Interactions and competition processes among tree species in young experimental mixed forests, assessed with chlorophyll fluorescence and leaf morphology.

Chlorophyll a fluorescence (ChlF) and leaf morphology were assessed in two sites in Europe (Kaltenborn, Germany, and Satakunta, Finland) within a forest diversity experiment. Trees at Satakunta, planted in 1999, form a stratified canopy, while in Kaltenborn the trees are 7 years old, with no apparent canopy connection among broadleaf species. The following ChlF parameters from measured OJIP transient curves were examined: F(V)/F(M) (a proxy for maximum quantum yield); ΨEo (a proxy for efficiency in transferring an electron from reduced QA to the electron transport chain); I-P phase (a proxy for efficiency of reducing final acceptors beyond PSI); and PItot (total performance index for potential energy conservation from photons absorbed by PSII to reduction of PSI end acceptors). At Satakunta F(V)/F(M) and ΨEo in Betula pendula were higher in monocultures and lower in mixed plots, perhaps due to increasing light availability in mixed plots, which can induce photoinhibition. The opposite trend was observed in Picea abies, which was shaded in mixed plots. At Kaltenborn F(V)/F(M) decreased in Fagus sylvatica and P. abies in mixed plots due to competition both above- and belowground. At Satakunta LMA increased in B. pendula leaves with increasing species richness. Leaf area of ten leaves was reduced in F. sylvatica in mixed plots at Kaltenborn. By up-scaling the overall fluorescence response to plot level (PItot_plot ), a significant positive correlation with tree diversity was found at Kaltenborn, but not at Satakunta. This could suggest that competition/facilitation processes in mixed stands play a significant role in the early stages of forest establishment, but then tend to be compensated in more mature stands.

Covariation of fluctuating asymmetry, herbivory and chemistry during birch leaf expansion.

Fluctuating asymmetry (FA) is used to describe developmental instability in bilateral structures. In trees, high FA of leaves has been assumed to indicate the level of environmental or genetic stress, and for herbivores leaves from such trees have been shown to be in some cases (though not invariably) of higher quality compared to trees with symmetrical leaves. We demonstrated that FA of birch leaves correlated positively with growth rate of leaves, and with the amount of leaf biomass consumed by larvae of the geometrid Epirrita autumnata. Since asymmetry per se cannot define leaf quality for a herbivore, we determined the biochemical compounds which covary with the degree of foliage FA, in order to elucidate relationships between leaf FA, chemistry and herbivory. High foliar FA was characteristic of birches with high initial concentrations, and rapid seasonal decline in the concentrations of gallic acid and hydrolysable tannins, and with rapid seasonal changes in the concentrations of flavonoid-glycosides and sugars. In contrast, leaf FA was not related to concentrations of proanthocyanidins, protein-bound amino acids or soluble phenylalanine, the precursor of proanthocyanidins and proteins with aromatic amino acids. The positive correlation between leaf FA and consumption by E. autumnata was presumably related to the previously demonstrated compensatory consumption of E. autumnata to high concentrations of foliar gallotannins. Furthermore, sugars are well-known feeding stimulants. We propose that the variable results in studies correlating leaf FA and herbivory may stem from variable chemical associations of FA in different plants and of species-specific effects of compounds on insects.

Insect performance on experimentally stressed woody plants: a meta-analysis.

In this review, we test the hypothesis that abiotic stress increases the suitability of plants as food for herbivores. We conducted a meta-analysis that included 70 experimental studies in which insect performance was measured on woody plants subjected to water stress, pollution, and/or shading. Overall, plant stress had no significant effect on insect growth rate, fecundity, survival, or colonization density. We found great variation, however, in the magnitude and direction of insect responses among studies, most of which was related to insect feeding guild. In general, boring and sucking insects performed better on stressed plants, whereas plant stress adversely affected gall-makers and chewing insects. Reduction in performance of chewers was greater on stressed slow-growing plants than on stressed fast growers. Reproductive potential of sucking insects was increased by pollution but reduced by water stress. In some cases where sample sizes were small or the treatment periods short, apparent differences in insect responses to stress were probably artifacts due to inappropriate experimental design.

Antioxidant responses to simulated acid rain and heavy metal deposition in birch seedlings.

This study measured the responses of different anti-oxidants in 2-year-old birch (Betula pendula Roth) seedlings subjected to simulated acid rain (pH 4.0) and heavy metals (Cu/Ni), applied alone or in combination for 2 months. The applied concentrations of pollutants did not significantly affect seedling biomass or total glutathione levels. Acid rain alone increased superoxide dismutase (SOD) activity both in leaves and roots, while heavy metals alone inhibited SOD activity in roots. Both acid rain and heavy metals applied singly increased ascorbate peroxidase (APX) and guaiacol peroxidase (GPX) activities in leaves but decreased activities in roots. In contrast, acid rain and heavy metal treatments increased glutathione reductase (GR) activity in roots but not in leaves. Spraying birch seedlings with a mixture of acid rain and heavy metals increased SOD, APX and GPX activities in leaves and GR activity in roots. However, the effects of mixed pollutants on enzyme activities usually were less than the summed effects of individual pollutants. Enzyme responses also depended on where pollutants were applied: spraying pollutants onto the shoots initiated higher responses in SOD, APX and GPX than did application to the soil surface, while the opposite was true for GR.

Density patterns of gall mites (Acarina:Eriophyidae) in a polluted area.

Trends in the densities of six species of gall mites on European aspen (Populus tremula) and on two birch species (Betula pubescens and B. pendula) were compared in an air pollution gradient from the Harjavalta copper-nickel smelter, SW Finland. The densities of gall mites on both birch species decreased towards the smelter and were negatively correlated with the levels of copper and nickel in the birch leaves. In contrast, the densities of aspen mites correlated neither with distance from the pollution source nor with the content of heavy metals in aspen leaves. Both birch and aspen trees tended to produce smaller leaves near the smelter, but a significant correlation between gall mite densities and site-specific leaf areas was only found for one species of mite.

Variations in chemical composition of birch foliage under air pollution stress and their consequences for Eriocrania miners.

The relationships between chemical composition of birch foliage, individual performance and population density of both solitary and gregarious species of Eriocrania leafminers (Lepidoptera: Eriocraniidae) were studied in the vicinity of the Harjavalta copper-nickel smelter, southwestern Finland. The contents of heavy metals (Cu, Ni, Fe and Zn) in birch foliage exponentially decreased with the distance from the factory whereas the content of manganese showed the opposite trend. Performance parameters of Eriocrania miners were correlated neither with the distance from the pollution source nor with the foliar content of heavy metals. However, larval masses of both solitary and gregarious species were highest at the zone of moderate pollution. The levels of total nitrogen and total carbon and the carbon: nitrogen ratio in birch leaves showed no clear patterns in relation to distance from the factory. Feeding efficiency of solitary larvae increased with higher foliar nitrogen content but decreased when the carbon: nitrogen ratio was high. Survival of solitary Eriocrania species was negatively correlated with total leaf carbon content. However, the population density of solitary miners showed a negative correlation with total foliar nitrogen and a positive correlation with the carbon:nitrogen ratio. Performance of the gregarious E. haworthi was not correlated with the chemical composition of birch leaves, whereas population density was highest close to the factory complex and correlated positively with the levels of copper and nickel in birch leaves. Consequently, pollution-induced changes in measured host plant chemicals were unlikely to affect population densities of Eriocrania miners via altered larval performance.