Concentration of atmospheric CO2 and temperature have both been rising for the last three decades. In this century, the temperature has been predicted to rise by 2–5 °C and the CO2 concentration to double. These changes may affect the primary and secondary metabolism of plants and thus have implications for other trophic levels. However, the biotic interactions in changing climate conditions are poorly known. In this study, two questions were addressed: (i) How will climate change affect growth and the amounts of secondary compounds in flexible plant species? and (ii) How will this affect herbivores living on this species. Four clones of the dark‐leaved willow (Salix myrsinifolia (Salisb.)) seedlings were grown in closed‐top chambers with two controlled factors: concentration of atmospheric CO2 and temperature (T). There were four combinations of these factors, each combination replicated four times (total of 16 chambers): (i) Control CO2 (350 ppm) and control T, (ii) Elevated CO2 (700 ppm) and control T, (iii) Control CO2 and elevated T (2 °C), and (iv) Elevated CO2 and elevated T. Stem growth and aerial biomass of the plants were determined; and the leaf phenolics, nitrogen and water concentrations were analysed. In addition the growth rate of larvae and feeding preference of adults of a specialist herbivore, the chrysomelid beetle Phratora vitellinae (L.), on the treated willow leaves were measured. Elevated temperature and CO2 concentration increased the stem biomass and elevated CO2 increased leaf biomass and total aerial biomass of the willows. Patterns of biomass allocation were different in different temperature treatments. At elevated temperature there was less branch and leaf material in relation to stems than at the control temperature. Moreover, patterns of biomass allocation differed among clones. CO2 enhancement increased the specific leaf weight (SLW) and reduced both water and nitrogen content of the leaves, however, leaf area was unaffected by the treatments. Carbon dioxide (CO2) and T enhancement reduced the concentrations of several phenolic compounds in the leaves. Phenolic compounds, nutrients, and water in the leaves might be diluted partly due to increased carbon allocation to different structures (e.g. thickening of cell wall and increase of trichomes, etc.). In some cases plant clones showed specific responses to treatments. The CO2 enhancement reduced the relative growth rate (RGR) of the beetle larvae, and in contrast, temperature elevation increased it. Adult beetles did not clearly discriminate between willow leaves grown in different T and CO2 environments, but tended to eat more leaf material from chambers with doubled CO2 concentration. At elevated CO2 adult beetles may need to eat more leaf material in order to reproduce, which may in turn prolong the life cycles, increasing the risk of being eaten and possibly affecting ability to overwinter successfully. Overall, climate change may significantly modify the dynamic interaction between willow and beetle populations.
The relationship between the food selection of four leaf beetle species (Phratora vitellinae, Plagiodera versicolora, Lochmaea capreae, Galerucella lineola) and the phenolic glycosides of willow (Salix spp.) leaves was tested in laboratory food choice experiments. Four willow species native to the study area (Eastern Finland) and four introduced, cultivated willows were tested.The willow species exhibited profound differences in their phenolic glycoside composition and total concentration. The food selection patterns of the leaf beetles followed closely the phenolic glycoside spectra of the willow species. Both the total amount and the composition of phenolic glycosides affected the feeding by the beetles. Phenolic glycosides apparently have both stimulatory and inhibitory influences on leaf beetle feeding depending on the degree of adaptation of a particular insect. Very rare glycosides or exceptional combination of several glycoside types seem to provide certain willow species with high level of resistance against most herbivorous insects. Analogously the average absolute amount of leaf beetle feeding was lower on the introduced willows than on the native species to which the local herbivores have a good opportunity to become adapted.
The ability to induce galls on plants has evolved independently in many insect orders, but the adaptive significance and evolutionary consequences of gall induction are still largely unknown. We studied these questions by analyzing the concentrations of various plant defense compounds in willow leaves and sawfly galls. We found that the galls are probably nutritionally beneficial for the sawfly larvae, because the concentrations of most defensive phenolics are substantially lower in gall interiors than in leaves. More importantly, changes in chemistry occur in a similar coordinated pattern in all studied willow species, which suggests that the insects control the phenolic biosynthesis in their hosts. The resulting convergence of the chemical properties of the galls both within and between host species indicates that the role of plant chemistry in the evolution of host shifts may be fundamentally less significant in gallers than in other phytophagous insects.
Phenological synchrony between budburst and emergence of larvae is critical for the fitness of many spring-feeding insect herbivores. Therefore, large intraspecific variation in timing of budburst of the host may have a negative effect on the herbivore. We studied how asynchrony between emergence of larvae and budburst affects the fitness of Operophtera brumata (Lepidoptera: Geometridae), a major defoliator of Quercus robur, which can adapt to the phenology of a single tree. It is known that, in maturing leaves of Q. robur, accumulation of condensed tannins has a negative effect on growth of O. brumata. However, there is no information about the effect of hydrolysable tannins and other phenolics that are potential antifeedants. In this study, we also analysed changes in secondary chemistry of the foliage of Q. robur and how different compounds are correlated with growth and survival of O. brumata. The effect of asynchrony on O. brumata was studied in rearing experiments. The neonate larvae were incubated without food for different periods of time. The decline in nutritional quality of foliage was estimated by rearing cohorts of larvae with manipulated hatching times on the leaves of ten individual Q. robur trees. For the chemical analysis, the foliage of these trees was sampled at regular intervals. In the absence of foliage, mortality of neonate larvae started to increase exponentially soon after the larvae emerged. If the larvae missed budburst, the decline in nutritional quality of the foliage led to increased mortality and lower body mass (= fecundity). Hydrolysable tannins were not significantly correlated with performance of the larvae. Only condensed tannins were found to correlate negatively with the growth and survival of O. brumata. Certain individual trees were unsuitable hosts for O. brumata because the decline in quality of the foliage was very rapid. Based on regression equations for increasing rate of mortality and decreasing fecundity, we calculated that a relatively small mismatch of +/-30 degree days between budburst and hatching of larvae leads to a 50% decrease in the fitness of O. brumata. Thus, large phenological variation within a Q. robur stand can limit the colonisation of neighbouring trees by dispersing larvae. Furthermore, the hybridisation of moths adapted to phenologically different trees may lead to maladapted phenology of their offspring.
Seedlings of European white birch (Betula pendula Roth) were grown in growth chambers for one growth season under four carbon dioxide regimes (350, 700, 1050 and 1400 ppm) and at three fertilization levels (0, 100 and 500 kg ha monthly). The soluble carbohydrates and secondary phenolics in the leaves and stems were analysed. It was found that fertilizer addition reduced the amounts of glucose and fructose while sucrose remained almost unaffected. The sugar content of leaves increased at 700 ppm and 1050 ppm of CO and decreased at the highest CO concentration (1400 ppm). The amounts of proanthocyanidins and flavonoids in leaves decreased with fertilization addition and increased with CO enrichment. The production of simple phenolic glucosides varied according to the fertilization and CO treatments. The triterpenoid content of stems seemed to increase with fertilization and CO-addition. Our results indicate that the production of phytochemicals in the birch seedlings is very sensitive to both fertilization and CO addition, which is in agreement with earlier studies, and thus provide some support for the hypothesis of carbon allocation to plant defence when there is an excess of carbon and nutrient. The considerable variation in the production of secondary components may indicate that the synthesis of these defensive metabolites can be regulated by a plant to certain extent, depending on the ability of the plant to acclimate to changes in the physical environment.
Summary1. Community genetics research has posited a genetic basis to the assembly of ecological communities. For arthropod herbivores in particular, there is strong support that genetic variation in host plants is a key factor shaping their diversity and composition. However, the specific plant phenotypes underlying herbivore responses remain poorly explored for most systems. 2. We address this knowledge gap by examining the influence of both genetic and phenotypic variation in a dominant host-plant species, Salix hookeriana, on its associated arthropod herbivore community in a common garden experiment. Specifically, we surveyed herbivore responses among five different arthropod feeding guilds to 26 distinct S. hookeriana genotypes. Moreover, we quantified the heritability of a suite of plant traits that determine leaf quality (e.g. phenolic compounds, trichomes, specific leaf area, C : N) and whole-plant architecture, to identify which traits best accounted for herbivore community responses to S. hookeriana genotype. 3. We found that total herbivore abundance and community composition differed considerably among S. hookeriana genotypes, with strong and independent responses of several species and feeding guilds driving these patterns. We also found that leaf phenolic chemistry displayed extensive heritable variation, whereas leaf physiology and plant architecture tended to be less heritable. Of these traits, herbivore responses were primarily associated with leaf phenolics and plant architecture; however, different herbivore species and feeding guilds were associated with different sets of traits. Despite our thorough trait survey, plant genotype remained a significant predictor of herbivore responses in most trait association analyses, suggesting that unmeasured host-plant characteristics and/or interspecific interactions were also contributing factors. 4. Taken together, our results support that the genetic basis of herbivore community assembly occurs through a suite of plant traits for different herbivore species and feeding guilds. Still, identifying these phenotypic mechanisms requires measuring a broad range of plant traits and likely further consideration of how these traits affect interspecific interactions.
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