Library

Notes: The effects of elevated CO2 and temperature on the resource allocation pattern and resistance against mammalian herbivores of silver birch (Betula pendula Roth) were studied. Birch seedlings were grown through two growing seasons in closed-top chambers exposed to four different treatments: ambient CO2 and temperature, elevated atmospheric CO2 (700 ppm) and ambient temperature, elevated temperature (+3°C above ambient) and ambient CO2, and a combination of elevated CO2 and temperature. After winter hardening of the seedlings, the growth of the seedlings was measured and the concentration of secondary compounds such as phenolics and papyriferic acid determined. The top parts of the stem were fed to hares, and the basal parts of the same stems were offered to voles.Elevated CO2 increased the height and basal diameter of the shoots, shoot biomass and total biomass of the seedlings but did not have any effect on secondary chemistry. Elevated temperature increased the height and shoot biomass, but did not have a significant effect on the total biomass of the seedlings. Elevated temperature decreased the concentration of condensed tannins and their precursor, (+)-catechin, in the top part of the stems, but only the concentration of (+)-catechin in the basal part of the stems. There were no significant interactive effects between CO2 and temperature on phenolics in the stems, while the concentration of papyriferic acid showed significant interaction in the top part of the stems. This indicates high accumulation of papyriferic acid in ambient CO2 under increased temperature. Consequently, elevated temperature increased the resistance of birch against hares, but did not affect the resistance of the basal parts of the same birches to voles. Our results indicate that the predicted climatic change will not necessarily lead to increased browsing damage by the mountain hare and the field vole to silver birch.

Notes: 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.

Notes: Summary As ramets of the willow, Salix cinerea L. (Salicaceae) aged shoot length decreased in the six populations studied in S.E. Finland. Many traits correlated positively with shoot length: basal diameter, number of internodes, internode length, leaf size, and length of growing period. The bud-galling sawfly, Euura mucronata (Hartig) Man. (Churchill) (Hymenoptera: Tenthredinidae), responded positively to shoot length or correlated traits, and negatively to ramet age in three forest populations. This herbivore attacked the most vigorous plants in a population, and numbers of attacks declined as ramets aged and senesced. The generality of this kind of herbivore response to plant quality is emphasized.

Notes: Abstract The effect of CO2 enrichment (700 and 1050 ppm) on phytomass, soluble sugars, leaf nitrogen and secondary chemicals of three Salix myrsinifolia clones was studied in plants cultivated at very poor (sand seedlings) and moderate (peat seedlings) nutrient availability and under low illumination. The total shoot phytomass production of sand scedlings was less than 10% of that of the peat seedlings. Carbon dioxide increased the total shoot phytomass of peat seedlings. When the ambient carbon supply was doubled (to 700 ppm) the growth of sand seedlings was slightly enhanced but 1050 ppm CO2 gave growth figures similar to those at the control CO2 level. Leaf nitrogen content and total soluble sugar contents were significantly higher in peat seedlings than in sand seedlings. Leaf nitrogen showed a decreasing trend in relation to CO2 increase. On the other hand, CO2 did not have any clear-cut effect on total sugars. At the control CO2 level the content of salicortin, which is a dynamic phenolic, was higher in the peat seedlings than in the sand seedlings, but salicin showed the opposite trend. CO2 enrichment considerably decreased these phenolics in the peat seedlings. At the control CO2 level, the content of more static phenolics, such as proanthocyanidins, was higher in sand seedlings. An increased carbon supply considerably increased static phenolics in the peat seedlings. Willow defence against generalist herbivores is moderately decreased by enhancement of atmospheric carbon dioxide.

Notes: Summary Overwintering mountain hares (Lepus timidus) fed selectively on the shoots of a number of northern willow (Salix spp.) species. The hares preferred certain species over others and mature shoots over juvenile ones. There was a negative correlation between the phenolic glycoside concentration in the twigs and the hares' feeding. This correlation was substantiated by feeding experiments in which oat grains treated with purified phenoglycoside and catechin fractions of willow bark were offered along with untreated control oat grains to free-ranging mountain hares. Both fractions in concentrations normally found in willow twigs inhibited hare feeding. The results suggest that these phenolic compounds play a crucial role in the building up of resistance patterns among the willows. The decline in resistance in mature shoots of tall willows indicates that the juvenile resistance can be, perhaps secondarily, an adaptation against mammals browsing from ground level. Accordingly, low willow species retain a high level of resistance also in maturity.

Notes: Summary The bud-galling sawfly, Euura mucronata, attacked longer shoot length classes on its host, Salix cinerea, more frequently than shorter shoots. Shoot length accounted for 76 to 93 percent of the variance in number of galls per 100 shoots in three habitats: forest, watermeadow, and lakeside. The reasons for this pattern were addressed with studies on shoot length in relation to: 1. Number of resources (buds) per shoot; 2. Success in establishment of larvae in galls; 3. Gall size and resources per gall; and 4. Survival of larvae after establishment as influenced by plant resistance and natural enemy attack. The most important factors proved to be success in establishment of larvae, with percent of variance accounted for ranging from 57 to 77 percent in three of four sites where relationships were significant, and survival after establishment of larvae, with variance accounted for ranging from 40 to 54 percent in the same three sites. The pattern of survival was dictated by plant resistance and not by natural enemies. These two additive factors resulted in a general relationship across all sites of increasing emergence of fully developed larvae per cohort as shoot length increased, accounting for 78 percent of the variance. These adaptive advantages to attacking longer shoots are sufficient to account for the pattern of increased probability of shoots being attacked as they increase in length.

Notes: Summary 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.

Notes: Abstract The effects of individual phenolic glucosides and total glucoside fractions on the feeding behaviour of three willowfeeding leaf beetles (Coleoptera: Chrysomelidae) were tested in the laboratory. Feeding preferences of the tested leaf beetles were strongly influenced by certain phenolic glucosides which are typical secondary compounds of willows (Salicaceae:Salix). Two of the tested leaf beetles,Galerucella lineola andLochmaea capreae showed rather similar responses to glucoside treatments. Both of them were strongly stimulated by total glucoside fractions fromSalix triandra and by its major glucoside salidroside. The third species,Phatora vitellinae, was attracted most by the fractions fromS. myrsinifolia andS. pentandra, and by two related salicylate glucosides, tremulacin and salicortin. Food selection pattern of the tested beetles in the laboratory concords fairly well with their distributions in the field and with the occurrence of phenolic glucosides in their host willows. Phenolic glucoside extracts stimulated more feeding than individual pure glucosides. This indicates that different compounds have synergistic effects in the feeding behaviour of leaf beetles. Our results clearly show that willow leaf beetles select their food based on phenolic glucosides of their host plants.