Polar day syndrome: differences in growth, photosynthetic traits and sink-size patterns between northern and southern Finnish silver birch (Betula pendula Roth) provenances in native and non-native photoperiods.

Continuous light (CL) is available throughout the polar day for plants in the Arctic during the growing season, whereas provenances of the same species experience a very different environment with non-CL (NCL) just a few latitudes to the south. Both provenances need to acclimate to climate warming, yet we lack comprehensive understanding of how their growth, photosynthesis and leaf traits differ. Further, the provenances presumably have morphological and physiological adaptations to their native environments and therefore differ in response to photoperiod. We tested the height growth, leaf longevity, biomass accumulation, biomass allocation and rates of gas exchange of northern (67°N) and southern (61°N) Finnish silver birch (Betula pendula Roth) origins in CL- and NCL-treatments in a 4-month chamber experiment. Irrespective of photoperiod, 67°N had higher area-based photosynthetic rate (Anet), stomatal conductance (gs) and relative height growth rate (RGR), but lower stomatal density and fewer branches and leaves than 61°N. Photoperiod affected height growth cessation, biomass and photosynthetic traits, whereas leaf longevity and many leaf functional traits remained unchanged. In CL, both provenances had lower gs, higher RGR, increased shoot:root ratio and increased sink sizes (more branching, more leaves, increased total plant dry weight) compared with NCL. In NCL, 67°N ceased height growth earlier than in CL, which altered biomass accumulation and distribution patterns. Northern conditions impose challenges for plant growth and physiology. Whether a provenance inhabits and is adapted to an area with or without CL can also affect its response to the changing climate. Northern birches may have adapted to CL and the short growing season with a 'polar day syndrome' of traits, including relatively high gas exchange rates with low leaf biomass and growth traits that are mainly limited by the environment and the earlier growth cessation (to avoid frost damage).

Strategic roadmap to assess forest vulnerability under air pollution and climate change.

Although it is an integral part of global change, most of the research addressing the effects of climate change on forests have overlooked the role of environmental pollution. Similarly, most studies investigating the effects of air pollutants on forests have generally neglected the impacts of climate change. We review the current knowledge on combined air pollution and climate change effects on global forest ecosystems and identify several key research priorities as a roadmap for the future. Specifically, we recommend (1) the establishment of much denser array of monitoring sites, particularly in the South Hemisphere; (2) further integration of ground and satellite monitoring; (3) generation of flux-based standards and critical levels taking into account the sensitivity of dominant forest tree species; (4) long-term monitoring of N, S, P cycles and base cations deposition together at global scale; (5) intensification of experimental studies, addressing the combined effects of different abiotic factors on forests by assuring a better representation of taxonomic and functional diversity across the ~73,000 tree species on Earth; (6) more experimental focus on phenomics and genomics; (7) improved knowledge on key processes regulating the dynamics of radionuclides in forest systems; and (8) development of models integrating air pollution and climate change data from long-term monitoring programs.

Climate and Competitive Status Modulate the Variation in Secondary Metabolites More in Leaves Than in Fine Roots of Betula pendula.

Plant secondary metabolites have many important functions; they also determine the productivity and resilience of trees under climate change. The effects of environmental factors on secondary metabolites are much better understood in above-ground than in below-ground part of the tree. Competition is a crucial biotic stress factor, but little is known about the interaction effect of climate and competition on the secondary chemistry of trees. Moreover, competition effect is usually overlooked when analyzing the sources of variation in the secondary chemistry. Our aim was to clarify the effects of competitive status, within-crown light environment, and climate on the secondary chemistry of silver birch (Betula pendula Roth). We sampled leaves (from upper and lower crown) and fine roots from competitively dominant and suppressed B. pendula trees in plantations along a latitudinal gradient (56-67° N) in Fennoscandia, with mean annual temperature (MAT) range: -1 to 8°C. Secondary metabolites in leaves (SML) and fine roots (SMFR) were determined with an HPLC-qTOF mass spectrometer. We found that SML content increased significantly with MAT. The effect of competitive stress on SML strengthened in colder climates (MAT<4°C). Competition and shade initiated a few similar responses in SML. SMFR varied less with MAT. Suppressed trees allocated relatively more resources to SML in warmer climates and to SMFR in colder ones. Our study revealed that the content and profile of secondary metabolites (mostly phenolic defense compounds and growth regulators) in leaves of B. pendula varied with climate and reflected the trees' defense requirements against herbivory, exposure to irradiance, and competitive status (resource supply). The metabolic profile of fine roots reflected, besides defense requirements, also different below-ground competition strategies in warmer and colder climates. An increase in carbon assimilation to secondary compounds can be expected at northern latitudes due to climate change.

Strategy by latitude? Higher photosynthetic capacity and root mass fraction in northern than southern silver birch (Betula pendula Roth) in uniform growing conditions.

Growth of northern trees is limited by short growing seasons. In multi-year trials, northern trees usually grow less than southern ones but can have higher gas exchange, whereas differences in biomass allocation and its relation to photosynthesis are less known. We characterized silver birch (Betula pendula Roth) provenances from southern (latitude 61°) and northern (latitude 67°) Finland in uniform chamber conditions. In a time-series experiment, we measured traits related to growth, biomass allocation and photosynthesis, and determined gas exchange responses to temperature and light. We found provenance differences in photosynthetic capacity and growth. The northern provenance allocated relatively more to roots, having a higher root mass fraction and lower shoot:root ratio than the southern provenance. On the other hand, the northern provenance had fewer leaves and lower total leaf dry weight (DW) than the southern provenance. The northern provenance attained higher rates of net photosynthesis (Anet) and higher stomatal conductance (gs) in all measured temperatures and higher photosynthesis at the optimum temperature (Aopt) than the southern provenance, but there was no difference in the optimum temperature of photosynthesis (Topt, 18.3 °C for the southern provenance vs 18.9 °C for the northern one). Photosynthetic light response curves showed no between-provenance differences. In a time-series, the northern provenance had higher Anet than the southern provenance, but gs was similar. The northern provenance had higher maximum quantum yield of photosystem II photochemistry (Fv/Fm) than the southern provenance. There were no differences between provenances in height, total plant DW, shoot DW, root DW or shoot mass fraction. Our results suggest that the provenances occupy a common thermal niche, or can at least relatively quickly acclimate to a common growth temperature. Thus, carbon assimilation of these northern trees may not be significantly affected by rising temperatures alone. In an equal photoperiod and optimal conditions, we found different one-season biomass accumulation strategies: southern trees grow with more leaves, while northern trees reach similar total assimilation (total DW, height) with more efficient photosynthetic capacity per leaf area (higher gas exchange, higher Fv/Fm) and relatively more investment in the below-ground fraction of the plant.

RPA-PCR couple: an approach to expedite plant diagnostics and overcome PCR inhibitors.

DNA extraction can be lengthy and sometimes ends up with amplification inhibitors. We present the potential of recombinase polymerase amplification (RPA) to replace plant DNA extraction. In our rapid 'RPA-PCR couple' concept, RPA is tuned to slower reaction kinetics to promote amplification of long targets. RPA primers amplify target and some flanking regions directly from simple plant macerates. Then PCR primers exponentially amplify the target directly from the RPA reaction. We present the coupling of RPA with conventional, TaqMan and SYBR Green PCR assays. We applied the concept to strawberry Phytophthora pathogens and the Phytophthora identification marker atp9-nad9. We found RPA-PCR couple specific, sensitive and reliable. The approach may also benefit other difficult samples such as food, feces and ancient samples.

Ozone affects plant, insect, and soil microbial communities: A threat to terrestrial ecosystems and biodiversity.

Elevated tropospheric ozone concentrations induce adverse effects in plants. We reviewed how ozone affects (i) the composition and diversity of plant communities by affecting key physiological traits; (ii) foliar chemistry and the emission of volatiles, thereby affecting plant-plant competition, plant-insect interactions, and the composition of insect communities; and (iii) plant-soil-microbe interactions and the composition of soil communities by disrupting plant litterfall and altering root exudation, soil enzymatic activities, decomposition, and nutrient cycling. The community composition of soil microbes is consequently changed, and alpha diversity is often reduced. The effects depend on the environment and vary across space and time. We suggest that Atlantic islands in the Northern Hemisphere, the Mediterranean Basin, equatorial Africa, Ethiopia, the Indian coastline, the Himalayan region, southern Asia, and Japan have high endemic richness at high ozone risk by 2100.

Insect herbivory dampens Subarctic birch forest C sink response to warming.

Climate warming is anticipated to make high latitude ecosystems stronger C sinks through increasing plant production. This effect might, however, be dampened by insect herbivores whose damage to plants at their background, non-outbreak densities may more than double under climate warming. Here, using an open-air warming experiment among Subarctic birch forest field layer vegetation, supplemented with birch plantlets, we show that a 2.3 °C air and 1.2 °C soil temperature increase can advance the growing season by 1-4 days, enhance soil N availability, leaf chlorophyll concentrations and plant growth up to 400%, 160% and 50% respectively, and lead up to 122% greater ecosystem CO2 uptake potential. However, comparable positive effects are also found when insect herbivory is reduced, and the effect of warming on C sink potential is intensified under reduced herbivory. Our results confirm the expected warming-induced increase in high latitude plant growth and CO2 uptake, but also reveal that herbivorous insects may significantly dampen the strengthening of the CO2 sink under climate warming.

Elevated temperature and ozone modify structural characteristics of silver birch (Betula pendula) leaves.

To study the effects of slightly elevated temperature and ozone (O3) on leaf structural characteristics of silver birch (Betula pendula Roth), saplings of four clonal genotypes of this species were exposed to elevated temperature (ambient air temperature +0.8-1.0 °C) and elevated O3 (1.3-1.4× ambient O3), alone and in combination, in an open-air exposure field over two growing seasons (2007 and 2008). So far, the impacts of moderate elevation of temperature or the combination of elevated temperature and O3 on leaf structure of silver birch have not been intensively studied, thus showing the urgent need for this type of studies. Elevated temperature significantly increased leaf size, reduced non-glandular trichome density, decreased epidermis thickness and increased plastoglobuli size in birch leaves during one or both growing seasons. During the second growing season, O3 elevation reduced leaf size, increased palisade layer thickness and decreased the number of plastoglobuli in spongy cells. Certain leaf structural changes observed under a single treatment of elevated temperature or O3, such as increase in the amount of chloroplasts or vacuole, were no longer detected at the combined treatment. Leaf structural responses to O3 and rising temperature may also depend on timing of the exposure during the plant and leaf development as indicated by the distinct changes in leaf structure along the experiment. Genotype-dependent cellular responses to the treatments were detected particularly in the palisade cells. Overall, this study showed that even a slight but realistic elevation in ambient temperature can notably modify leaf structure of silver birch saplings. Leaf structure, in turn, influences leaf function, thus potentially affecting acclimation capacity under changing climate.

Differences in growth and gas exchange between southern and northern provenances of silver birch (Betula pendula Roth) in northern Europe.

Due to its ubiquity across northern latitudes, silver birch (Betula pendula Roth) is an attractive model species for studying geographical trait variation and acclimation capacity. Six birch provenances from 60 to 67°N across Finland were grown in a common garden and studied for provenance and genotype variation. We looked for differences in height growth, photosynthetic gas exchange and chlorophyll content index (CCI) and compared the gas exchange of early and late leaves on short and long shoots, respectively. The provenances stratified into southern and northern groups. Northern provenances attained less height growth increment and had higher stomatal conductance (gs) and lower intrinsic water-use efficiency (WUE, Anet/gs) than southern provenances, whereas net photosynthesis (Anet) or CCI did not show clear grouping. Short shoot leaves had lower gs and higher WUE than long shoot leaves in all provenances, but there was no difference in Anet between shoot types. The separation of the provenances into two groups according to their physiological responses might reflect the evolutionary history of B. pendula. Latitudinal differences in gas exchange and water use traits can have plausible consequences for global carbon and water fluxes in a warming climate.

Low vapor pressure deficit reduces glandular trichome density and modifies the chemical composition of cuticular waxes in silver birch leaves.

Cuticular wax layer is the first barrier against the outside environment and the first defense encountered by herbivores and pathogens. The effects of environmental factors on cuticular chemistry, and on the formation of glandular trichomes that account for the storage and secretion of lipophilic compounds to the leaf surface are poorly understood. Low vapor pressure deficit (VPD) has shown to reduce the nitrogen (N) status of plants. Thus, we studied the effects of elevated air humidity, indicated as VPD, and the effect of N fertilization on cuticular waxes and glandular trichome density in silver birch (Betula pendula Roth). Experiments were carried out in growth chambers with juvenile plants and in a long-term field experiment with older trees. Low VPD reduced the glandular trichome density in both experiments, in chamber and in field. The contents of the major triterpenoid and flavonoid aglycones correlated positively with glandular trichome density, which supports the role of trichomes in the exudation of secondary compounds to the leaf surface. A closer examination of the cuticular wax chemistry in the chamber experiment revealed that low VPD and N supply affected the composition of cuticular waxes, but not the total wax content. The deposition of different wax compounds followed a co-ordinated pattern in birch leaves, but different compound groups varied in their responses to N fertilization and low VPD. Low VPD reduced the hydrophobicity of cuticular waxes, as demonstrated by lower alkane content and less hydrophobic flavonoid profile in low VPD than in high VPD. Reduced hydrophobicity of the wax layer is presumed to increase leaf wettability. Together with reduced trichome density in low VPD it may enhance the susceptibility of trees to fungal pathogens and herbivores. High N supply under low VPD reduced the effect of low VPD on the cuticular wax composition. Total fatty acid content and the expression of ß-amyrin synthase were lower under high N supply than under moderate N supply irrespective of VPD treatment. Nitrogen availability and decreasing VPD will modify leaf surface properties in silver birch and thereby affect tree defence against abiotic and biotic stress factors that emerge under climate change.

Artificially decreased vapour pressure deficit in field conditions modifies foliar metabolite profiles in birch and aspen.

Relative air humidity (RH) is expected to increase in northern Europe due to climate change. Increasing RH reduces the difference of water vapour pressure deficit (VPD) between the leaf and the atmosphere, and affects the gas exchange of plants. Little is known about the effects of decreased VPD on plant metabolism, especially under field conditions. This study was conducted to determine the effects of artificially decreased VPD on silver birch (Betula pendula Roth.) and hybrid aspen (Populus tremula L.×P. tremuloides Michx.) foliar metabolite and nutrient profiles in a unique free air humidity manipulation (FAHM) field experiment during the fourth season of humidity manipulation, in 2011. Long-term exposure to decreased VPD modified nutrient homeostasis in tree leaves, as demonstrated by a lower N concentration and N:P ratio in aspen leaves, and higher Na concentration and lower K:Na ratio in the leaves of both species in decreased VPD than in ambient VPD. Decreased VPD caused a shift in foliar metabolite profiles of both species, affecting primary and secondary metabolites. Metabolic adjustment to decreased VPD included elevated levels of starch and heptulose sugars, sorbitol, hemiterpenoid and phenolic glycosides, and α-tocopherol. High levels of carbon reserves, phenolic compounds, and antioxidants under decreased VPD may modify plant resistance to environmental stresses emerging under changing climate.

Low vapour pressure deficit affects nitrogen nutrition and foliar metabolites in silver birch.

Air humidity indicated as vapour pressure deficit (VPD) is directly related to transpiration and stomatal function of plants. We studied the effects of VPD and nitrogen (N) supply on leaf metabolites, plant growth, and mineral nutrition with young micropropagated silver birches (Betula pendula Roth.) in a growth chamber experiment. Plants that were grown under low VPD for 26 d had higher biomass, larger stem diameter, more leaves, fewer fallen leaves, and larger total leaf area than plants that were grown under high VPD. Initially, low VPD increased height growth rate and stomatal conductance; however, the effect was transient and the differences between low and high VPD plants became smaller with time. Metabolic adjustment to low VPD reflected N deficiency. The concentrations of N, iron, chlorophyll, amino acids, and soluble carbohydrates were lower and the levels of starch, quercetin glycosides, and raffinose were higher in the leaves that had developed under low VPD compared with high VPD. Additional N supply did not fully overcome the negative effect of low VPD on nutrient status but it diminished the effects of low VPD on leaf metabolism. Thus, with high N supply, the glutamine to glutamate ratio and starch production under low VPD became comparable with the levels under high VPD. The present study demonstrates that low VPD affects carbon and nutrient homeostasis and modifies N allocation of plants.

Searching for common responsive parameters for ozone tolerance in 18 rice cultivars in India: Results from ethylenediurea studies.

Eighteen rice (Oryza sativa) cultivars were screened for ozone (O3) tolerance and for the most responsive parameters with ethylenediurea (EDU) treatments at two experimental sites experiencing high ambient O3 conditions in the Indo-Gangetic Plains (IGP) of India. EDU was applied at 15 day intervals until the final harvest phase as a foliar spray at 300 ppm in order to protect the plants from the adverse effects of O3. Antioxidant activity, malondialdehyde content (MDA), chlorophyll content, gas exchange, and chlorophyll fluorescence (Fv/Fm) at the vegetative and flowering phases and harvest-related parameters were studied, for a total of 24 parameters. Seven of the studied cultivars had higher than average grainweightplant(-1) in all site and treatment combinations and can be recommended for cultivation in areas suffering from high O3 concentrations. The most responsive parameters with EDU treatment in high O3 across all cultivars were superoxide dismutase (SOD) and catalase (CAT) activities, the contents of oxidised (GSSG) and reduced (GSH) glutathione and MDA, and shoot weight plant(-1). These results indicated that the O3 scavenging activity of EDU is mediated through an antioxidant defence system rather than a direct effect on physiological parameters, such as photosynthesis and stomatal conductance.

Volatile organic compounds emitted from silver birch of different provenances across a latitudinal gradient in Finland.

Climate warming is having an impact on distribution, acclimation and defence capability of plants. We compared the emission rate and composition of volatile organic compounds (VOCs) from silver birch (Betula pendula (Roth)) provenances along a latitudinal gradient in a common garden experiment over the years 2012 and 2013. Micropropagated silver birch saplings from three provenances were acquired along a gradient of 7° latitude and planted at central (Joensuu 62°N) and northern (Kolari 67°N) sites. We collected VOCs emitted by shoots and assessed levels of herbivore damage of three genotypes of each provenance on three occasions at the central site and four occasions at the northern site. In 2012, trees of all provenances growing at the central site had higher total VOC emission rates than the same provenances growing at the northern site; in 2013 the reverse was true, thus indicating a variable effect of latitude. Trees of the southern provenance had lower VOC emission rates than trees of the central and northern provenances during both sampling years. However, northward or southward translocation itself had no significant effect on the total VOC emission rates, and no clear effect on insect herbivore damage. When VOC blend composition was studied, trees of all provenances usually emitted more green leaf volatiles at the northern site and more sesquiterpenes at the central site. The monoterpene composition of emissions from trees of the central provenance was distinct from that of the other provenances. In summary, provenance translocation did not have a clear effect in the short-term on VOC emissions and herbivory was not usually intense at the lower latitude. Our data did not support the hypothesis that trees growing at lower latitudes would experience more intense herbivory, and therefore allocate resources to chemical defence in the form of inducible VOC emissions.

Ozone affects ascorbate and glutathione biosynthesis as well as amino acid contents in three Euramerican poplar genotypes.

Ozone is an air pollutant that causes oxidative stress by generation of reactive oxygen species (ROS) within the leaf. The capacity to detoxify ROS and repair ROS-induced damage may contribute to ozone tolerance. Ascorbate and glutathione are known to be key players in detoxification. Ozone effects on their biosynthesis and on amino acid metabolism were investigated in three Euramerican poplar genotypes (Populus deltoides Bartr. × Populus nigra L.) differing in ozone sensitivity. Total ascorbate and glutathione contents were increased in response to ozone in all genotypes, with the most resistant genotype (Carpaccio) showing an increase of up to 70%. Reduced ascorbate (ASA) concentration at least doubled in the two most resistant genotypes (Carpaccio and Cima), whereas the most sensitive genotype (Robusta) seemed unable to regenerate ASA from oxidized ascorbate (DHA), leading to an increase of 80% of the oxidized form. Increased ascorbate (ASA + DHA) content correlated with the increase in gene expression in its biosynthetic pathway, especially the putative gene of GDP-l-galactose phosphorylase VTC2. Increased cysteine availability combined with increased expression of γ-glutamylcysteine synthetase (GSH1) and glutathione synthetase (GSH2) genes allows higher glutathione biosynthesis in response to ozone, particularly in Carpaccio. In addition, ozone caused a remobilization of amino acids with a decreased pool of total amino acids and an increase of Cys and putrescine, especially in Carpaccio. In addition, the expression of genes encoding threonine aldolase was strongly induced only in the most tolerant genotype, Carpaccio. Reduced ascorbate levels could partly explain the sensitivity to ozone for Robusta but not for Cima. Reduced ascorbate level alone is not sufficient to account for ozone tolerance in poplar, and it is necessary to consider several other factors including glutathione content.

Genetic and environmental determinants of insect herbivore community structure in a Betula pendula population.

A number of recent studies have shown that intraspecific genetic variation of plants may have a profound effect on the herbivorous communities which depend on them. However less is known about the relative importance of intraspecific variation compared to other ecological factors, for example environmental variation or the effects of herbivore damage. We randomly selected 22 Betula pendula genotypes from a local population (< 0.9 ha), cloned them and planted cloned seedlings on two study sites separated at a regional scale (distance between sites about 30 km) to examine an insect community of 23-27 species on these genotypes. B. pendula genotypes did not differ in their species richness, but the total mean abundance and the structure of the insect herbivore community was significantly affected by the genotype, which could account for up to 27% of the total variation in community structure. B. pendula genotype accounted for two to four times more variation in the arthropod community structure than did environmental (block) variation on a local scale, while on a regional scale, genotypic and environmental (site) variation accounted for 4-14% of the arthropod community structure. The genetic effects were modified by environmental variation on both a local and regional scale over one study year, and locally, the largest part of the variation (38%) could be explained by the genotype × environment (block) interactions. Suppression of insect herbivores during one growing season led to changed arthropod community structure in the following growing season, but this effect was minimal and could explain only 4% of the total variation in insect community structure. Our results suggest that both genetic and environmental factors are important determinants of the community structure of herbivorous insects. Together these mechanisms appear to maintain the high diversity of insects in B. pendula forest ecosystems.

Carbohydrate concentrations and freezing stress resistance of silver birch buds grown under elevated temperature and ozone.

The effects of slightly elevated temperature (+0.8 °C), ozone (O3) concentration (1.3 × ambient O3 concentration) and their combination on over-wintering buds of Betula pendula Roth were studied after two growing seasons of exposure in the field. Carbohydrate concentrations, freezing stress resistance (FSR), bud dry weight to fresh weight ratio, and transcript levels of cytochrome oxidase (COX), alternative oxidase (AOX) and dehydrin (LTI36) genes were studied in two clones (clones 12 and 25) in December. Elevated temperature increased the bud dry weight to fresh weight ratio and the ratio of raffinose family oligosaccharides to sucrose and the transcript levels of the dehydrin (LTI36) gene (in clone 12 only), but did not alter the FSR of the buds. Genotype-specific alterations in carbohydrate metabolism were found in the buds grown under elevated O3. The treatments did not significantly affect the transcript level of the COX or AOX genes. No clear pattern of an interactive effect between elevated temperature and O3 concentration was found. According to these data, the increase in autumnal temperatures and slightly increasing O3 concentrations do not increase the risk for freeze-induced damage in winter in silver birch buds, although some alterations in bud physiology occur.

Needle metabolome, freezing tolerance and gas exchange in Norway spruce seedlings exposed to elevated temperature and ozone concentration.

Northern forests are currently experiencing increasing mean temperatures, especially during autumn and spring. Consequently, alterations in carbon sequestration, leaf biochemical quality and freezing tolerance (FT) are likely to occur. The interactive effects of elevated temperature and ozone (O(3)), the most harmful phytotoxic air pollutant, on Norway spruce (Picea abies (L.) Karst.) seedlings were studied by analysing phenology, metabolite concentrations in the needles, FT and gas exchange. Sampling was performed in September and May. The seedlings were exposed to a year-round elevated temperature (+1.3 °C), and to 1.4× ambient O(3) concentration during the growing season in the field. Elevated temperature increased the concentrations of amino acids, organic acids of the citric acid cycle and some carbohydrates, and reduced the concentrations of phenolic compounds, some organic acids of the shikimic acid pathway, sucrose, cyclitols and steroids, depending on the timing of the sampling. Although growth onset occurred earlier at elevated temperature, the temperature of 50% lethality (LT(50)) was similar in the treatments. Photosynthesis and the ratio of photosynthesis to dark respiration were reduced by elevated temperature. Elevated concentrations of O(3) reduced the total concentration of soluble sugars, and tended to reduce LT(50) of the needles in September. These results show that alterations in needle chemical quality can be expected at elevated temperatures, but the seedlings' sensitivity to autumn and spring frosts is not altered. Elevated O(3) has the potential to disturb cold hardening of Norway spruce seedlings in autumn, and to alter the water balance of the seedling through changes in stomatal conductance (g(s)), while elevated temperature is likely to reduce g(s) and consequently reduce the O(3)-flux inside the leaves.

Interactive effects of elevated ozone and temperature on carbon allocation of silver birch (Betula pendula) genotypes in an open-air field exposure.

In the present experiment, the single and combined effects of elevated temperature and ozone (O(3)) on four silver birch genotypes (gt12, gt14, gt15 and gt25) were studied in an open-air field exposure design. Above- and below-ground biomass accumulation, stem growth and soil respiration were measured in 2008. In addition, a (13)C-labelling experiment was conducted with gt15 trees. After the second exposure season, elevated temperature increased silver birch above- and below-ground growth and soil respiration rates. However, some of these variables showed that the temperature effect was modified by tree genotype and prevailing O(3) level. For instance, in gt14 soil respiration was increased in elevated temperature alone (T) and in elevated O(3) and elevated temperature in combination (O(3) + T) treatments, but in other genotypes O(3) either partly (gt12) or totally nullified (gt25) temperature effects on soil respiration, or acted synergistically with temperature (gt15). Before leaf abscission, all genotypes had the largest leaf biomass in T and O(3) + T treatments, whereas at the end of the season temperature effects on leaf biomass depended on the prevailing O(3) level. Temperature increase thus delayed and O(3) accelerated leaf senescence, and in combination treatment O(3) reduced the temperature effect. Photosynthetic : non-photosynthetic tissue ratios (P : nP ratios) showed that elevated temperature increased foliage biomass relative to woody mass, particularly in gt14 and gt12, whereas O(3) and O(3) + T decreased it most clearly in gt25. O(3)-caused stem growth reductions were clearest in the fastest-growing gt14 and gt25, whereas mycorrhizal root growth and sporocarp production increased under O(3) in all genotypes. A labelling experiment showed that temperature increased tree total biomass and hence (13)C fixation in the foliage and roots and also label return was highest under elevated temperature. Ozone seemed to change tree (13)C allocation, as it decreased foliar (13)C excess amount, simultaneously increasing (13)C excess obtained from the soil. The present results suggest that warming has potential to increase silver birch growth and hence carbon (C) accumulation in tree biomass, but the final magnitude of this C sink strength is partly counteracted by temperature-induced increase in soil respiration rates and simultaneous O(3) stress. Silver birch populations' response to climate change will also largely depend on their genotype composition.