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The impact of atmospheric pollution on the growth of European forest tree species, particularly European beech, Silver fir and Norway spruce, is examined in five mesic forests in the Czech Republic. Analyzing of basal area increment (BAI) patterns using linear mixed effect models reveals a complex interplay between atmospheric nitrogen (N) and sulphur (S) deposition, climatic variables and changing CO 2 concentrations. Beech BAI responds positively to N deposition (in tandem with air CO 2 concentration), with soil phosphorus (P) availability emerging as a significant factor influencing overall growth rates. Fir BAI, on the other hand, was particularly negatively influenced by S deposition, although recent growth acceleration suggests growth resilience in post-pollution period. This fir growth surge likely coincides with stimulation of P acquisition following the decline of acidic pollution. The consequence is the current highest productivity among the studied tree species. The growth dynamics of both conifers were closely linked to the stoichiometric imbalance of phosphorus in needles, indicating the possible sensitivity of exogenous controls on nutrient uptake. Furthermore, spruce BAI was positively linked to calcium availability across sites. Despite enhanced water-use efficiency under elevated CO 2 , spruce growth is constrained by precipitation deficit and demonstrates weakening resilience to increasing growing season air temperatures. Overall, these findings underscore the intricate relationships between atmospheric pollution, nutrient availability, and climatic factors in shaping the growth dynamics of European forest ecosystems. Thus, incorporating biogeochemical context of nutrient availability is essential for realistic modelling of tree growth in a changing climate., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Tropospheric ozone (O 3 ) causes widespread damage to vegetation; however, monitoring of O 3 induced damage is often reliant on manual leaf inspection. Reflectance spectroscopy of vegetation can identify and detect unique spectral signatures of different abiotic and biotic stressors. In this study, we tested the use of hyperspectral leaf reflectance to detect O 3 stress in alder, beech, birch, crab apple, and oak saplings exposed to five long-term O 3 regimes (ranging from daily target maxima of 30 ppb O 3 to 110 ppb). Hyperspectral reflectance varied significantly between O 3 treatments, both in whole spectra analysis and when simplified to representative components. O 3 damage had a multivariate impact on leaf reflectance, underpinned by changes in pigment balance, water content and structural composition. Vegetation indices derived from reflectance which characterised the visible green peak were able to differentiate between O 3 treatments. Iterative normalised difference spectral indices across the hyperspectral wavelength range were correlated to visual damage scores to identify significant wavelengths for O 3 damage detection. We propose a new Ozone Damage Index (OzDI), which characterises the reflectance peak in the shortwave infrared region and outperformed existing vegetation indices in terms of correlation to O 3 treatment. These results demonstrate the potential application of hyperspectral reflectance as a high throughput method of O 3 damage detection in a range of common broadleaf. species., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Background: Climate change is leading to severe and long-term droughts in European forest ecosystems. can have profound effects on various physiological processes, including photosynthesis, gene expression patterns, and nutrient uptake at the developmental stage of young trees. Objectives: Our study aimed to test the hypothesis that the application of silica (SiO 2 ) influences photosynthetic efficiency and gene expression in 1- to 2-year-old Fagus sylvatica (L.) seedlings. Additionally, we aimed to assess whether silicon application positively influences the structural properties of leaves and roots. To determine whether the plant physiological responses are genotype-specific, seedlings of four geographically different provenances were subjected to a one-year evaluation under greenhouse conditions. Methods: We used the Kruskal-Wallis test followed by Wilcoxon's test to evaluate the differences in silicon content and ANOVA followed by Tukey's test to evaluate the physiological responses of seedlings depending on treatment and provenance. Results: Our results showed a significantly higher Si content in the roots compared with the leaves, regardless of provenance and treatment. The most significant differences in photosynthetic performance were found in trees exposed to Si treatment, but the physiological responses were generally nuanced and provenance-dependent. Expression of hsp 70 and hsp 90 was also increased in leaf tissues of all provenances. These results provide practical insights that Si can improve the overall health and resilience of beech seedlings in nursery and forest ecosystems, with possible differences in the beneficial role of silicon application arising from the large differences in wild populations of forest tree species.

Microplastics have the capacity to accumulate in soil due to their high resistance to degradation, consequently altering soil properties and influencing plant growth. This study focused on assessing the impact of various types and doses of microplastics on beech seedling growth. In our experiment, we used polypropylene and styrene granules with diameter of 4.0 mm in quantities of 2.5% and 7%. The hypothesis was that microplastics significantly affect seedlings' nutritional status and growth characteristics. The research analysed seedlings' nutrition, root morphological features, above-ground growth, and enzymatic activity in the substrate. Results confirmed the importance of microplastics in shaping the nutritional status of young beech trees. Microplastic type significantly impacted N/P and Ca/Mg stoichiometry, while microplastic quantity influenced Ca/Al and Ca + K + Mg/Al stoichiometry. Notably, only in the case of root diameter were significantly thicker roots noted in the control variant, whereas microplastics played a role in shaping the leaves' characteristics of the species studied. The leaf area was significantly larger in the control variant compared to the variant with polypropylene in the amount of 2.5% and styrene in the amount of 7%. Additionally, the study indicates a significant impact of microplastics on enzyme activity. In the case of CB and SP, the activity was twice as high in the control variant compared to the variants with microplastics. In the case of BG, the activity in the control variant was higher in relation to the variants used in the experiment. Research on the impact of microplastics on the growth of beech seedlings is crucial for enhancing our understanding of the effects of environmental pollution on forest ecosystems. Such studies are integral in shaping forestry management practices and fostering a broader public understanding of the ecological implications of plastic pollution., (© 2024. The Author(s).)

Research into trees plays a very important role in evaluations of soil contamination with diesel oil. Trees are ideal for reclaiming contaminated soils because their large biomass renders them more resistant to higher concentrations of pollutants. In the literature, there is a general scarcity of long-term studies performed on trees, in particular European beeches. The aim of this study was to evaluate the responses of Scots pines and European beeches grown for 8 years on soil contaminated with diesel oil. Selected morphological and physiological parameters of trees were analyzed. The biomass yield of Scots pines was not significantly correlated with increasing concentrations of diesel oil, but it was more than 700% higher than in European beeches. Scots pines were taller and had a larger stem diameter than European beeches during the 8-year study. The diameter of trees grown on the most contaminated soil was reduced 1.5-fold in Scots pines and more than twofold in European beeches. The length of Scots pine needles from the most contaminated treatment decreased by 50% relative to control needles. The shortest needles were heaviest. The fluctuating asymmetry (FA) of needle length was highest in Scots pines grown on the most contaminated soil, whereas the reverse was noted in the FA of needle weight. Diesel oil decreased the concentrations of chlorophylls a and b, total chlorophyll, and carotenoids. The Fv/Fm ratio of needles and leaves was influenced by the tested concentrations of diesel oil. The results of the study indicate that the Scots pine better adapts (grows more rapidly and produces higher biomass) to long-term soil contamination with diesel oil than the European beech. In European beeches, growth inhibition and leaf discoloration (a decrease in chlorophyll content) were observed already after the first year of the experiment, which indicates that 1-year-old seedlings of European beech are robust bioindicators of soil contamination with diesel oil.

To test the hypothesis that ultraviolet radiation (UV) modulates photosynthetic responses to elevated CO 2 concentration ([CO 2 ]) in plants, saplings of European beech were grown for two vegetation seasons under ambient (400 ppm) and elevated (700 ppm) atmospheric [CO 2 ]. From April to November the saplings were exposed to (i) ambient UV radiation, (ii) excluded and (iii) enhanced UV (150% of ambient). Gas-exchange and chlorophyll fluorescence techniques were used throughout the second vegetation season together with biochemical analyses of the amount and activity of the Rubisco enzyme. We found support for the hypothesis that an impact of elevated [CO 2 ] on photosynthesis is substantially modulated by UV radiation. Moreover, we found that the [CO 2 ] × UV interaction is changing along the vegetation season: an enhanced UV radiation stimulated a positive effect of elevated [CO 2 ] on plant photosynthesis at the beginning of the vegetation season (short-term effect), whilst long-term cultivation reduced the stimulatory effect of elevated [CO 2 ] (a clear down-regulation of photosynthesis). Down-regulation was, however, not found in plants grown under the conditions of excluded UV radiation. We found evidence that the down-regulation of photosynthesis is associated with a complex acclimation at different hierarchical and functional levels, including an acclimation of primary photochemical reactions, carboxylation activity of Rubisco enzyme, and stomatal conductance., (Copyright © 2018 Elsevier Masson SAS. All rights reserved.)

Under the conditions of ongoing climate change, terrestrial ecosystems will be simultaneously exposed to a permanent rise in atmospheric CO 2 concentration and increasing variability of such environmental factors as temperature, precipitation, and UV radiation. This will result in numerous interactions. The interactive effects caused by exposure to such multiple environmental factors are not yet well understood. We tested the hypotheses that enhanced UV radiation reduces the stimulatory effect of elevated CO 2 concentration on plant biomass production and that it alters biomass allocation in broadleaved European beech (Fagus sylvatica L.) saplings. Our results after 2 years of exposure confirmed interactive effects of CO 2 concentration and UV radiation on biomass production, and particularly on biomass allocation to roots and aboveground biomass. The strongest stimulatory effect of elevated CO 2 on aboveground biomass and roots was found under ambient UV radiation, while both low and high UV doses reduced this stimulation. Nitrogen content in the roots and the distribution of nitrogen among leaves and roots were also significantly affected by interaction of CO 2 concentration and UV radiation. The observed changes in leaf and root C:N stoichiometry were associated with altered morphological traits, and particularly with a change in the proportion of fine roots. As the biomass allocation and especially the proportion of fine roots can play an important role in effective water and nutrient use and acclimation to future climates, it is essential to obtain a deeper understanding of the links between C:N stoichiometry and biomass accumulation., (Copyright © 2018 Elsevier Masson SAS. All rights reserved.)

The primary objective of this study was to describe parameters related to the leaf biochemical assimilation capacity of photosynthesis, such as the maximum rates of carboxylation (V cmax ) and electron transport (J max ), as a function of the leaf nitrogen content throughout a canopy of Siebold's beech and Japanese oak grown under elevated ozone (O 3 ) conditions during a growing season. To this end, we investigated the relationship between photosynthetic traits and leaf nitrogen content in various canopy positions of two tree species under free-air O 3 exposure (60 nmol mol -1 , during daylight hours) in June, August, and October 2012. We observed O 3 -induced reduction in V cmax and J max without reduction of leaf nitrogen content in both tree species. In Siebold's beech, V cmax and J max in leaves with higher N area were largely decreased by O 3 from August, while little effect of O 3 was observed in leaves with lower N area . On the other hand, there was no difference in the extent of O 3 -induced reduction in V cmax and J max across the range of N area in leaves of Japanese oak. Reduction of leaf nitrogen content under elevated O 3 conditions was observed only in Siebold's beech in October. These results indicated that the decrease in the efficiency of photosynthetic nitrogen use is in an earlier step in O 3 -induced decline of photosynthesis in Siebold's beech and Japanese oak. Based on these results, we emphasize the importance of integration of O 3 effects into the conventional estimation of V cmax and J max from leaf nitrogen content for evaluating canopy photosynthesis under current and future elevated O 3 conditions., (Copyright © 2018. Published by Elsevier B.V.)

The aim of this study was to determine the content, distribution and behaviour of Al in soils under beech forest with different parent rock, and to assess the role of herbaceous vegetation on soil Al behaviour. We hypothesize that the contents of elements in the soil sorption complex (Al etc.) are strongly influenced by vegetation cover. Also, low molecular mass organic acids (LMMOA) can be considered as an indicator of soil organic matter (SOM) decomposition and vegetation litter turnover. Speciation of LMMOA, nutrition content (PO 4 3- , Ca 2+ , K + ) and element composition in aqueous extracts were determined by means of ion chromatography and inductively coupled plasma - optical emission spectrometry (ICP-OES) respectively. Active and exchangeable pH, sorption characteristics and exchangeable Al (Al ex ) were determined in BaCl 2 extracts by ICP-OES. Elemental composition of parent rocks was assessed by means of X-ray fluorescence spectroscopy. Herb-poor localities showed lower pH, less nutrients (PO 4 3- , Ca 2+ , K + ), less LMMOA, a larger stock of SOM and greater cation exchange capacity. There was also lower mobilisation of Al in organic horizons, which explains the larger pools of Al. Generally, we can conclude that LMMOA, and thus soil vegetation cover, play an important role in the Al soil cycle., (Copyright © 2017 Elsevier Inc. All rights reserved.)

A unique database of stand volume growth, estimated as periodic annual volume increment (in m 3  ha -1 per year over the period 2001-2005) from 728 European beech (Fagus sylvatica L.) sites distributed across Italy, was used to assess the effects of ambient ozone (O 3 ), expressed as annual average (M24), accumulated exposure above a 40 ppb hourly threshold (AOT40), and total stomatal ozone flux (POD0). Growth data were from the National forest inventory of Italy, while climate data and ozone concentrations were computed by the WRF and CHIMERE models, respectively. Results show that the growth increased with increasing solar radiation and air temperature and decreased with increasing number of cold days, while effects of soil water content and O 3 were not significant. In contrast, the literature results suggest that European beech is sensitive to both drought and O 3 . Ozone levels resulted to be very high (48 ppb M24, 51,200 ppb h AOT40, 21.08 mmol m -2 POD0, on average) and thus able to potentially affect European beech growth. We hypothesize that the high-frequency signals of soil water and O 3 got lost when averaged over 5 years and recommended finer time-resolution investigations and inclusion of other factors of variability, e.g., thinning, tree age, and size.

Global change affects the functioning of forest ecosystems and the services they provide, but little is known about the interactive effects of co-occurring global change drivers on important functions such as tree growth and vitality. In the present study we quantified the interactive (i.e. synergistic or antagonistic) effects of atmospheric nitrogen (N) deposition and climatic variables (temperature, precipitation) on tree growth (in terms of tree-ring width, TRW), taking forest ecosystems with European beech (Fagus sylvatica L.) as an example. We hypothesised that (i) N deposition and climatic variables can evoke non-additive responses of the radial increment of beech trees, and (ii) N loads have the potential to strengthen the trees' sensitivity to climate change. In young stands, we found a synergistic positive effect of N deposition and annual mean temperature on TRW, possibly linked to the alleviation of an N shortage in young stands. In mature stands, however, high N deposition significantly increased the trees' sensitivity to increasing annual mean temperatures (antagonistic effect on TRW), possibly due to increased fine root dieback, decreasing mycorrhizal colonization or shifts in biomass allocation patterns (aboveground vs. belowground). Accordingly, N deposition and climatic variables caused both synergistic and antagonistic effects on the radial increment of beech trees, depending on tree age and stand characteristics. Hence, the nature of interactions could mediate the long-term effects of global change drivers (including N deposition) on forest carbon sequestration. In conclusion, our findings illustrate that interaction processes between climatic variables and N deposition are complex and have the potential to impair growth and performance of European beech. This in turn emphasises the importance of multiple-factor studies to foster an integrated understanding and models aiming at improved projections of tree growth responses to co-occurring drivers of global change., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

We aimed to clarify the effects of ozone (O 3 ) on photosynthetic ability of upper and lower canopy leaves of Fagus crenata Blume seedlings grown under different soil nutrient conditions. To accomplish this objective, we analyzed the response of photosynthetic parameters such as maximum carboxylation rate (V cmax ) to cumulative stomatal O 3 uptake (ΣF st ) and reduction rate of V cmax per unit ΣF st as an index of detoxification capacity for O 3 . The seedlings of Fagus crenata were grown for two growing seasons (2014-2015) in nine treatments comprised of a combination of three levels of gas treatments (charcoal-filtered air or 1.0- or 1.5-times ambient O 3 concentration) and three levels of soil nutrient treatments (non-fertilized or a supply of relatively low or high concentrations of compound fertilizer). The nutrient supply significantly increased the degree of O 3 -induced reduction in V cmax in September. However, nutrient supply did not significantly increase ΣF st and reduce the detoxification capacity for O 3 . On the other hand, the degree of O 3 -induced reduction in V cmax of upper canopy leaves was higher as compared with that of lower canopy leaves in August due to the higher ΣF st . However, the reduction rate of V cmax per unit ΣF st in lower canopy leaves was higher than that in upper canopy leaves, indicating lower detoxification capacity for O 3 in lower canopy leaves. Reduction rate of V cmax per unit ΣF st over the threshold, which is assumed to be proportional to gross photosynthetic rate, was similar between upper and lower canopy leaves. Therefore, capacity of photosynthetic CO 2 assimilation is likely to be associated with detoxification capacity for O 3 in upper and lower canopy leaves of F. crenata seedlings grown under different soil nutrient conditions., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

This work evaluates the possibility of using beech wood (Fagus sylvatica) dilute-acid (H 2 SO 4 ) hydrolysate as a feedstock for Chlorella sorokiniana growth, fatty acid and pigment production. Neutralized wood acid hydrolysate, containing organic and mineral compounds, was tested on Chlorella growth at different concentrations and compared to growth under phototrophic conditions. Chlorella growth was improved at lower loadings and inhibited at higher loadings. Based on these results, a 12% neutralized wood acid hydrolysate (Hyd12%) loading was selected to investigate its impact on Chlorella growth, fatty acid and pigment production. Hyd12% improved microalgal biomass, fatty acid and pigment productivities both in light and in dark, when compared to photoautotrophic control. Light intensity had substantial influence on fatty acid and pigment composition in Chlorella culture during Hyd12%-based growth. Moreover, heterotrophic Chlorella cultivation with Hyd12% also showed that wood hydrolysate can constitute an attractive feedstock for microalgae cultivation in case of lack of light., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

This study investigated microwave-assisted formic acid (MW-FA) pretreatment as a possible way to improve aromatic production from catalytic fast pyrolysis (CFP) of lignocellulosic biomass. Results showed that short duration of MW-FA pretreatment (5-10min) could effectively disrupt the recalcitrant structure of beech wood and selectively remove its hemicellulose and lignin components. This increased the accessibility of cellulose component of biomass to subsequent thermal conversion in CFP. Consequently, the MW-FA pretreated beech wood produced 14.0-28.3% higher yields (26.4-29.8C%) for valuable aromatic products in CFP than the untreated control (23.2C%). In addition, the yields of undesired solid residue (char/coke) decreased from 33.1C% for the untreated control to 28.6-29.8C% for the MW-FA pretreated samples. These results demonstrate that MW-FA pretreatment can provide an effective way to improve the product distribution from CFP of lignocellulose., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

The purpose of this study was to obtain basic information on acclimation capacity of photosynthesis in Siebold's beech seedlings to increasing light intensity under future elevated CO2 conditions. We monitored leaf photosynthetic traits of these seedlings in changing light conditions (before removal of shade trees, the year after removal of shade trees and after acclimation to open conditions) in a 10-year free air CO2 enrichment experiment in northern Japan. Elevated CO2 did not affect photosynthetic traits such as leaf mass per area, nitrogen content and biochemical photosynthetic capacity of chloroplasts (i.e. maximum rate of carboxylation and maximum rate of electron transport) before removal of the shade trees and after acclimation to open conditions; in fact, a higher net photosynthetic rate was maintained under elevated CO2 . However, in the year after removal of the shade trees, there was no increase in photosynthesis rate under elevated CO2 conditions. This was not due to photoinhibition. In ambient CO2 conditions, leaf mass per area and nitrogen content were higher in the year after removal of shade trees than before, whereas there was no increase under elevated CO2 conditions. These results indicate that elevated CO2 delays the acclimation of photosynthetic traits of Siebold's beech seedlings to increasing light intensity., (© 2015 German Botanical Society and The Royal Botanical Society of the Netherlands.)

Rhizospheric nitric oxide (NO) and carbon dioxide (CO2) are signalling compounds known to affect physiological processes in plants. Their joint influence on tree nitrogen (N) nutrition, however, is still unknown. Therefore, this study investigated, for the first time, the combined effect of rhizospheric NO and CO2 levels on N uptake and N pools in European beech (Fagus sylvatica L.) seedlings depending on N availability. For this purpose, roots of seedlings were exposed to one of the nine combinations (i.e., low, ambient, high NO plus CO2 concentration) at either low or high N availability. Our results indicate a significant effect of rhizospheric NO and/or CO2 concentration on organic and inorganic N uptake. However, this effect depends strongly on NO and CO2 concentration, N availability and N source. Similarly, allocation of N to different N pools in the fine roots of beech seedlings also shifted with varying rhizospheric gas concentrations and N availability., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Soil microbial communities play an important role in forest ecosystem functioning, but how climate change will affect the community composition and consequently bacterial functions is poorly understood. We assessed the effects of reduced precipitation with the aim of simulating realistic future drought conditions for one growing season on the bacterial community and its relation to soil properties and forest management. We manipulated precipitation in beech and conifer forest plots managed at different levels of intensity in three different regions across Germany. The precipitation reduction decreased soil water content across the growing season by between 2 to 8% depending on plot and region. T-RFLP analysis and pyrosequencing of the 16S rRNA gene were used to study the total soil bacterial community and its active members after six months of precipitation reduction. The effect of reduced precipitation on the total bacterial community structure was negligible while significant effects could be observed for the active bacteria. However, the effect was secondary to the stronger influence of specific soil characteristics across the three regions and management selection of overstorey tree species and their respective understorey vegetation. The impact of reduced precipitation differed between the studied plots; however, we could not determine the particular parameters being able to modify the response of the active bacterial community among plots. We conclude that the moderate drought induced by the precipitation manipulation treatment started to affect the active but not the total bacterial community, which points to an adequate resistance of the soil microbial system over one growing season.

Allocation of recent photoassimilates of juvenile beech and spruce in response to twice-ambient ozone (2 × O(3)) and plant competition (i.e. intra vs. inter-specific) was examined in a phytotron study. To this end, we employed continuous (13)CO(2)/(12)CO(2) labeling during late summer and pursued tracer kinetics in CO(2) released from stems. In beech, allocation of recent photoassimilates to stems was significantly lowered under 2 × O(3) and increased in spruce when grown in mixed culture. As total tree biomass was not yet affected by the treatments, C allocation reflected incipient tree responses providing the mechanistic basis for biomass partitioning as observed in longer experiments. Compartmental modeling characterized functional properties of substrate pools supplying respiratory C demand. Respiration of spruce appeared to be exclusively supplied by recent photoassimilates. In beech, older C, putatively located in stem parenchyma cells, was a major source of respiratory substrate, reflecting the fundamental anatomical disparity between angiosperm beech and gymnosperm spruce., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

The effect of long-term exposure of twice-ambient O(3) (2 × O(3)) on whole-tree nitrogen (N) uptake and partitioning of adult beech and spruce was studied in a mixed forest stand, SE-Germany. N uptake as (15)N tracer and N pools were calculated using N concentrations and biomass of tree compartments. Whole-tree N uptake tended to be lower under 2 × O(3) in both species compared to trees under ambient O(3) (1 × O(3)). Internal partitioning in beech showed significantly higher allocation of new N to roots, with mycorrhizal root tips and fine roots together receiving about 17% of new N (2 × O(3)) versus 7% (1 × O(3)). Conversely, in spruce, N allocation to roots was decreased under 2 × O(3). These contrasting effects on belowground N partitioning and pool sizes, being largely consistent with the pattern of N concentrations, suggest enhanced N demand and consumption of stored N with higher relevance for tree-internal N cycling in beech than in spruce., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Stomata tend to narrow under ozone (O(3)) impact, leading to limitation of stomatal O(3) influx. Here, we review stomatal response under recently conducted free-air O(3) exposure experiments on two species of the same tree genus: Fagus sylvatica at Kranzberg Forest (Germany) and F. crenata at Sapporo Experimental Forest (Japan). Both beeches exhibited reduction in stomatal conductance (gs) by 10-20% under experimentally enhanced O(3) regimes throughout the summer relative to ambient-air controls. Stomatal narrowing occurred, in early summer, in the absence of reduced carboxylation capacity of Rubisco, although photosynthetic net CO(2) uptake rate temporarily reflected restriction to some minor extent. Observed stomatal narrowing was, however, diminished in autumn, suggesting gradual loss of stomatal regulation by O(3). Monotonic decline in gs with cumulative O(3) exposure or flux in current modeling concepts appear to be unrealistic in beech., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

To construct stomatal conductance models and estimate stomatal O3 uptake for Fagus crenata, Quercus serrata, Quercus mongolica var. crispula and Betula platyphylla, stomatal conductance (gs) was measured in seedlings of the four tree species. Better estimates of gs were made by incorporating the acute effects of O3 on gs into the models and the models could explain 34-52% of the variability in gs. Although the O3 concentration was relatively high in spring from April to May, COU of F. crenata, Q. serrata and Q. mongolica var. crispula were relatively low and the ratios of COU in spring to total COU in one year were 16.8% in all tree species because of low gs limited mainly by leaf pre-maturation and/or low temperature. The COU of B. platyphylla were relatively high mainly because of rapid leaf maturation and lower optimal temperature for stomatal opening., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

The estimate of growth losses by ozone exposure of forest trees is a significant part in current C sequestration calculations and will also be important in future modeling. It is therefore important to know if the relationship between ozone flux and growth reduction of young trees, used to derive a Critical Level for ozone, is also valid for mature trees. Epidemiological analysis of stem increment data from Fagus sylvatica L. and Picea abies Karst. observed in Swiss forest plots was used to test this hypothesis. The results confirm the validity of the flux-response relationship at least for beech and therefore enable estimating forest growth losses by ozone on a country-wide scale. For Switzerland, these estimates amount to 19.5% growth reduction for deciduous forests, 6.6% for coniferous forests and 11.0% for all forested areas based on annual ozone stomatal uptake during the time period 1991-2011., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Background and Aims: Stomatal density (SD) generally decreases with rising atmospheric CO2 concentration, Ca. However, SD is also affected by light, air humidity and drought, all under systemic signalling from older leaves. This makes our understanding of how Ca controls SD incomplete. This study tested the hypotheses that SD is affected by the internal CO2 concentration of the leaf, Ci, rather than Ca, and that cotyledons, as the first plant assimilation organs, lack the systemic signal., Methods: Sunflower (Helianthus annuus), beech (Fagus sylvatica), arabidopsis (Arabidopsis thaliana) and garden cress (Lepidium sativum) were grown under contrasting environmental conditions that affected Ci while Ca was kept constant. The SD, pavement cell density (PCD) and stomatal index (SI) responses to Ci in cotyledons and the first leaves of garden cress were compared. (13)C abundance (δ(13)C) in leaf dry matter was used to estimate the effective Ci during leaf development. The SD was estimated from leaf imprints., Key Results: SD correlated negatively with Ci in leaves of all four species and under three different treatments (irradiance, abscisic acid and osmotic stress). PCD in arabidopsis and garden cress responded similarly, so that SI was largely unaffected. However, SD and PCD of cotyledons were insensitive to Ci, indicating an essential role for systemic signalling., Conclusions: It is proposed that Ci or a Ci-linked factor plays an important role in modulating SD and PCD during epidermis development and leaf expansion. The absence of a Ci-SD relationship in the cotyledons of garden cress indicates the key role of lower-insertion CO2 assimilation organs in signal perception and its long-distance transport., (© The Author 2014. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Phytophthora plurivora causes severe damage on Fagus sylvatica and is responsible for the extensive decline of European Beech throughout Europe. Unfortunately, no effective treatment against this disease is available. Phosphite (Phi) is known to protect plants against Phytophthora species; however, its mode of action towards P. plurivora is still unknown. To discover the effect of Phi on root infection, leaves were sprayed with Phi and roots were subsequently inoculated with P. plurivora zoospores. Seedling physiology, defense responses, colonization of root tissue by the pathogen and mortality were monitored. Additionally the Phi concentration in roots was quantified. Finally, the effect of Phi on mycelial growth and zoospore formation was recorded. Phi treatment was remarkably efficient in protecting beech against P. plurivora; all Phi treated plants survived infection. Phi treated and infected seedlings showed a strong up-regulation of several defense genes in jasmonate, salicylic acid and ethylene pathways. Moreover, all physiological parameters measured were comparable to control plants. The local Phi concentration detected in roots was high enough to inhibit pathogen growth. Phi treatment alone did not harm seedling physiology or induce defense responses. The up-regulation of defense genes could be explained either by priming or by facilitation of pathogen recognition of the host.

A multiplicative stomatal conductance model was constructed to estimate stomatal O3 uptake of Fagus crenata exposed to O3 under different N loads to the soil. Our stomatal conductance model included environmental functions such as the stomatal responses of F. crenata to diurnal changes, chronic O3 stress (AOT0), acute O3 stress (O3 concentration), and nitrogen load to soil. The model could explain 62% of the variability in stomatal conductance. We suggest therefore that stomatal closure induced by O3 and N load-induced soil acidification must be taken into account in developing a stomatal conductance model for estimating stomatal O3 uptake for future risk assessment of O3 impact on Japanese forest tree species such as F. crenata., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

To determine the effects of ozone (O3) on the canopy carbon budget, we investigated photosynthesis and respiration of leaves of Siebold's beech saplings under free air O3 exposure (60 nmol mol(-1), during daytime) in relation to the within-canopy light gradient; we then calculated the canopy-level photosynthetic carbon gain (PCG) and respiratory carbon loss (RCL) using a canopy photosynthesis model. Susceptibilities of photosynthesis and respiration to O3 were greater in leaves of upper canopy than in the lower canopy. The canopy net carbon gain (NCG) was reduced by O3 by 12.4% during one growing season. The increased RCL was the main factor for the O3-induced reduction in NCG in late summer, while contributions of the reduced PCG and the increased RCL to the NCG were almost the same in autumn. These results indicate contributions of changes in PCG and RCL under O3 to NCG were different between seasons., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

In recent years, many studies have focused on the limiting role of mesophyll conductance (gm ) to photosynthesis (An ) under water stress, but no studies have examined the effect of drought on gm through the forest canopy. We investigated limitations to An on leaves at different heights in a mixed adult stand of sessile oak (Quercus petraea) and beech (Fagus sylvatica) trees during a moderately dry summer. Moderate drought decreased An of top and lowest beech canopy leaves much more than in leaves located in the mid canopy; whereas in oak, An of the lower canopy was decreased more than in sunlit leaves. The decrease of An was probably not due to leaf-level biochemistry given that VCmax was generally unaffected by drought. The reduction in An was instead associated with reduction in stomatal and mesophyll conductances. Drought-induced increases in stomatal limitations were largest in leaves from the top canopy, whereas drought-induced increases in mesophyll limitations were largest in leaves from the lowest canopy. Sensitivity analysis highlighted the need to decompose the canopy into different leaf layers and to incorporate the limitation imposed by gm when assessing the impact of drought on the gas exchange of tree canopies., (© 2013 John Wiley & Sons Ltd.)

Background and Aims: Resistance of plants to ozone stress can be classified as either avoidance or tolerance. Avoidance of ozone stress may be explained by decreased stomatal conductance during ozone exposure because stomata are the principal interface for entry of ozone into plants. In this study, a coupled photosynthesis-stomatal model was modified to test whether the presence of ozone can induce avoidance of ozone stress by stomatal closure., Methods: The response of Siebold's beech (Fagus crenata), a representative deciduous tree species, to ozone was studied in a free-air ozone exposure experiment in Japan. Photosynthesis and stomatal conductance were measured under ambient and elevated ozone. An optimization model of stomata involving water, CO2 and ozone flux was tested using the leaf gas exchange data., Key Results: The data suggest that there are two phases in the avoidance of ozone stress via stomatal closure for Siebold's beech: (1) in early summer ozone influx is efficiently limited by a reduction in stomatal conductance, without any clear effect on photosynthetic capacity; and (2) in late summer and autumn the efficiency of ozone stress avoidance was decreased because the decrease in stomatal conductance was small and accompanied by an ozone-induced decline of photosynthetic capacity., Conclusions: Ozone-induced stomatal closure in Siebold's beech during early summer reduces ozone influx and allows the maximum photosynthetic capacity to be reached, but is not sufficient in older leaves to protect the photosynthetic system.

For the first time until now, the results from a prediction model (Atmospheric Dispersion Modelling System (ADMS)-Road) of pollutant dispersion in a street canyon were compared to the results obtained from biomonitors. In particular, the instrumental monitoring of particulate matter (PM10) and the biomonitoring of 14 polycyclic aromatic hydrocarbons (PAHs) and 11 metals by Quercus ilex leaves and Hypnum cupressiforme moss bags, acting as long- and short-term accumulators, respectively, were carried out. For both PAHs and metals, similar bioaccumulation trends were observed, with higher concentrations in biomonitors exposed at the leeward canyon side, affected by primary air vortex. The major pollutant accumulation at the leeward side was also predicted by the ADMS-Road model, on the basis of the prevailing wind direction that determines different exposure of the street canyon sides to pollutants emitted by vehicular traffic. A clear vertical (3, 6 and 9 m) distribution gradient of pollutants was not observed, so that both the model and biomonitoring results suggested that local air turbulences in the street canyon could contribute to uniform pollutant distribution at different heights.

The saccharification of beech wood using alkaline polyol pulping (AlkaPolP) and enzymatic hydrolysis was investigated. It will be demonstrated that the AlkaPolP process yields high quality pulps which can easily be hydrolyzed by cellulases. In order to find optimum reaction conditions chips of Fagus sylvatica were pretreated by alkaline glycerol at temperatures between 190 and 230 °C for 15, 20, and 25 min. The impacts of temperature and time were expressed using a severity factor R0. The dependencies of the conversion during enzymatic hydrolysis on severity, pulp yield, delignification and pulp composition are shown. In further experiments it was investigated if the sugar yields can be increased by the application of ultrasound or surfactants before enzyme addition. Up to 95% of the initial cellulose in wood were converted into glucose using cellulases from Trichoderma reesei and β-glucosidase from Aspergillus niger., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

We set up a free-air ozone (O(3)) exposure system for determining the photosynthetic responses of Siebold's beech (Fagus crenata) and oak (Quercus mongolica var. crispula) to O(3) under field conditions. Ten-year-old saplings of beech and oak were exposed to an elevated O(3) concentration (60 nmol mol(-1)) during daytime from 6 August to 11 November 2011. Ozone significantly reduced the net photosynthetic rate in leaves of both species in October, by 46% for beech and 15% for oak. In beech there were significant decreases in maximum rate of carboxylation, maximum rate of electron transport in photosynthesis, nitrogen content and photosynthetic nitrogen use efficiency, but not in oak. Stomatal limitation of photosynthesis was unaffected by O(3). We therefore concluded photosynthesis in beech is more sensitive to O(3) than that in oak, and the O(3)-induced reduction of photosynthetic activity in beech was due not to stomatal closure, but to biochemical limitation., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

• Increasing atmospheric concentrations of phytotoxic ozone (O(3) ) can constrain growth and carbon sink strength of forest trees, potentially exacerbating global radiative forcing. Despite progress in the conceptual understanding of the impact of O(3) on plants, it is still difficult to detect response patterns at the leaf level. • Here, we employed principal component analysis (PCA) to analyse a database containing physiological leaf-level parameters of 60-yr-old Fagus sylvatica (European beech) trees. Data were collected over two climatically contrasting years under ambient and twice-ambient O(3) regimes in a free-air forest environment. • The first principal component (PC1) of the PCA was consistently responsive to O(3) and crown position within the trees over both years. Only a few of the original parameters showed an O(3) effect. PC1 was related to parameters indicative of oxidative stress signalling and changes in carbohydrate metabolism. PC1 correlated with cumulative O(3) uptake over preceding days. • PC1 represents an O(3) -responsive multivariate pattern detectable in the absence of consistently measurable O(3) effects on individual leaf-level parameters. An underlying effect of O(3) on physiological processes is indicated, providing experimental confirmation of theoretical O(3) response patterns suggested previously., (© 2012 The Authors. New Phytologist © 2012 New Phytologist Trust.)

Impacts of elevated ground-level ozone (O(3)) on nitrogen (N) uptake and allocation were studied on mature European beech (Fagus sylvatica L.) and Norway spruce (Picea abies [L.] Karst.) in a forest stand, hypothesizing that: (i) chronically elevated O(3) limits nutrient uptake, and (ii) beech responds more sensitively to elevated O(3) than spruce, as previously found for juvenile trees. Tree canopies were exposed to twice-ambient O(3) concentrations (2 × O(3)) by a free-air fumigation system, with trees under ambient O(3) serving as control. After 5 years of O(3) fumigation, (15)NH(4)(15)NO(3) was applied to soil, and concentrations of newly acquired N (N(labelled)) and total N (N(total)) in plant compartments and soil measured. Under 2 × O(3), N(labelled) and N(total) were increased in the bulk soil and tended to be lower in fine and coarse roots of both species across the soil horizons, supporting hypothesis (i). N(labelled) was reduced in beech foliage by up to 60%, and by up to 50% in buds under 2 × O(3). Similarly, N(labelled) in stem bark and phloem was reduced. No such reduction was observed in spruce, reflecting a stronger effect on N acquisition in beech in accordance with hypothesis (ii). In spruce, 2 × O(3) tended to favour allocation of new N to foliage. N(labelled) in beech foliage correlated with cumulative seasonal transpiration, indicating impaired N acquisition was probably caused by reduced stomatal conductance and, hence, water transport under elevated O(3). Stimulated fine root growth under 2 × O(3) with a possible increase of below-ground N sink strength may also have accounted for lowered N allocation to above-ground organs. Reduced N uptake and altered allocation may enhance the use of stored N for growth, possibly affecting long-term stand nutrition.

Previously, laboratory experiments have revealed that freely diluted azole fungicides potentiate the direct toxic effect of pyrethroid insecticides on Daphnia magna. More ecologically relevant exposure scenarios where pesticides are adsorbed have not been addressed. In this study we exposed beech leaves (Fagus sylvatica) to the azole fungicide propiconazole (50 or 500 μg L(-1)), the pyrethroid insecticide alpha-cypermethrin (0.1 or 1 μg L(-1)) or any combination of the two for 3h. Exposed leaves were transferred to aquaria with or without an assemblage of macroinvertebrate shredders, and we studied treatment effects on rates of microbial leaf decomposition, microbial biomass (using C:N ratio as a surrogate measure) and macroinvertebrate shredding activity during 26 days post-exposure. Microbial leaf decomposition rates were significantly reduced in the propiconazole treatments, and the reduction in microbial activity was significantly correlated with loss of microbial biomass (increased C:N ratio). Macroinvertebrate shredding activity was significantly reduced in the alpha-cypermethrin treatments. In addition, the macroinvertebrate assemblage responded to the propiconazole treatments by increasing their consumption of leaf litter with lower microbial biomass, probably to compensate for the reduced nutritional quality of this leaf litter. We found no interaction between the two pesticides on macroinvertebrate shredding activity, using Independent Action as a reference model. In terms of microbial leaf decomposition rates, however, alpha-cypermethrin acted as an antagonist on propiconazole. Based on these results we emphasise the importance of considering indirect effects of pesticides in the risk assessment of surface water ecosystems., (Copyright © 2012 Elsevier B.V. All rights reserved.)

The hypothesis was tested that O(3)-induced changes in leaf-level photosynthetic parameters have the capacity of limiting the seasonal photosynthetic carbon gain of adult beech trees. To this end, canopy-level photosynthetic carbon gain and respiratory carbon loss were assessed in European beech (Fagus sylvatica) by using a physiologically based model, integrating environmental and photosynthetic parameters. The latter were derived from leaves at various canopy positions under the ambient O(3) regime, as prevailing at the forest site (control), or under an experimental twice-ambient O(3) regime (elevated O(3)), as released through a free-air canopy O(3) fumigation system. Gross carbon gain at the canopy-level declined by 1.7%, while respiratory carbon loss increased by 4.6% under elevated O(3). As this outcome only partly accounts for the decline in stem growth, O(3)-induced changes in allocation are referred to and discussed as crucial in quantitatively linking carbon gain with stem growth., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

We investigated the effects of ozone and leaf senescence on steady-state stomatal conductance and stomatal response to light variation. Measurements were carried out in a free-air ozone exposure experiment on a representative deciduous broadleaved tree species in Japan (Fagus crenata). Both steady-state and dynamic stomatal response to light variation varied intrinsically with season due to leaf senescence. Ozone induced the decrease in steady-state leaf gas exchange and the sluggish stomatal closure progressively. These findings suggest that ozone reduces the ability of plants to adapt to a fluctuating light environment under natural conditions, and therefore impairs plant growth and ability to control water loss., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

We measured the effect of elevated atmospheric CO(2) on atmospheric nitrogen (N(2)) fixation in the tree species Alnus glutinosa growing in monoculture or in mixture with the non-N(2)-fixing tree species Betula pendula and Fagus sylvatica. We addressed the hypotheses that (1) N(2) fixation in A. glutinosa will increase in response to increased atmospheric CO(2) concentrations, when growing in monoculture, (2) the impact of elevated CO(2) on N(2) fixation in A. glutinosa is the same in mixture and in monoculture and (3) the impacts of elevated CO(2) on N cycling will be evident by a decrease in leaf δ(15)N and by the soil-leaf enrichment factor (EF), and that these impacts will not differ between mixed and single species stands. Trees were grown in a forest plantation on former agricultural fields for four growing seasons, after which the trees were on average 3.8 m tall and canopy closure had occurred. Atmospheric CO(2) concentrations were maintained at either ambient or elevated (by 200 ppm) concentrations using a free-air CO(2) enrichment (FACE) system. Leaf δ(15)N was measured and used to estimate the amount (N(dfa)) and proportion (%N(dfa)) of N derived from atmospheric fixation. On average, 62% of the N in A. glutinosa leaves was from fixation. The %N(dfa) and N(dfa) for A. glutinosa trees in monoculture did not increase under elevated CO(2), despite higher growth rates. However, N(2) fixation did increase for trees growing in mixture, despite the absence of significant growth stimulation. There was evidence that fixed N(2) was transferred from A. glutinosa to F. sylvatica and B. pendula, but no evidence that this affected their CO(2) response. The results of this study show that N(2) fixation in A. glutinosa may be higher in a future elevated CO(2) world, but that this effect will only occur where the trees are growing in mixed species stands.

The ozone effect on Norway spruce (Picea abies (L) Karst.) and European beech (Fagus sylvatica L.) was studied on 48 monitoring plots in 2005-2008. These plots represent two major forest tree species stands of different ages in eight regions of the Czech Republic. The forest conditions were represented by defoliation and the annual radial increment of individual trees. The ozone exposure was assessed by using modeled values of mean annual O(3) concentration and the AOT40 index. The malondialdehyde (MDA) content of the foliage was analysed and used as an indicator of oxidative stress. The correlation analysis showed a significant relation of Norway spruce defoliation to the AOT40 exposure index, and European beech defoliation to the MDA level. The radial increment response to ozone was significant only for the European beech: (a) the correlation analysis showed its decrease with increasing AOT40; (b) the regression model showed its decrease with increasing mean annual ozone concentration only at lower altitudes (<700 m a.s.l.).

Ozone levels in the Jizerske hory Mts. measured at 13 sites by diffusive samplers during the 2006 and 2007 vegetation seasons are presented. A significant ozone gradient (5.4 ppb in 2006 and 4.0 ppb in 2007) per 100 m difference in altitude between 370 and 1,100 m a.s.l. was recorded. High-resolution maps of phytotoxic potential were developed. The AOT40 threshold (5 ppm h) was exceeded over the entire area with the highest levels exceeding this threshold by 12 times in the upper portions of the mountains. Ozone visible injury was evaluated at four of the monitoring sites on seven native plant and tree species. Four species showed ozone-like symptoms, two of which (Rubus idaeus and Fagus sylvatica) were confirmed as ozone-induced. Our results indicate that ambient ozone is likely to have a much lower impact on the Jizerske hory Mts. vegetation than expected, considering the measured ambient ozone exposures and favourable environmental conditions for ozone uptake.

In June 2008, 303,000 L of hydrofracturing fluid from a natural gas well were applied to a 0.20-ha area of mixed hardwood forest on the Fernow Experimental Forest, West Virginia. During application, severe damage and mortality of ground vegetation was observed, followed about 10 d later by premature leaf drop by the overstory trees. Two years after fluid application, 56% of the trees within the fluid application area were dead. Ehrh. was the tree species with the highest mortality, and L. was the least affected, although all tree species present on the site showed damage symptoms and mortality. Surface soils (0-10 cm) were sampled in July and October 2008, June and October 2009, and May 2010 on the fluid application area and an adjacent reference area to evaluate the effects of the hydrofracturing fluid on soil chemistry and to attempt to identify the main chemical constituents of the hydrofracturing fluid. Surface soil concentrations of sodium and chloride increased 50-fold as a result of the land application of hydrofracturing fluids and declined over time. Soil acidity in the fluid application area declined with time, perhaps from altered organic matter cycling. This case study identifies the need for further research to help understand the nature and the environmental impacts of hydrofracturing fluids to devise optimal, safe disposal strategies., (Copyright © by the American Society of Agronomy, Crop Science Society of America, and Soil Science Society of America, Inc.)

Background and Aims: Masting, i.e. synchronous but highly variable interannual seed production, is a strong sink for carbon and nutrients. It may, therefore, compete with vegetative growth. It is currently unknown whether increased atmospheric CO(2) concentrations will affect the carbon balance (or that of other nutrients) between reproduction and vegetative growth of forest species. In this study, reproduction and vegetative growth of shoots of mature beech (Fagus sylvatica) trees grown at ambient and elevated atmospheric CO(2) concentrations were quantified. It was hypothesized that within a shoot, fruiting has a negative effect on vegetative growth, and that this effect is ameliorated at increased CO(2) concentrations., Methods: Reproduction and its competition with leaf and shoot production were examined during two masting events (in 2007 and 2009) in F. sylvatica trees that had been exposed to either ambient or elevated CO(2) concentrations (530 µmol mol(-1)) for eight consecutive years, between 2000 and 2008., Key Results: The number of leaves per shoot and the length of terminal shoots was smaller or shorter in the two masting years compared with the one non-masting year (2008) investigated, but they were unaffected by elevated CO(2) concentrations. The dry mass of terminal shoots was approx. 2-fold lower in the masting year (2007) than in the non-masting year in trees growing at ambient CO(2) concentrations, but this decline was not observed in trees exposed to elevated CO(2) concentrations. In both the CO(2) treatments, fruiting significantly decreased nitrogen concentration by 25 % in leaves and xylem tissue of 1- to 3-year-old branches in 2009., Conclusions: Our findings indicate that there is competition for resources between reproduction and shoot growth. Elevated CO(2) concentrations reduced this competition, indicating effects on the balance of resource allocation between reproduction and vegetative growth in shoots with rising atmospheric CO(2) concentrations.

For a quantitative estimate of the ozone effect on vegetation reliable models for ozone uptake through the stomata are needed. Because of the analogy of ozone uptake and transpiration it is possible to utilize measurements of water loss such as sap flow for quantification of ozone uptake. This technique was applied in three beech (Fagus sylvatica) stands in Switzerland. A canopy conductance was calculated from sap flow velocity and normalized to values between 0 and 1. It represents mainly stomatal conductance as the boundary layer resistance in forests is usually small. Based on this relative conductance, stomatal functions to describe the dependence on light, temperature, vapour pressure deficit and soil moisture were derived using multivariate nonlinear regression. These functions were validated by comparison with conductance values directly estimated from sap flow. The results corroborate the current flux parameterization for beech used in the DO3SE model., (Copyright (c) 2010 Elsevier Ltd. All rights reserved.)

Canopies of adult European beech (Fagus sylvatica) and Norway spruce (Picea abies) were labeled with CO(2) depleted in (13)C to evaluate carbon allocation belowground. One-half the trees were exposed to elevated O(3) for 6 yrs prior to and during the experiment. Soil-gas sampling wells were placed at 8 and 15 cm and soil CO(2) was sampled during labeling in mid-late August, 2006. In beech, delta(13)CO(2) at both depths decreased approximately 50 h after labeling, reflecting rapid translocation of fixed C to roots and release through respiration. In spruce, label was detected in fine-root tissue, but there was no evidence of label in delta(13)CO(2). The results show that C fixed in the canopy rapidly reaches respiratory pools in beech roots, and suggest that spruce may allocate very little of recently-fixed carbon into root respiration during late summer. A change in carbon allocation belowground due to long-term O(3) exposure was not observed.

Stomatal O3 fluxes to a mixed beech/spruce stand (Fagus sylvatica/Picea abies) in Central Europe were determined using two different approaches. The sap flow technique yielded the tree-level transpiration, whereas the eddy covariance method provided the stand-level evapotranspiration. Both data were then converted into stomatal ozone fluxes, exemplifying this novel concept for July 2007. Sap flow-based stomatal O3 flux was 33% of the total O3 flux, whereas derivation from evapotranspiration rates in combination with the Penman-Monteith algorithm amounted to 47%. In addition to this proportional difference, the sap flow-based assessment yielded lower levels of stomatal O3 flux and reflected stomatal regulation rather than O3 exposure, paralleling the daily courses of canopy conductance for water vapor and eddy covariance-based total stand-level O3 flux. The demonstrated combination of sap flow and eddy covariance approaches supports the development of O3 risk assessment in forests from O3 exposure towards flux-based concepts., (Copyright 2009 Elsevier Ltd. All rights reserved.)

In 2006, a controlled infection study was performed in the 'Kranzberger Forst' to address the following questions: (1) Will massive artificial inoculation with Apiognomonia errabunda override the previously observed inhibitory effect of chronic ozone? (2) Can biochemical or molecular markers be detected to account for the action of ozone? To this end six adult beech trees were chosen, three ozone fumigated (2x ozone) and three control trees (ambient = 1x ozone). Spore-sprayed branches of sun and shade crown positions of each of the trees, and uninoculated control branches, were enclosed in 100-L plastic bags for one night to facilitate infection initiation. Samples were taken within a five-week period after inoculation. A. errabunda infestation levels quantified by real-time PCR increased in leaves that were not fumigated with additional ozone. Cell wall components and ACC (ethylene precursor 1-amino cyclopropane-1-carboxylic acid) increased upon ozone fumigation and may in part lead to the repression of fungal infection., (2009 Elsevier Ltd. All rights reserved.)

In a 50- to 70-year-old mixed stand of Norway spruce (Picea abies (L.) Karst.) and European beech (Fagus sylvatica L.) in Germany, tree cohorts have been exposed to double ambient ozone (2xO(3)) from 2000 through 2007 and can be compared with trees in the same stand under the ambient ozone regime (1xO(3)). Annual diameter growth, allocation pattern, stem form, and stem volume were quantified at the individual tree and stand level. Ozone fumigation induced a shift in the resource allocation into height growth at the expense of diameter growth. This change in allometry leads to rather cone-shaped stem forms and reduced stem stability in the case of spruce, and even neiloidal stem shapes in the case of beech. Neglect of such ozone-induced changes in stem shape may lead to a flawed estimation of volume growth. On the stand level, 2xO(3) caused, on average, a decrease of 10.2 m(3) ha(-1) yr(-1) in European beech., (2009 Elsevier Ltd. All rights reserved.)

The growth-differentiation balance hypothesis (GDBH) predicts changes in susceptibility of plants against herbivores with changing resource availability. In the presented study we tested the validity of the GDBH for trees infected with a root pathogen. For this purpose Fagus sylvatica seedlings grown under different atmospheric CO(2)- and soil nitrogen regimes were infected with the root pathogen Phytophthora citricola. High nitrogen supply increased total biomass of beech regardless of the CO(2)-treatment, whereas elevated CO(2) enhanced biomass only in the high nitrogen treatment. The responses of beech under the different growing regimes to the Phytophthora root infection were not in line with the predictions of the GDBH. Enhanced susceptibility of beech against P. citricola was found in seedlings grown under elevated CO(2) and low nitrogen supply. Fifteen months after inoculation these plants were characterized by enhanced water use efficiency, by altered root-shoot ratios, and by enhanced specific root tip densities., (2009 Elsevier Ltd. All rights reserved.)

The effects of experimentally elevated O(3) on soil respiration rates, standing fine-root biomass, fine-root production and delta(13)C signature of newly produced fine roots were investigated in an adult European beech/Norway spruce forest in Germany during two subsequent years with contrasting rainfall patterns. During humid 2002, soil respiration rate was enhanced under elevated O(3) under beech and spruce, and was related to O(3)-stimulated fine-root production only in beech. During dry 2003, the stimulating effect of O(3) on soil respiration rate vanished under spruce, which was correlated with decreased fine-root production in spruce under drought, irrespective of the O(3) regime. delta(13)C signature of newly formed fine-roots was consistent with the differing g(s) of beech and spruce, and indicated stomatal limitation by O(3) in beech and by drought in spruce. Our study showed that drought can override the stimulating O(3) effects on fine-root dynamics and soil respiration in mature beech and spruce forests., (2009 Elsevier Ltd. All rights reserved.)

Here we synthesize key findings from a series of experiments to gain new insight on inter-plant competition between juvenile beech (Fagus sylvatica) and spruce (Picea abies) under the influence of increased O(3) and CO(2) concentrations. Competitiveness of plants was quantified and mechanistically interpreted as space-related resource investments and gains. Stable isotopes were addressed as temporal integrators of plant performance, such as photosynthesis and its relation to water use and nitrogen uptake. In the weaker competitor, beech, efficiency in space-related aboveground resource investment was decreased in competition with spruce and positively related to Delta(13)C, as well as stomatal conductance, but negatively related to delta(18)O. Likewise, our synthesis revealed that strong belowground competition for water in spruce was paralleled in this species by high N assimilation capacity. We suggest combining the time-integrative potential of stable isotopes with space-related investigations of competitiveness to accomplish mechanistic understanding of plant competition for resources., (2009 Elsevier Ltd. All rights reserved.)

Tropospheric ozone causes severe oxidative stress in plants. To investigate the transcriptional responsiveness of adult trees to ozone, fully-expanded sun and shade leaves of mature beech trees were harvested at four time points over the entire vegetation period in 2005 and 2006. Microarray analyses were conducted on leaves from trees grown in the field under ambient and twice-ambient ozone concentrations at Kranzberger Forst (Bavaria). Beech trees changed their transcript levels in response to ozone. In the years 2005 and 2006 different transcription patterns were observed; this may have been a result of different weather conditions and ozone uptake. Furthermore, we obtained differences in mRNA expression patterns between shade and sun leaves. In the ozone-treated sun leaves of 2005, slightly up- and down-regulated transcript levels were detected, particularly in the spring and autumn, whereas shade leaves clearly exhibited reduced mRNA levels, particularly at the end of the vegetation period. In 2006, this pattern could not be confirmed, and in the autumn, four other transcripts were slightly up-regulated in ozone-treated shade leaves. In addition, two other transcripts were found to be influenced in sun leaves in the spring/summer. While we detected changes in the levels of only a few transcripts, the observed effects were not identical in both years. In conclusion, elevated ozone exhibited very small influence on the transcription levels of genes of mature beech trees., (2009 Elsevier Ltd. All rights reserved.)