We propose an alternative hypothesis to the handicap‐signalling hypothesis, to explain the high number of specialist aphids on tree species having bright autumn colour. Since birch aphids actively seek the first yellowing leaves for breeding in autumn, it is obvious that autumn colour of foliage does not repel migrating aphids. We suggest that aphids use bright colours as a cue to detect individual trees and leaves that are good sources of nitrogen in the form of amino acids in autumn. The active formation of bright‐coloured pigments in leaves is needed to protect them from photo inhibition during energy consuming nutrient retranslocation under cold autumn conditions. During nutrient export from leaves, nitrogen is in the form of amino acids in the sieve elements and easily available for aphids. Therefore, bright colours may act as a signal of easily available high‐quality food for viviparous aphid migrants that are selecting suitable trees for their sexual offspring reproduction. The females of sexual generation grown on the better quality food probably can oviposit the over‐wintering eggs to the twigs in higher numbers, which may have an adaptive advantage in competition with conspecific females.

Atmospheric change may affect plant phenolic compounds, which play an important part in plant survival. Therefore, we studied the impacts of CO 2 and O 3 on the accumulation of 27 phenolic compounds in the short-shoot leaves of two European silver birch (Betula pendula Roth) clones (clones 4 and 80). Seven-year-old soil-grown trees were exposed in open-top chambers over three growing seasons to ambient and twice ambient CO 2 and O 3 concentrations singly and in combination in central Finland.Elevated CO 2 increased the concentration of the phenolic acids (1 25%), myricetin glycosides (1 18%), catechin derivatives (1 13%) and soluble condensed tannins (1 19%) by increasing their accumulation in the leaves of the silver birch trees, but decreased the flavone aglycons (À7%) by growth dilution. Elevated O 3 increased the concentration of 3,4 0 -dihydroxypropiophenone 3-b-D-glucoside (1 22%), chlorogenic acid (1 19%) and flavone aglycons (1 4%) by inducing their accumulation possibly as a response to increased oxidative stress in the leaf cells. Nevertheless, this induction of antioxidant phenolic compounds did not seem to protect the birch leaves from detrimental O 3 effects on leaf weight and area, but may have even exacerbated them. On the other hand, elevated CO 2 did seem to protect the leaves from elevated O 3 because all the O 3 -derived effects on the leaf phenolics and traits were prevented by elevated CO 2 . The effects of the chamber and elevated CO 2 on some compounds changed over time in response to the changes in the leaf traits, which implies that the trees were acclimatizing to the altered environmental conditions. Although the two clones used possessed different composition and concentrations of phenolic compounds, which could be related to their different latitudinal origin and physiological characteristics, they responded similarly to the treatments. However, in some cases the variation in phenolic concentrations caused by genotype or chamber environment was much larger than the changes caused by either elevated CO 2 or O 3 .

We studied the effects of elevated concentrations of carbon dioxide ([CO2]) and ozone ([O3]) on growth, biomass allocation and leaf area of field-grown O3-tolerant (Clone 4) and O3-sensitive clones (Clone 80) of European silver birch (Betula pendula Roth) trees during 1999-2001. Seven-year-old trees of Clones 4 and 80 growing outside in open-top chambers were exposed for 3 years to the following treatments: outside control (OC); chamber control (CC); 2 x ambient [CO2] (EC); 2 x ambient [O3] (EO); and 2 x ambient [CO2] + 2 x ambient [O3] (EC+EO). When the results for the two clones were analyzed together, elevated [CO2] increased tree growth and biomass, but had no effect on biomass allocation. Total leaf area increased and leaf abscission was delayed in response to elevated [CO2]. Elevated [O3] decreased dry mass of roots and branches and mean leaf size and induced earlier leaf abscission in the autumn; otherwise, the effects of elevated [O3] were small across the clones. However, there were significant interactions between elevated [CO2] and elevated [O3]. When results for the clones were analyzed separately, stem diameter, volume growth and total biomass of Clone 80 were increased by elevated [CO2] and the stimulatory effects of elevated [CO2] on stem volume growth and total leaf area increased during the 3-year study. Clone 80 was unaffected by elevated [O3]. In Clone 4, elevated [O3] decreased root and branch biomass by 38 and 29%, respectively, whereas this clone showed few responses to elevated [CO2]. Elevated [CO2] significantly increased total leaf area in Clone 80 only, which may partly explain the smaller growth responses to elevated [CO2] of Clone 4 compared with Clone 80. Although we observed responses to elevated [O3], the responses to the EC+EO and EC treatments were similar, indicating that the trees only responded to elevated [O3] under ambient [CO2] conditions, perhaps reflecting a greater quantity of carbohydrates available for detoxification and repair in elevated [CO2].

The production of volatile organic compounds (VOCs) through the activation of different signaltransduction pathways may be induced in various biotic and abiotic stress situations having importance e.g. in insect and disease resistance. We compared the emission of VOCs emitted from silver birch Betula pendula Roth (clones 4 and 80) twigs damaged either by larvae of Epirrita autumnata, or infected with pathogenic leaf spot causing fungus Marssonina betulae. We also analysed whether local herbivore damage can systemically induce the release of VOCs from the undamaged top of same sapling. The emissions of methylsalicylate (MeSA), (Z)-ocimene, (E)-b-ocimene, (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT) and linalool were induced from the twigs after 72 h feeding damage by E. autumnata larvae. However, 48 h feeding damage did not induce rapid systemic release of VOCs from undamaged top leaves of the same twigs. Pathogen-infected birch twigs had significantly greater emission of (Z)-ocimene and (E)-b-ocimene than intact control twigs. The emission of DMNT was not significantly induced and MeSA was not found at all after pathogen infection, both being significantly different from herbivore damaged twigs. According to our results leaf fungal pathogen induces VOC emission profile differs from that of arthropod herbivore-damaged leaves, suggesting that birch is able to transmit parasite-specific information via VOC emissions to conspecifics and natural enemies of herbivores.

Long-term effects of elevated CO2 and O3 concentrations on gene expression in silver birch (Betula pendula Roth) leaves were studied during the end of the growing season. Two birch genotypes, clones 4 and 80, with different ozone growth responses, were exposed to 2¥ ambient CO2 and/or O3 in open-top chambers (OTCs). Microarray analyses were performed after 2 years of exposure, and the transcriptional profiles were compared to key physiological characteristics during leaf senescence. There were genotypic differences in the responses to CO2 and O3. Clone 80 exhibited greater transcriptional response and capacity to alter metabolism, resulting in better stress tolerance. The gene expression patterns of birch leaves indicated contrasting responses of senescence-related genes to elevated CO2 and O3. Elevated CO2 delayed leaf senescence and reduced associated transcriptional changes, whereas elevated O3 advanced leaf senescence because of increased oxidative stress. The combined treatment demonstrated that elevated CO2 only temporarily alleviated the negative effects of O3. Gene expression data alone were insufficient to explain the O3 response in birch, and additional physiological and biochemical data were required to understand the true O3 sensitivity of these clones.

The effect of atmospheric change on birch aphid (Euceraphis betulae Koch) oviposition preference was examined and plant characteristics that are possibly responsible for the observed effects were investigated. It was hypothesized that the increasing concentrations of CO 2 and O 3 affect singly or in combination the oviposition of birch aphids via changes in host plant characteristics. Two genotypes of field-growing silver birch (Betula pendula Roth) trees (clones 4 and 80), which were exposed to doubled ambient concentration of CO 2 and O 3 , singly and in combination, in a 3-year open-top chamber experiment, were used in an aphid oviposition preference test. It was found that elevated CO 2 , irrespective of ozone concentration, increased the number of aphid eggs laid on clone 4, but not in clone 80. Several flavonoid aglycones were identified from the exudate coating of birch buds. Although elevated CO 2 and O 3 affected these phenolic compounds in clone 4, the effects did not correlate with the observed changes in aphid oviposition. It is suggested that neither bud length, which was not affected by the treatments, nor surface exudate phenolics mediate birch aphid oviposition preference.

Seasonal changes in growth, photosynthesis, and related biochemical properties and leaf structure were determined for two clones (4 and 80, 20 trees per clone) of 7-year-old Betula pendula Roth trees during the growing season of 1998. Differences between the two genotypes were determined to characterize the physiological traits that might affect growth and productivity and that might differ between the genotypes. Net photosynthesis of the short shoot leaves varied between 11 and 15 µmol·m–2·s–1 and decreased only slightly towards the end of the summer. However, our results showed more marked decreases in the amount of Rubisco (ribulose biphosphate carboxylase/oxygenase) and leaf N and increases in the total leaf, palisade and spongy layer thickness, chloroplast and starch grain size, and diameter of plastoglobuli in both clones in response to leaf ageing and changes in growth environment. Height and biomass were greater in clone 80 than in clone 4. This was related to slightly more efficient net photosynthesis and higher stomatal conductance and density as well as higher activity of Rubisco and content of foliar nutrients (other than N). We conclude that clone 80 is characterized by faster gas exchange, higher Rubisco activity, stomatal conductance, and density, and earlier leaf ageing, which may be related to the higher ozone sensitivity determined previously in pot experiments with younger saplings.

Ozone sensitivity of silver birch ( BETULA PENDULA Roth) has been thoroughly investigated since early 1990's in Finland. In our long-term open-field experiments the annual percentage reduction in basal diameter and stem volume increment were the best non-destructive growth indicators for ozone impact when plotted against AOTX. Remarkable differences in defence strategies, stomatal conductance, and defence compounds (phenolics), clearly indicate that external exposure indices are ineffective for accurate risk assessment for birch. For flux-based approaches, site-specific values for G(max) and G(dark) are necessary, and determinants for detoxification capacity, ageing of leaves, and cumulative ozone impact would be needed for further model development. Increasing CO(2) seems to counteract negative ozone responses in birch, whereas exposure to spring time frost may seriously exacerbate ozone damage in northern conditions. Therefore, we need to proceed towards incorporating the most important climate change factors in any attempts for ozone risk assessment.