher abundance and species richness of sessile phloem feeders compared to sciomorphic ones. The parameters of some phloem and mesophyll feeder taxa also correlated with non-glandular trichome length and density, type of trichomes and epicuticular wax layer, and the presence of calcium oxalate crystals in the epidermis. Results of the study suggest that the leaf architecture-related herbivory trends under greenhouse conditions are similar to those occurring in natural ecosystems when considering the functional significance of particular leaf traits, and remain relevant at the scale of particular plant taxa with disregard of spatial factor.     

Adult beetle grazing induces willow trichome defence against subsequent larval feeding

Induced increases in trichome density to grazing by herbivores have been suggested to increase plant resistance to future herbivore attack. In this paper we present results which show that plants of Salix cinerea L. respond to adult leaf beetle (Phratora vulgatissima L.) grazing by developing new leaves with an increased trichome density. The same plants are usually attacked again later in the season when the next generation of larvae hatches on the plants. The effect of the induced response was studied by comparing larval growth and feeding on newly developed leaves of plants exposed to different defoliation treatments. Larvae on plants previously exposed to adult grazing consumed less total leaf area and showed more dispersed feeding than larvae on plants protected from previous grazing. Larvae on plants exposed to mechanical defoliation responded intermediately. These results corresponded to the increased trichome density of defoliated plants. However, we found this larval response only in whole plant tests--when reared on single, excised leaves in petri dishes, larvae in all treatments behaved similarly. This discrepancy between the on-plant experiment and that in petri dishes highlights how experimental design may alter the conclusion of a study. We suggest that the induced response to adult grazing may act as a defence against subsequent larval feeding.     

Different mechanisms explain feeding type-specific patterns of latitudinal variation in herbivore damage among diverse feeding types of herbivorous insects

To determine whether latitudinal variation in herbivore impact exists, we examined three major herbivorous insect feeding types (chewers, gallers, and miners) on/in leaves of Japanese beech. Herbivores were collected with litter traps deployed in forests across a latitudinal gradient of 10°. Leaf litter analyses demonstrated that chewing herbivory increased with increasing latitude of collection site. However, the densities of miners and gallers decreased with latitude. To test whether latitudinal variation in herbivore damage occurs in the absence of geographically differentiated environmental cueing (e.g., physical stresses or herbivore damage), we measured both genetically determined constitutive leaf traits and herbivore damage in a common-garden experiment. In this experiment, miner density decreased with latitude, but chewing herbivory did not vary latitudinally. Galler density was higher on trees from native provenances than on trees from unrelated provenances likely because of local adaptations. Leaf mass per unit area (LMA), tannin, and phenolics all decreased with latitude of provenance. The latitudinal variation in one constitutive leaf trait (LMA) best explained latitudinal variation in chewing herbivory. Thus, different mechanisms account for feeding type-specific patterns of latitudinal variation in herbivore damage among different herbivore feeding types.     

Impacts of urbanization on insect herbivory and plant defences in oak trees

Systematic comparisons of species interactions in urban versus rural environments can improve our understanding of shifts in ecological processes due to urbanization. However, such studies are relatively uncommon and the mechanisms driving urbanization effects on species interactions (e.g. between plants and insect herbivores) remain elusive. Here we investigated the effects of urbanization on leaf herbivory by insect chewers and miners associated with the English oak Quercus robur by sampling trees in rural and urban areas throughout most of the latitudinal distribution of this species. In performing these comparisons, we also controlled for the size of the urban areas (18 cities) and gathered data on CO₂ emissions. In addition, we assessed whether urbanization affected leaf chemical defences (phenolic compounds) and nutritional traits (phosphorus and nitrogen), and whether such changes correlated with herbivory levels. Urbanization significantly reduced leaf chewer damage but did not affect leaf miners. In addition, we found that leaves from urban locations had lower levels of chemical defences (condensed and hydrolysable tannins) and higher levels of nutrients (nitrogen and phosphorus) compared to leaves in rural locations. The magnitude of urbanization effects on herbivory and leaf defences was not contingent upon city size. Importantly, while the effects of urbanization on chemical defences were associated with CO₂ emissions, changes in leaf chewer damage were not associated with either leaf traits or CO₂ levels. These results suggest that effects of urbanization on herbivory occur through mechanisms other than changes in the plant traits measured here. Overall, our simultaneous assessment of insect herbivory, plant traits and abiotic correlates advances our understanding of the main drivers of urbanization effects on plant–herbivore interactions.     

Attraction of willow warblers to sawfly‐damaged mountain birches: novel function of inducible plant defences?

Plants wounded by invertebrate herbivores emit volatile compounds which invertebrate predators and parasitoids can utilize in locating herbivore prey or hosts. We studied the possibility that an analogical phenomenon might operate between plants and avian insectivores. We show that foliar damage by sawfly larvae on the mountain birch led a passerine bird (willow warbler) to prefer intact branches from trees with introduced larvae over intact branches from control trees. Besides olfaction, the UV vision of birds offers a possible mechanism, as some herbivore‐inducible leaf compounds, e.g. surface flavonoids, have UV spectral maxima well within the range of birds’ UV vision.