Flux-based response of sucrose and starch in leaves of adult beech trees (Fagus sylvatica L.) under chronic free-air O3 fumigation

Investigations on sucrose and starch contents in leaves of 60-year-old beech trees ( FAGUS SYLVATICA L.) are the focus of the present study. Five trees were exposed to a twice ambient ozone regime (2 x O(3)) with a free-air canopy exposure system throughout the seasons and five trees under the prevailing ambient ozone regime served as controls (1 x O(3)). In order to examine chronic ozone (O(3)) effects, leaf samples from the sun and shade crowns of the trees were analyzed five times throughout the growing seasons in 2003 and 2004. Sucrose concentrations of leaves collected in 2004 were consistently lower than those taken in 2003, regardless of the O(3) treatment and crown position. However, the opposite was found for starch. O(3) caused a reduction of sucrose and starch contents of sun leaves in both years. Due to the fact that O(3)-responsiveness depends on the O(3) uptake through stomata during the season, all carbohydrate data were related to the cumulative O(3) uptake (COU). Little differences were found comparing sucrose and starch contents in leaves of trees grown under ambient or elevated O (3) regimes, possibly indicating the high capacity of leaves of adult beech to cope with rising O(3) exposure. Even under 2 x O(3), leaves were still able to regulate the O(3) intake by narrowing their stomata at the cost of CO(2)-uptake and sugar synthesis. In order to clarify whole-tree response patterns carbohydrate data were compared with photosynthesis, stomatal conductance and electron transport rates. In 2004 all parameters revealed a significant common response pattern to COU that indicated a reduction for all parameters under 2 x O(3).     

O3 flux-related responsiveness of photosynthesis, respiration, and stomatal conductance of adult Fagus sylvatica to experimentally enhanced free-air O3 exposure

Knowledge of responses of photosynthesis, respiration, and stomatal conductance to cumulative ozone uptake (COU) is still scarce, and this is particularly the case for adult trees. The effect of ozone (O(3)) exposure on trees was examined with 60-year-old beech trees (FAGUS SYLVATICA) at a forest site of southern Germany. Trees were exposed to the ambient O(3) regime (1 x O(3)) or an experimentally elevated twice-ambient O(3) regime (2 x O(3)). The elevated 2 x O (3) regime was provided by means of a free-air O(3) canopy exposure system. The hypotheses were tested that (1) gas exchange is negatively affected by O(3) and (2) the effects of O(3) are dose-dependent and thus the sizes of differences between treatments are positively related to COU. Gas exchange (light-saturated CO(2) uptake rate A(max), stomatal conductance g (s), maximum rate of carboxylation Vc (max), ribulose-1,5-bisphosphate turnover limited rate of photosynthesis J (max), CO(2) compensation point CP, apparent quantum yield of net CO(2) uptake AQ, carboxylation efficiency CE, day- and nighttime respiration) and chlorophyll fluorescence (electron transfer rate, ETR) were measured IN SITU on attached sun and shade leaves. Measurements were made periodically throughout the growing seasons of 2003 (an exceptionally dry year) and 2004 (a year with average rainfall). In 2004 Vc(max), J(max), and CE were lower in trees receiving 2 x O(3) compared with the ambient O(3) regime (1 x O(3)). Treatment differences in Vc (max), J (max), CE were rather small in 2004 (i.e., parameter levels were lower by 10 - 30 % in 2 x O(3) than 1 x O(3)) and not significant in 2003. In 2004 COU was positively correlated with the difference between treatments in A (max), g (s), and ETR (i.e., consistent with the dose-dependence of O(3)'s deleterious effects). However, in 2003, differences in A(max), g (s), and ETR between the two O(3) regimes were smaller at the end of the dry summer 2003 (i.e., when COU was greatest). The relationship of COU with effects on gas exchange can apparently be complex and, in fact, varied between years and within the growing season. In addition, high doses of O(3) did not always have significant effects on leaf gas exchange. In view of the key findings, both hypotheses were to be rejected.     

Gas exchange and antioxidative compounds in young beech trees under free-air ozone exposure and comparisons to adult trees

Three-year-old beech (Fagus sylvatica) seedlings growing in containers were placed into the sun and shade crown of a mature beech stand exposed to ambient (1 x O(3)) and double ambient (2 x O(3)) ozone concentrations at a free-air exposure system ("Kranzberg Forst", Germany). Pigments, alpha-tocopherol, glutathione, ascorbate, and gas exchange were measured in leaves during 2003 (a drought year) and 2004 (an average year). Sun-exposed seedlings showed higher contents of antioxidants, xanthophylls, and beta-carotene and lower contents of chlorophyll, alpha-carotene, and neoxanthin than shade-exposed seedlings. In 2003 sun-exposed seedlings showed higher contents of carotenoids and total glutathione and lower net photosynthesis rates (A(max)) compared to 2004. O(3) exposure generally affected the content of chlorophyll, the xanthophyll cycle, and the intercellular CO(2) concentration (c(i)). Seedlings differed from the adjacent adult trees in most biochemical and physiological parameters investigated: Sun exposed seedlings showed higher contents of alpha-tocopherol and xanthophylls and lower contents of ascorbate, chlorophyll, neoxanthin, and alpha-carotene compared to adult trees. Shade exposed seedlings had lower contents of xanthophylls, alpha-carotene, and alpha-tocopherol than shade leaves of old-growth trees. In 2003, seedlings had higher A(max), stomatal conductance (g(s)), and c(i) under 2 x O(3) than adult trees. The results showed that shade acclimated beech seedlings are more sensitive to O(3), possibly due to a lower antioxidative capacity per O(3) uptake. We conclude that beech seedlings are uncertain surrogates for adult beech trees.     

Effects of long-term, free-air ozone fumigation on the cytokinin content of mature beech trees

We present the results of a study of the effects of chronic exposure to elevated ozone on the cytokinins of mature beech trees. Methods for analysing the cytokinin (CK) content of beech (FAGUS SYLVATICA) were developed using seven enzyme-linked immunosorbent assays (ELISAs). Samples taken during 2003 and 2004 from 10 mature beech trees in Kranzberg forest, 5 trees exposed to twice ambient ozone (2 x O(3)) by free-air fumigation and 5 control trees (1 x O(3)), were analysed. In 2003 and 2004 the cytokinin content of leaf samples followed a similar seasonal pattern. In leaf samples, the content of aromatic types was equal to that of the isoprenoid types. In root samples, the level of aromatic types was no different from leaves, but that of the isoprenoid types was much higher. Leaf and phloem cytokinin contents for 2 x O(3) trees were lower than for 1 x O(3) at almost all sampling times. The effect of ozone was greater for leaves in the sun crown than for leaves in the shade crown. By contrast, the root and xylem contents of cytokinin for 2 x O(3) trees were greatly elevated over the values for 1 x O(3) trees early in the growing season. We propose that O(3)-associated CK destruction in leaves reduces CK-mediated root growth suppression. The resulting increases in root growth and ectomycorrhiza, reported by other groups in the Kranzberg forest project, are likely to be responsible for the increased CK export in xylem, although O(3)-associated CK destruction in the leaves appears to nullify this increase.     

Effects of long-term free-air ozone fumigation on delta15N and total N in Fagus sylvatica and associated mycorrhizal fungi

Patterns of nitrogen (N) isotope composition (delta(15)N) and total N contents were determined in leaves, fine roots, root-associated ectomycorrhizal fungi (ECM) of adult beech trees (FAGUS SYLVATICA), and soil material under ambient (1 x O(3)) and double ambient (2 x O(3)) atmospheric ozone concentrations over a period of two years. From fine root to leaf material delta(15)N decreased consecutively. Under enhanced ozone concentrations total N was reduced in fine roots and delta(15)N showed a decrease in roots and leaves. In the soil and in most types of mycorrhizae, delta(15)N and total N were not altered due to ozone fumigation. The number of vital ectomycorrhizal root tips increased and the mycorrhizal community structure changed in 2 x O(3). Simultaneously, the specific rate of inorganic N-uptake by the roots was reduced under the double ozone regime. From these results it is assumed that 2 x O(3) changes N-nutrition of the trees at the level of N-acquisition, as indicated by enhanced mycorrhizal root tip density, altered mycorrhizal species composition, and reduced specific N-uptake rates.     

Plant-mediated nitrous oxide emissions from beech (Fagus sylvatica) leaves

Nitrous oxide (N2O) emission estimates from forest ecosystems are based currently on emission measurements using soil enclosures. Such enclosures exclude emissions via tall plants and trees and may therefore underestimate the whole-ecosystem N2O emissions. Here, we measured plant-mediated N2O emissions from the leaves of potted beech (Fagus sylvatica) seedlings after fertilizing the soil with 15N-labelled ammonium nitrate (15NH4(15)NO3), and after exposing the roots to elevated concentrations of N2O. Ammonium nitrate fertilization induced N2O + 15N2O emissions from beech leaves. Likewise, the foliage emitted N2O after beech roots were exposed to elevated concentrations of N2O. The average N2O emissions from the fertilization and the root exposure experiments were 0.4 and 2.0 microg N m(-2) leaf area h(-1), respectively. Higher than ambient atmospheric concentrations of N2O in the leaves of the forest trees indicate a potential for canopy N2O emissions in the forest. Our experiments demonstrate the existence of a previously overlooked pathway of N2O to the atmosphere in forest ecosystems, and bring about a need to investigate the magnitude of this phenomenon at larger scales.     

Changes in the community of ectomycorrhizal fungi and increased fine root number under adult beech trees chronically fumigated with double ambient ozone concentration

Forest soils are an important but under-studied part of forest ecosystems. The effects of O(3) on below-ground processes in a mature forest have only received limited attention so far. In our study, we have analysed the community of ectomycorrhizal fungi and beech fine root dynamics over two growing seasons (2003 - 2004) in a 70-year old mixed spruce-beech forest stand, in which two groups of five adult beech trees were either fumigated by 2 x ambient ozone concentration or used as control. The main difference between previous studies and our approach was that previous studies were performed on seedlings in pot experiments or in closed or open top chambers, and not IN SITU, in a mature forest stand. Although beech is a relatively unresponsive species to tropospheric O(3), we found a pronounced effect of 2 x O(3) on the number of vital ectomycorrhizal root tips and non-turgescent fine roots. Both categories of roots were significantly increased when compared to controls in two consecutive years at each sampling event. The number of types of ectomycorrhizae and species richness increased in 2004, but not in the extremely dry year 2003. We hypothesised that the observed changes might be an expression of a transitional state in below-ground succession of niches caused by an O(3) induced effect on carbon allocation to roots and the rhizosphere. We have detected changes in ectomycorrhizal species level, however Shannon-Weavers species diversity index and percentage of types of ectomycorrhizae did not change significantly in any sampling year thus indicating our results cannot be unequivocally explained by summer drought in year 2003 or by O(3) exposure alone.     

Evidence that branch cuvettes are reasonable surrogates for estimating O3 effects in entire tree crowns

Within the scope of quantifying ozone (O(3)) effects on forest tree crowns it is still an open question whether cuvette branches of adult trees are reasonable surrogates for O(3) responses of entire tree crowns and whether twigs exhibit autonomy in defense metabolism in addition to carbon autonomy. Therefore, cuvette-enclosed branches of mature beech (Fagus sylvatica) trees were compared with branches exposed to the same and different ozone regimes by a free-air fumigation system under natural stand conditions by means of a VICE VERSA experiment. For this purpose, cuvettes receiving 1 x O(3) air were mounted in trees exposed to 2 x O(3) and cuvettes receiving 2 x O(3) air were mounted in trees exposed to 1 x O (3) in the upper sun crown. At the end of the fumigation period in September 2004, leaves were examined for differences in gas exchange parameters, pigments, antioxidants, carbohydrates, and stable isotope ratios. No significant differences in foliar gas exchange, total carbohydrates, stable isotope ratios, pigment, and antioxidant contents were found as a consequence of cuvette enclosure (cuvette versus free-air branches) of the same O(3) concentrations besides increase of glucose inside the cuvettes and reduction of the de-epoxidation state of the xanthophyll cycle pigments. No significant ozone effect was found for the investigated gas exchange and most biochemical parameters. The total and oxidized glutathione level of the leaves was increased by the 2 x O(3) treatment in the cuvette and the free-air branches, but this effect was significant only for the free-air branches. From these results we conclude that cuvette branches are useful surrogates for examining the response of entire tree crowns to elevated O(3) and that the defence metabolism of twigs seems to be at least partially autonomous.     

Chronic ozone exposure affects leaf senescence of adult beech trees: a chlorophyll fluorescence approach

Accelerated leaf senescence is one of the harmful effects of elevated tropospheric ozone concentrations ([O(3)]) on plants. The number of studies dealing with mature forest trees is scarce however. Therefore, five 66-year-old beech trees (Fagus sylvatica L.) have been exposed to twice-ambient (2xambient) [O(3)] levels by means of a free-air canopy O(3) exposure system. During the sixth year of exposure, the hypothesis of accelerated leaf senescence in 2xambient [O(3)] compared with ambient [O(3)] trees was tested for both sun and shade leaves. Chlorophyll (chl) fluorescence was used to assess the photosynthetic quantum yield, and chl fluorescence images were processed to compare functional leaf homogeneity and the proportion of O(3)-injured leaf area (stipples) under ambient and 2xambient [O(3)] regimes. Based on the analysis of chl fluorescence images, sun leaves of both ambient and 2xambient [O(3)] trees had apparently developed typical necrotic O(3) stipples during high O(3) episodes in summer, while accelerated senescence was only observed with sun leaves of 2xambient [O(3)] trees. This latter effect was indicated along with a faster decrease of photosynthetic quantum yield, but without evidence of changes in non-photochemical quenching. Overall, treatment effects were small and varied among trees. Therefore, compared with ambient [O(3)], the consequence of the observed O(3)-induced accelerated leaf senescence for the carbon budget is likely limited.     

CASIROZ: Root parameters and types of ectomycorrhiza of young beech plants exposed to different ozone and light regimes

Tropospheric ozone (O(3)) triggers physiological changes in leaves that affect carbon source strength leading to decreased carbon allocation below-ground, thus affecting roots and root symbionts. The effects of O(3) depend on the maturity-related physiological state of the plant, therefore adult and young forest trees might react differently. To test the applicability of young beech plants for studying the effects of O(3) on forest trees and forest stands, beech seedlings were planted in containers and exposed for two years in the Kranzberg forest FACOS experiment (Free-Air Canopy O(3) Exposure System, http://www.casiroz.de ) to enhanced ozone concentration regime (ambient [control] and double ambient concentration, not exceeding 150 ppb) under different light conditions (sun and shade). After two growing seasons the biomass of the above- and below-ground parts, beech roots (using WinRhizo programme), anatomical and molecular (ITS-RFLP and sequencing) identification of ectomycorrhizal types and nutrient concentrations were assessed. The mycorrhization of beech seedlings was very low ( CA. 5 % in shade, 10 % in sun-grown plants), no trends were observed in mycorrhization (%) due to ozone treatment. The number of Cenococcum geophilum type of ectomycorrhiza, as an indicator of stress in the forest stands, was not significantly different under different ozone treatments. It was predominantly occurring in sun-exposed plants, while its majority share was replaced by Genea hispidula in shade-grown plants. Different light regimes significantly influenced all parameters except shoot/root ratio and number of ectomycorrhizal types. In the ozone fumigated plants the number of types, number of root tips per length of 1 to 2 mm root diameter, root length density per volume of soil and concentration of Mg were significantly lower than in control plants. Trends to a decrease were found in root, shoot, leaf, and total dry weights, total number of root tips, number of vital mycorrhizal root tips, fine root (mass) density, root tip density per surface, root area index, concentration of Zn, and Ca/Al ratio. Due to the general reduction in root growth indices and nutrient cycling in ozone-fumigated plants, alterations in soil carbon pools could be predicted.     

Spatiotemporal changes of beetle communities across a tree diversity gradient

Aim Plant and arthropod diversity are often related, but data on the role of mature tree diversity on canopy insect communities are fragmentary. We compare species richness of canopy beetles across a tree diversity gradient ranging from mono‐dominant beech to mixed stands within a deciduous forest, and analyse community composition changes across space and time. Location Germany’s largest exclusively deciduous forest, the Hainich National Park (Thuringia). Methods We used flight interception traps to assess the beetle fauna of various tree species, and applied additive partitioning to examine spatiotemporal patterns of diversity. Results Species richness of beetle communities increased across the tree diversity gradient from 99 to 181 species per forest stand. Intra‐ and interspecific spatial turnover among trees contributed more than temporal turnover among months to the total γ‐beetle diversity of the sampled stands. However, due to parallel increases in the number of habitat generalists and the number of species in each feeding guild (herbivores, predators and fungivores), no proportional changes in community composition could be observed. If only beech trees were analysed across the gradient, patterns were similar but temporal (monthly) species turnover was higher compared to spatial turnover among trees and not related to tree diversity. Main conclusions The changes in species richness and community composition across the gradient can be explained by habitat heterogeneity, which increased with the mix of tree species. We conclude that understanding temporal and spatial species turnover is the key to understanding biodiversity patterns. Mono‐dominant beech stands are insufficient to conserve fully the regional species richness of the remaining semi‐natural deciduous forest habitats in Central Europe, and analysing beech alone would have resulted in the misleading conclusion that temporal (monthly) turnover contributes more to beetle diversity than spatial turnover among different tree species or tree individuals.     

Stem growth and canopy dynamics in a world-wide range of Fagus forests

Abstract. Forests dominated by Fagus (beech) occur widely in the Northern Hemisphere. Tree species dominant together with beech vary in tolerance of understorey conditions. They are deciduous broad-leaved, evergreen broad-leaved or evergreen coniferous. The frequency and intensity of events that reduce the forest canopy cover are important determinants of the ratio of beech to other species in the canopy. For trees in the understorey and the canopy, stem diameter growth rate is determined by light regime and growing space which in turn are determined by canopy cover. We evaluated increase in stem diameter growth rate as an indicator of sudden reductions in canopy cover and canopy dynamics. We used tree-ring chronologies and calculated an index of growth rate increase (GI) to compare the canopy dynamics of 11 natural beech forests. Per site, the annual average value of GI poorly reflected the effects of dry or cool summers, and it clearly reflected events like tornados and hurricanes that removed substantial canopy cover. Among groups of sites average values of GI were significantly different. In the sites with a lower level of average GI, the establishment of the more shade tolerant tree and shrub species in the understorey was favoured, and subcanopy layers became more dense. On the other hand, higher levels of average GI allowed for more light demanding tree species to reach the canopy.     

Tree species distribution in canopy gaps and mature forest in an area of cloud forest of the Ibitipoca Range,south-eastern Brazil

The tree community of both canopy gaps and mature forest was surveyed in a 5 ha plot of cloud forest in the Ibitipoca Range, south-eastern Brazil, aiming at: (a) comparing the tree community structure of canopy gaps with that of three strata of the mature forest, and (b) relating the tree community structure of canopy gaps with environmental and biotic variables. All saplings of canopy trees with 1–5 m of height established in 31 canopy gaps found within the plot were identified and measured. Mature forest trees with dbh 3 cm were sampled in four 40×40 quadrats laid on the four soil sites recognised in the local soil catena. All surveyed trees were identified, measured and distributed into three forest strata: understorey (<5 m of height), sub-canopy (5.1–15 m) and canopy (15.1–30 m). The following variables were obtained for each gap: mode of formation, age, soil site, slope grade, size, canopy openness and abundance of bamboos and lianas. A detrended correspondence analysis indicated that the tree community structure of gaps in all soil sites was more similar to that of the mature forest understorey, suggesting that the bank of immatures plays an important role in rebuilding the forest canopy and that gap phases may be important for understorey shade-tolerant species. There was evidence of gap-dependence for establishment for only one canopy tree species. Both canonical correspondence analysis and correlation analysis demonstrated for a number of tree species that the distribution of their saplings in canopy gaps was significantly correlated with two variables: soil site and canopy openness. The future forest structure at each gap is probably highly influenced by both the present structure of the adjacent mature forest and the gap creation event.     

Variability of stomatal conductance, leaf anatomy, and seasonal leaf wettability of young and adult European beech leaves along a vertical canopy gradient

This study assessed the variation of leaf anatomy, chlorophyll content index (CCI), maximal stomatal conductance (g _s ^max ) and leaf wettability within the canopy of an adult European beech tree (Fagus sylvatica L.) and for beech saplings placed along the vertical gradient in the canopy. At the top canopy level (CL_28m) of the adult beech, CCI and leaf anatomy reflected higher light stress, while g _s ^max increased with height, reflecting the importance of gas exchange in the upper canopy layer. Leaf wettability, measured as drop contact angle, decreased from 85.5°?±?1.6° (summer) to 57.5°?±?2.8° (autumn) at CL_28m of the adult tree. At CL_22m, adult beech leaves seemed to be better optimized for photosynthesis than the CL_28m leaves because of a large leaf thickness with less protective and impregnated substances, and a higher CCI. The beech saplings, in contrast, did not adapt their stomatal characteristics and leaf anatomy according to the same strategy as the adult beech leaves. Consequently, care is needed when scaling up experimental results from seedlings to adult trees.     

Insect attraction to herbivore-induced beech volatiles under different forest management regimes

Insect herbivore enemies such as parasitoids and predators are important in controlling herbivore pests. From agricultural systems we know that land-use intensification can negatively impact biological control as an important ecosystem service. The aim of our study was to investigate the importance of management regime for natural enemy pressure and biological control possibilities in forests dominated by European beech. We hypothesize that the volatile blend released from herbivore-infested beech trees functions as a signal, attracting parasitoids and herbivore enemies. Furthermore, we hypothesize that forest management regime influences the composition of species attracted by these herbivore-induced beech volatiles. We installed flight-interception traps next to Lymantria dispar caterpillar-infested young beech trees releasing herbivore-induced volatiles and next to non-infested control trees. Significantly more parasitoids were captured next to caterpillar-infested trees compared to non-infested controls, irrespective of forest type. However, the composition of the trophic guilds in the traps did vary in response to forest management regime. While the proportion of chewing insects was highest in non-managed forests, the proportion of sucking insects peaked in forests with low management and of parasitoids in young, highly managed, forest stands. Neither the number of naturally occurring beech saplings nor herbivory levels in the proximity of our experiment affected the abundance and diversity of parasitoids caught. Our data show that herbivore-induced beech volatiles attract herbivore enemies under field conditions. They further suggest that differences in the structural complexity of forests as a consequence of management regime only play a minor role in parasitoid activity and thus in indirect tree defense.     

Beech forest management does not affect the infestation rate of the beech scale Cryptococcus fagisuga across three regions in Germany

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Canopy surface topography in a French Guiana forest and the folded forest theory

The canopy surface is an undulating surface that follows the irregular contours of the upper tree crowns and defines the inner and the outer limits of the forest volume. In French Guiana, the height of the canopy surface was surveyed in both a primary and a 20-years old clear-felled secondary forest plot. The topographic surface was displayed in a three-dimensional mesh, where X and Y are horizontal co-ordinates, and Z is the canopy height measured from the ground with an optical telemeter. The statistical dispersion of Z-data, and the spatial tree height variations, are interpreted at different levels of ecosystem organisation, from forest type (primary or secondary forest) to the trees themselves, following the folded forest model theory (Oldeman 1992, 1994). The vertical growth of trees creates a convex pattern in the relief of canopy surface, whereas gaps make concavities which delimit impact of perturbation on the forest structure. These events are either the result of the dynamic of single trees (emergent and decayed trees), or arise from the dynamic of a group of trees working together (group of emergent trees or complex gaps). At the plot scale, the elementary events, convexities and concavities, are gathered on similar topo-sequences, and form canopy units either higher or lower than the average canopy height. This study suggests that the topography of the canopy surface is defined by a complex nested system from trees, to groups of trees, to canopy units, within a delimited floristic and physical environment.     

Role of ethylene in the regulation of cell death and leaf loss in ozone-exposed European beech

To test the involvement of ethylene in mediating ozone-induced cell death and leaf loss in European beech ( Fagus sylvatica L.), tree seedlings were exposed to proportionally increased or decreased field ozone levels for up to 6 months. Ozone treatment caused cell death and accelerated leaf loss at higher than ambient levels, but had only minor effects at ambient and no effects at subambient ozone levels. The emission of ethylene, the levels of its precursor, 1-aminocyclopropane-1-carboxylate (ACC), and mRNA levels of specific ACC synthase ( FS-ACS2 ) and ACC oxidase ( FS-ACO1 ) isoforms showed a persistent increase and preceded cell death by approximately 2 weeks. Inhibition of ethylene biosynthesis led to reduced lesion formation whereas application of ACC accelerated ozone-induced cell death and leaf loss. Similar results were obtained when adult beech trees were exposed to 2 × ozone by a whole tree free-air canopy exposure system. The results suggest a role of ethylene in amplifying ozone effects under field conditions in this major European broad-leaved tree species.     

Water shortage affects the water and nitrogen balance in Central European beech forests

Whilst forest policy promotes cultivation and regeneration of beech dominated forest ecosystems, beech itself is a highly drought sensitive tree species likely to suffer from the climatic conditions prognosticated for the current century. Taking advantage of model ecosystems with cool-moist and warm-dry local climate, the latter assumed to be representative for future climatic conditions, the effects of climate and silvicultural treatment (different thinning regimes) on water status, nitrogen balance and growth parameters of adult beech trees and beech regeneration in the understorey were assessed. In addition, validation experiments with beech seedlings were carried out under controlled conditions, mainly in order to assess the effect of drought on the competitive abilities of beech. As measures of water availability xylem flow, shoot water potential, stomatal conductance as well as delta (13)C and delta (18)O in different tissues (leaves, phloem, wood) were analysed. For the assessment of nitrogen balance we determined the uptake of inorganic nitrogen by the roots as well as total N content and soluble N compounds in different tissues of adult and young trees. Retrospective and current analysis of delta (13)C, growth and meteorological parameters revealed that beech growing under warm-dry climatic conditions were impaired in growth and water balance during periods with low rain-fall. Thinning affected water, N balance and growth mostly of young beech, but in a different way under different local climatic conditions. Under cool, moist conditions, representative for the current climatic and edaphic conditions in beech forests of Central Europe, thinning improves nutrient and water status consistent to published literature and long-term experience of forest practitioners. However, beech regeneration was impaired as a result of thinning at higher temperatures and under reduced water availability, as expected in future climate.