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Forest trees are experiencing increasing frequency and intensity of drought events with climate change. We investigated xylem and phloem traits from mature Fagus sylvatica and Picea abies trees after 5 years of complete exclusion of throughfall precipitation during the growing season. Xylem and phloem anatomy, leaf and branch biomass were analysed along top branches of ~1.5 m lenght in 5 throughfall precipitation excluded (TE) and 5 control (CO) trees of both beech and spruce. Xylem traits were analysed on wood cores extracted from the stem at breast height. In the top branches of both species, the lumen diameter (or area) of xylem and phloem conduits did not differ between TE and CO trees. At breast height, TE trees of both species produced narrower xylem rings and conduits. While allocation to branch (BM) and needle biomass (LM) did not change between TE and CO in P. abies, TE F. sylvatica trees allocated proportionally more biomass to leaves (LM) than BM compared with CO. Despite artificial drought increased the mortality in the TE plots, our results revealed no changes in both xylem and phloem anatomies, undermining the hypothesis that successful acclimation to drought would primarily involve increased resistance against air embolism.

Laser scanning has revolutionized the ability to quantify single-tree morphologies and stand structural variables. In this study, we address the issue of occlusion when scanning a spruce (Picea abies (L.) H.Karst.) and beech (Fagus sylvatica L.) forest with a mobile laser scanner by making use of a unique study site setup. We scanned forest stands (1) from the ground only and (2) from the ground and from above by using a crane. We also examined the occlusion effect by scanning in the summer (leaf-on) and in the winter (leaf-off). Especially at the canopy level of the forest stands, occlusion was very pronounced, and we were able to quantify its impact in more detail. Occlusion was not as noticeable as expected for crown-related variables but, on average, resulted in smaller values for tree height in particular. Between the species, the total tree height underestimation for spruce was more pronounced than that for beech. At the stand level, significant information was lost in the canopy area when scanning from the ground alone. This information shortage is reflected in the relative point counts, the Clark–Evans index and the box dimension. Increasing the voxel size can compensate for this loss of information but comes with the trade-off of losing details in the point clouds. From our analysis, we conclude that the voxelization of point clouds prior to the extraction of stand or tree measurements with a voxel size of at least 20 cm is appropriate to reduce occlusion effects while still providing a high level of detail.

As climate change progresses, the frequency and duration of drought stress events are increasing. While the mechanisms of drought acclimation of trees has received considerable attention in recent years, the recovery processes remain critically understudied. We used a unique throughfall exclusion experiment in a mature temperate mixed forest consisting of the more isohydric Norway spruce and more anisohydric European beech, to study the recovery and resilience after drought release. We hypothesized that pre-dawn water potential (ΨPD) of both species will increase within 1 day after watering, while the recovery of stomatal conductance (gs) and the reversal of osmoregulation will be significantly delayed in the more isohydric spruce. Furthermore, we hypothesized that the xylem sap flow density (udaily) will not fully recover within the growing season due to the strong drought impact. After 5 years of summer drought, trees showed significantly reduced ΨPD, udaily and increased osmoregulation in leaves, but only isohydric spruce displayed increased leaf abscisic acid concentrations. In line with our hypothesis, ΨPD and gs recovered within 1 day in beech. Conversely, isohydric spruce showed delayed increases in ΨPD and gs. The delay in recovery of spruce was partially related to the replenishment of the stem water reservoir, as indicated by the missing response of udaily at the crown base compared with DBH level upon watering. However, udaily fully recovered only in the next growing season for beech and was still reduced in spruce. Nevertheless, in both species, osmotic acclimations of leaves were reversed within several weeks. While both species displayed full resilience to drought stress in water-related physiology, the recovery time was in several cases, e.g., udaily, ΨPD and gs, shorter for beech than for spruce. With future increases in the frequency of drought events under ongoing climate change, tree species that recover more quickly will be favored.

The role of trees in city cooling has warranted much attention based on concerns over climate change and urban expansion. Simultaneously, there is an interest in introducing species from dry habitats to cope with the increasing risks of drought under climate change. The general understanding is that the evolutionary adaptation to respective resource supplies in species' habitats affects their environmental tolerance. The physical performances of six frequently planted species, originating from two contrasting habitats, were tested in a drought experiment. We (1) investigated if species from drier habitats are more drought tolerant than species that have evolved in Central European woodlands under a temperate climate regime and (2) discussed the effect of tolerance on the cooling potential of these trees. Native species from mesic habitats maintained only 48% of their controls sap flux and of these species, Tilia cordata had the worst performance with premature leaf senescence. Species from drier habitats had less reduction in sap flux (60%) but lower stem growth, possibly favouring (fine) root development into deeper soil layers, as observed when comparing linden species. Higher stem water exploitation and stronger regulation of water use at high evaporative demand were further reaction patterns that likely helped species from dry habitats maintain good physiological functions. Therefore, even under sustained drought, we expect them to have a higher cooling capacity. As a conclusion, they should be favoured for planting in extreme urban environments. Systematic screening and testing of promising species from target habitats is recommended to diversify the choice of species.

Decreasing water availability due to climate change poses the question of whether and to what extent tree species are able to hydraulically acclimate and how hydraulic traits of stems and leaves are coordinated under drought. In a through-fall exclusion experiment, hydraulic acclimation was analyzed in a mixed forest stand of Fagus sylvatica L. and Picea abies (L.) Karst. In drought-stressed (TE, through-fall exclusion over 2 years) and control (CO) trees, hydraulic vulnerability was studied in branches as well as in leaves (F. sylvatica) and end-twigs (P. abies, entirely formed during the drought period) sampled at the same height in sun-exposed portions of the tree crown. In addition, relevant xylem anatomical traits and leaf pressure-volume relations were analyzed. The TE trees reached pre-dawn water potentials down to -1.6 MPa. In both species, water potentials at 50% loss of xylem hydraulic conductivity were ~0.4 MPa more negative in TE than in CO branches. Foliage hydraulic vulnerability (expressed as water potential at 50% loss of leaf/end-twig hydraulic conductance) and water potential at turgor loss point were also, respectively, 0.4 and 0.5 MPa lower in TE trees. Minor differences were observed in conduit mean hydraulic diameter and cell wall reinforcement. Our findings indicate significant and fast hydraulic acclimation under relatively mild drought in both tree species. Acclimation was well coordinated between branches and foliage, which might be essential for survival and productivity of mature trees under future drought periods.

Hydraulic redistribution (HR) of soil water through plant roots is a crucial phenomenon improving the water balance of plants and ecosystems. It is mostly described under severe drought, and not yet studied under moderate drought. We tested the potential of HR under moderate drought, hypothesizing that (H1) tree species redistribute soil water in their roots even under moderate drought and that (H2) neighboring plants are supported with water provided by redistributing plants. Trees were planted in split-root systems with one individual (i.e., split-root plant, SRP) having its roots divided between two pots with one additional tree each. Species were 2- to 4-year-old English oak (Quercus robur L.), European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst). A gradient in soil water potential (ψsoil) was established between the two pots (-0.55 ± 0.02 MPa and -0.29 ± 0.03 MPa), and HR was observed by labeling with deuterium-enriched water. Irrespective of species identity, 93% of the SRPs redistributed deuterium enriched water from the moist to the drier side, supporting H1. Eighty-eight percent of the plants in the drier pots were deuterium enriched in their roots, with 61 ± 6% of the root water originating from SRP roots. Differences in HR among species were related to their root anatomy with diffuse-porous xylem structure in both beech and-opposing the stem structure-oak roots. In spruce, we found exclusively tracheids. We conclude that water can be redistributed within roots of different tree species along a moderate ψsoil gradient, accentuating HR as an important water source for drought-stressed plants, with potential implications for ecohydrological and plant physiological sciences. It remains to be shown to what extent HR occurs under field conditions in Central Europe.

Key message Under severe drought, growth of Norway spruce suffered much more than European beech. Norway spruce benefited from growing in the environment of beech, and both species acclimated slightly to 5 years of experimentally extended drought. Abstract Recent studies show that the detrimental effects of drought on stand growth are mitigated when the stand contains mixed tree species. We analysed the growth responses of Norway spruce and European beech to episodic and experimentally extended drought in intra- and inter-specific mature stands. We used annual diameter growth records dating back to 1998 to determine the impact of the natural episodic drought in 2003 and 2015. To analyse extended drought, spruce and beech trees were exposed to extreme drought under automatic throughfall exclusion roofs from 2014 to 2018. The growth of spruce in an inter-specific environment with beech was 20–50% less affected by natural episodic drought compared with an intra-specific constellation. When beech grew in an inter-specific environment, it was by 23% more affected by drought compared to intra-specific conditions, but seemed to recover faster. The induced drought from 2014 to 2018 resulted in a strong growth reduction in the first year particularly for spruce, followed by a slight acclimation to the dry conditions. Beech acclimated and recovered faster than spruce across all growing conditions, while spruce only acclimatized faster in the environment of beech. Both species showed a higher mortality under induced drought compared with the controls; for spruce, the mortality rate was fivefold higher compared to the long-term mortality. The long-term moderate-growth stabilization and the growth increase after the 5-year exposure to drought suggest a gradual acclimation to drought by beech. The resistance and acclimation to drought of spruce when growing in mixture should be considered when designing resource efficient and productive mixed conifer-broadleaved stands for future climates.

Severe drought events threaten tree water transport system, productivity and survival. Woody angiosperms generally die when embolism-induced loss of hydraulic conductance (PLC) surpasses 80-90% under intense water shortage. However, the recovery capability and possible long-term carry-over effects of repeated drought events could dictate the fate of species' population under climate change scenarios. Potted saplings of European beech (Fagus sylvatica L.) were subjected to two drought cycles in two consecutive growing seasons, aiming to induce minimum leaf water potentials (Ψmd) of about -4 MPa, corresponding to hydraulic thresholds for survival of this species. In the first cycle, a well-irrigated (C) and a drought-stressed group (S) were formed, and, in the following summer, each group was divided in a well-irrigated and a drought-stressed one (four groups in total). The impact of the multiple drought events was assessed by measuring wood anatomical traits, biomass production, water relations, stem hydraulics and non-structural carbohydrate (NSC) content. We also investigated possible connections between stem hydraulics and carbon dynamics during the second drought event and following re-irrigation. S plants had lower Ψmd and maximum specific hydraulic conductivity (Ks) than C plants in the following growing season. Additionally, aboveground biomass production and leaf number were lower compared to C trees, resulting in lower water consumption. However, PLC was similar between groups, probably due to the production of new functional xylem in spring. The second drought event induced 85% PLC and promoted conversion of starch-to soluble sugars. Nevertheless, 1 week after re-irrigation, no embolism repair was observed and soluble sugars were reconverted to starch. The previous drought cycle did not influence the hydraulic performance during the second drought, and after re-irrigation S plants had 40% higher wood NSC content. Our data suggest that beech cannot recover from high embolism levels but multiple droughts might enhance stem NSC availability.

Forests are under pressure from accelerating global change. To cope with the multiple challenges related to global change but also to further improve forest management we need a better understanding of (1) the linkages between drivers of ecosystem change and the state and management of forest ecosystems as well as their capacity to adapt to ongoing global environmental changes, and (2) the interrelationships within and between the components of forest ecosystems. To address the resulting challenges for the state of forest ecosystems in Central Europe, we suggest 45 questions for future ecological research. We define forest ecology as studies on the abiotic and biotic components of forest ecosystems and their interactions on varying spatial and temporal scales. Our questions cover five thematic fields and correspond to the criteria selected for describing the state of Europe’s forests by policy makers, i.e. biogeochemical cycling, mortality and disturbances, productivity, biodiversity and biotic interactions, and regulation and protection. We conclude that an improved mechanistic understanding of forest ecosystems is essential for the further development of ecosystem-oriented multifunctional forest management in the face of accelerating global change. Forests are under pressure from accelerating global change. To cope with the multiple challenges related to global change but also to further improve forest management we need a better understanding of (1) the linkages between drivers of ecosystem change and the state and management of forest ecosystems as well as their capacity to adapt to ongoing global environmental changes, and (2) the interrelationships within and between the components of forest ecosystems. To address the resulting challenges for the state of forest ecosystems in Central Europe, we suggest 45 questions for future ecological research. We define forest ecology as studies on the abiotic and biotic components of forest ecosystems and their interactions on varying spatial and temporal scales. Our questions cover five thematic fields and correspond to the criteria selected for describing the state of Europe’s forests by policy makers, i.e. biogeochemical cycling, mortality and disturbances, productivity, biodiversity and biotic interactions, and regulation and protection. We conclude that an improved mechanistic understanding of forest ecosystems is essential for the further development of ecosystem-oriented multifunctional forest management in the face of accelerating global change.

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.

Under changing climatic conditions, drought may become a critical constraint for trees in urban areas, particularly at roadsides and highly paved squares. As healthy urban trees have proven to be an important mitigation and adaptation tool for climate change as well as a significant provider of ecosystem services, there is a need for planting species and cultivars capable of coping with the limited water supply. However, data on species&rsquo, and cultivars&rsquo, response to drought, particularly their water supplying root systems remains rare. To consider the whole plant responses to drought situations, we studied the growth and phenology of three frequently planted tree species and cultivars with a diameter of 5&ndash, 6 cm during a one-year rainfall exclusion experiment conducted in a nursery field as well as the dry biomass of the compartments branch, stem, and root after excavation. Our results revealed that species&rsquo, performance were linked to their within-plant carbon partitioning. A high tolerance to drought was noted for Acer campestre, with a particularly high ratio of root:shoot ratio, which made it presumably less susceptible to droughts. Tilia cordata &lsquo, Greenspire&rsquo, was highly affected by the reduced water availability visible through prematurely leaf senescence, while Carpinus betulus &lsquo, Fastigiata&rsquo, suffered from losing a considerable part of its root biomass, which resulted in the lowest root:shoot ratio of all species and cultivars. This study demonstrated the need for investigating the reaction patterns of species and cultivars by considering both the above-and the below-ground plant parts. We recommend that, for future tree plantings at harsh and challenging urban sites, an important selection criterion should be species&rsquo, capability to develop and retain strong and dense root systems even under limited water supply, as that is believed to be an important trait for drought tolerance.

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.

• 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.

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.

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.

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.

Fine root dynamics (diameter < 1 mm) in mature Fagus sylvatica, with the canopies exposed to ambient or twice-ambient ozone concentrations, were investigated throughout 2004. The focus was on the seasonal timing and extent of fine root dynamics (growth, mortality) in relation to the soil environment (water content, temperature). Under ambient ozone concentrations, a significant relationship was found between fine root turnover and soil environmental changes indicating accelerated fine root turnover under favourable soil conditions. In contrast, under elevated ozone, this relationship vanished as the result of an altered temporal pattern of fine root growth. Fine root survival and turnover rate did not differ significantly between the different ozone regimes, although a delay in current-year fine root shedding was found under the elevated ozone concentrations. The data indicate that increasing tropospheric ozone levels can alter the timing of fine root turnover in mature F. sylvatica but do not affect the turnover rate.

The effects of elevated O3 on photosynthetic properties in adult beech trees (Fagus sylvatica) were investigated in relation to leaf mass per area as a measure of the gradually changing, within-canopy light availability. Leaves under elevated O3 showed decreased stomatal conductance at unchanged carboxylation capacity of Rubisco, which was consistent with enhanced delta 13C of leaf organic matter, regardless of the light environment during growth. In parallel, increased energy demand for O3 detoxification and repair was suggested under elevated O3 owing to enhanced dark respiration. Only in shade-grown leaves,light-limited photosynthesis was reduced under elevated O3, this effect being accompanied by lowered F(v)/F(m). These results suggest that chronic O3 exposure primarily caused stomatal closure to adult beech trees in the field regardless of the within-canopy light gradient. However, light limitation apparently raised the O3 sensitivity of photosynthesis and accelerated senescence in shade leaves.

Summary Three-year-old Alnus glutinosa (L.) Gaertn. (alder) saplings were single or double inoculated at the stem base with Phytophthora alnisubsp. alni Brasier & S.A. Kirk under natural climatic conditions. Lesion formation on the bark showed a biphasic pattern of development, with extension occurring at a moderate rate in spring, and more rapidly during late summer. However, large variability was encountered in pathogen development within the population of infected saplings, ranging from high susceptibility to almost complete resistance. Infection resulted in severe growth retardation, and death within two years of inoculation in 75% of the saplings. During disease development, rates of transpiration and CO2 uptake were significantly reduced. Consequently, minimum leaf water potentials were less negative in infected saplings than in control saplings. Surviving saplings matched control trees in photosynthetic capacity, transpiration rate and water potential during the second year of infection. Leaf starch concentration of infected saplings was significantly higher than in control saplings, possibly indicating that the destruction of bark tissue by the pathogen impaired phloem transport from leaves to roots.

One of the decisive factors for the survival of forest trees is how well they can compete against their tree neighbours

The responsiveness of adult beech and spruce trees to chronic O3 stress was studied at a free-air O3 exposure experiment in Freising/Germany. Over three growing seasons, gas exchange characteristics, biochemical parameters, macroscopic O3 injury and the phenology of leaf organs were investigated, along with assessments of branch and stem growth as indications of tree performance. To assess response pattern to chronic O3 stress in adult forest trees, we introduce a new evaluation approach, which provides a comprehensive, readily accomplishable overview across several tree-internal scaling levels, different canopy regions and growing seasons. This new approach, based on a three-grade colour coding, combines statistical analysis and the proficient ability of the ‘‘human eye’’ in pattern recognition. 2005 Elsevier Ltd. All rights reserved.

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 acceler- ated 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.

Under the site conditions of a temperate forest, the exchange of short-chained oxygenated carbonyls (aldehydes, ketones) was assessed from leaves of adult European beech trees. The crowns of the trees were either exposed to an elevated O 3 regime as released by a free-air fumigation system (2 x O 3 ) or to the unchanged O 3 regime at the site (1 x O 3 , 'control'). Daily fluctuations of oxygenated carbonyls were quantified in relation to environmental and physiological factors. In particular, the effect of O 3 on carbonyl exchange was studied. Measurements of leaf gas exchange were performed with a dynamic cuvette system, and carbonyl fluxes were determined using 2,4-dinitrophenylhydrazine (DNPH)-coated silica gel cartridges. Leaves mainly emitted acetaldehyde, formaldehyde and acetone. Acetaldehyde dominated the emissions, amounting up to 100 nmol m -2 min -1 , followed by formaldehyde (approximately 80 nmol m -2 min -1 ) and acetone (approximately 60 nmol m -2 min -1 ). Carbonyl emissions were highest during midday and significantly lowered at night, irrespective of the O 3 exposure regime. Trees exposed to 2 x O 3 emitted acetaldehyde and acetone at enhanced rates. The findings are of particular significance for future climate change scenarios that assume increased O 3 levels.

Das Uberleben von Waldbaumen hangt auch davon ab, ob sie sich gegen ihre Nachbarpflanzen durchsetzen konnen

The current AOT40 concept for inferring risks in forest trees by ozone (O3) injury is based on an accumulated external O3 exposure rather than an internal O3 dose or uptake rate. AOT40 assumes O3 concentrations below 40 nl l−1 and night-time exposure to be negligible. Hence, this concept is rather inconsistent with observed forest conditions. In contrast, the flux concept of cumulative O3 uptake (CU) into the leaves has the potential of reflecting a physiologically meaningful internal O3 dose experienced by trees. In this paper, we relate AOT40 to cumulative O3 uptake into European beech (Fagus sylvatica), Norway spruce (Picea abies), European larch (Larix decidua) and cembran pine (Pinus cembra) trees differing in size, age and site conditions. We demonstrate that the flux concept can be extended to the tree and the stand level, making use of sap flow measurements through tree trunks. Although in both seedlings and adult trees AOT40 may show some linearity in correlations with average CU, the latter varies, at given AOT40, by 25±11% within and between species. This is because O3 flux is primarily influenced by stomatal aperture, the latter being affected by climate, canopy position, leaf and tree age while varying between species. In particular, if weighed by detoxification capacity, we suggest, therefore, O3 uptake related air quality indices to be promoted towards ecologically meaningful standards in forest protection, overcoming the shortcomings of exposure concepts. As O3 injury results from the balance between O3 uptake and detoxification in the leaf mesophyll, we conclude the flux concept in combination with measures of biochemical defence to have the capacity for predicting tree response to O3 stress.

International audience; Microclimate and tree age have been suggested to be factors influencing the defense capacity against oxidative stress. Therefore, 5-year-old Fagus sylvatica seedlings were grown on a scaffolding in the sun and shade crown of 55-year-old trees throughout one growing season. Independent of tree age sun leaves had a lower specific leaf area, lower pigment contents and a more capacitive antioxidative system than shade leaves. In addition, in the sun crown leaves of seedlings displayed a higher specific leaf area than leaves of adult trees. Age dependent changes in leaf morphology were related to changes in the defense capacity against oxidative stress, with area based concentrations of antioxidants and pigments increasing with tree age. Thus our results suggest that differences in the response to oxidative stress may be attributed to age and crown position related differences in the specific leaf area, the latter influencing the biochemical and physiological performance of Fagus sylvatica leaves.; L'influence du microclimat et de l'âge de l'arbre sur la capacité défensive du hêtre européen (Fagus sylvatica L.) contre le stress oxidatif. Le microclimat et l'âge de l'arbre sont considerés comme des facteurs qui influencent la capacité défensive contre le stress oxidatif. Par conséquent, des plants de Fagus sylvatica de cinq ans ont grandi sur un échafaudage à la couronne des arbres âgés de 55 ans, une fois au soleil, une autre fois à l'ombre pendant une période de croissance. Indépendamment de l'âge de l'arbre, le feuillage exposé au soleil avait une surface spécifique de la feuille inférieure, un contenu de pigment inférieur et un système antioxidatif plus puissant que le feuillage à l'ombre. De plus, au niveau de la couronne la plus haute, le feuillage des plants a développé une surface spécifique plus élevée que celui des arbres adultes. Des transformations de morphologie du feuillage, qui dépendent de l'âge des feuilles, ont été comparées avec des transformations de la capacité contre le stress oxidatif. À cela la teneur en substances antioxidatives relative à la surface de la feuille a augmenté avec l'âge de l'arbre. Donc nos résultats suggèrent que les différences de réaction au stress oxidatif peuvent être attribuées aux différences de la surface spécifique de la feuille - dépendant de l'âge et de la position à la couronne - ce qui influence la performance biochimique et physiologique des feuilles du Fagus sylvatica.

In mature trees of Picea abies, Fagus sylvatica and Quercus robur, photosynthesis and transpiration were assessed in response to the total solar eclipse that occurred in Central Europe during the late morning hours of August 11, 1999, a day with changing cloudiness. Measurements were conducted at three forest sites located in the totality zone and the 99% area of the eclipse within a radius of about 100 km around the city of Munich (southern Germany). The eclipse lasting 164 minutes lowered the photosynthetic photon flux density (PPFD) to about 1 μmol m -2 s -1 during the 2-minute totality period, when the sky was clear. During totality, photosynthesis was reduced to an extent that allowed CO 2 release to dominate the gas exchange of leaves. Effects on transpiration were less pronounced as the totality was apparently too short to induce distinct stomatal closure in response to low PPFD. Transpiration was strongly reduced, however, by increased air humidity and wet leaf surfaces during sporadic rain showers which preceded or succeeded the eclipse during the same day, whereas low PPFD through intermittent cloudiness during rain only moderately reduced photosynthesis. Although transpiration was lowered to a minor extent only by the eclipse, the latter affected the water transport through the whole tree, as reflected in a decline in the sap flow rate through the basal stem part with a time delay depending on the species. Nevertheless, trees responded in a synchronous way, regardless of the site, species or the percent degree of the eclipse.

summary Young trees of European beech (Fagus sylvatica) acclimated for one growing season to ambient (c. 367 l ll ’" )o r elevated CO # levels (c. 660 l ll ’") were exposed during the subsequent year to combinations of the same CO # regimes and ambient or twice-ambient ozone (O $ ) levels (generated from the database of a rural site). By the end of June, before the development of macroscopic leaf injury, the raised O $ levels had not aected the light and dark reactions of photosynthesis. However, acclimation to elevated CO # had resulted in lowered chlorophyll and nitrogen concentrations, whereas photosynthetic performance, examined over a wide range of parameters from light and dark reactions, remained unchanged or showed only slight reductions (e.g. apparent electron transport rate, ETR; apparent quantum yield of CO # gas exchange, U CO# ; apparent carboxylation eciency, CE; and photosynthetic capacity at light and CO # saturation, PC). In August, after the appearance of leaf necroses, plants grown under ambient CO # and twice-ambient O $ conditions declined in both the photosynthetic light reactions (optimum electron quantum yield, F v }F m , non-photochemical energy quenching, NPQ, reduction state of Q A , apparent electron quantum yield, U PSI I, maximum electron transport rates) and the dark reactions as reflected by CE, U CO# , as well as the maximum CO # uptake rate (i.e. PC). CE, U CO# and PC were reduced by c. 75, 40 and 75%, respectively, relative to plants exposed to ambient CO # and O $ levels. By contrast, plants exposed to twice-ambient O $ and elevated CO # levels maintained a photosynthetic performance similar to individuals grown either under ambient CO # and ambient O $ , or elevated CO # and ambient O $ conditions. The long-term exposure to elevated CO # therefore tended to counteract adverse chronic eects of enhanced O $ levels on photosynthesis. Possible reasons for this compensatory eect in F. sylvatica are discussed.

Spatio-temporally consistent O 3 doses are demonstrated in adult Fagus sylvatica from the Kranzberg Forest free-air fumigation experiment, covering cross-canopy and whole-seasonal scopes through sap flow measurement. Given O 3 -driven closure of stomata, we hypothesized enhanced whole-tree level O 3 influx to be prevented under enhanced O 3 exposure. Although foliage transpiration rate was lowered under twice-ambient O 3 around noon by 30% along with canopy conductance, the hypothesis was falsified, as O 3 influx was raised by 25%. Nevertheless, the twice-ambient/ambient ratio of O 3 uptake was smaller by about 20% than that of O 3 exposure, suggesting stomatal limitation of uptake. The O 3 response was traceable from leaves across branches to the canopy, where peak transpiration rates resembled those of shade rather than sun branches. Rainy/overcast-day and nightly O 3 uptake is quantified and discussed. Whole-seasonal canopy-level validation of modelled with sap flow-derived O 3 flux becomes available in assessing O 3 risk for forest trees.

Drought reduces the carbon (C) assimilation of trees and decouples aboveground from belowground carbon fluxes, but little is known about the response of drought-stressed trees to rewetting. This study aims to assess dynamics and patterns of C allocation in beech saplings under dry and rewetted soil conditions. In October 2010, 5-year-old beech saplings from a forest site were transplanted into 20 l pots. In 2011, the saplings were subjected to different levels of soil drought ranging from non-limiting water supply (control) to severe water limitation with soil water potentials of less than -1.5 MPa. As a physiologically relevant measure of drought, the cumulated soil water potential (i.e., drought stress dose (DSD)) was calculated for the growing season. In late August, the saplings were transferred into a climate chamber and pulse-labeled with (13)C-depleted CO2 (δ(13)C of -47‰). Isotopic signatures in leaf and soil respiration were repeatedly measured. Five days after soil rewetting, a second label was applied using 99 atom% (13)CO2. After another 12 days, the fate of assimilated C in each sapling was assessed by calculating the (13)C mass balance. Photosynthesis decreased by 60% in saplings under severe drought. The mean residence time (MRT) of recent assimilates in leaf respiration was more than three times longer than under non-limited conditions and was positively correlated to DSD. Also, the appearance of the label in soil respiration was delayed. Within 5 days after rewetting, photosynthesis, MRT of recent assimilates in leaf respiration and appearance of the label in soil respiration recovered fully. Despite the fast recovery, less label was recovered in the biomass of the previously drought-stressed plants, which also allocated less C to the root compartment (45 vs 64% in the control). We conclude that beech saplings quickly recover from extreme soil drought, although transitional after-effects prevail in C allocation, possibly due to repair-driven respiratory processes.

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.

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.

Ground-level ozone (O3) has gained awareness as an agent of climate change. In this respect, key results are comprehended from a unique 8-year free-air O3-fumigation experiment, conducted on adult beech (Fagus sylvatica) at Kranzberg Forest (Germany). A novel canopy O3 exposure methodology was employed that allowed whole-tree assessment in situ under twice-ambient O3 levels. Elevated O3 significantly weakened the C sink strength of the treesoil system as evidenced by lowered photosynthesis and 44% reduction in whole-stem growth, but increased soil respiration. Associated effects in leaves and roots at the gene, cell and organ level varied from year to year, with drought being a crucial determinant of O3 responsiveness. Regarding adult individuals of a late-successional tree species, empirical proof is provided first time in relation to recent modelling predictions that enhanced ground-level O3 can substantially mitigate the C sequestration of forests in view of climate change. Empirical proof corroborates substantial mitigation of carbon sequestration in the treesoil system of a forest site under enhanced O3 impact for adult beech.

We examined the effect of ozone (O(3)) on Norway spruce (Picea abies) needle epicuticular wax over three seasons at the Kranzberg Ozone Fumigation Experiment. Exposure to 2x ambient O(3) ranged from 64.5 to 74.2 microl O(3) l(-1) h AOT40, and 117.1 to 123.2 nl O(3) l(-1) 4th highest daily maximum 8-h average O(3) concentration. The proportion of current-year needle surface covered by wax tubes, tube aggregates, and plates decreased (P=0.011) under 2x O(3). Epistomatal chambers had increased deposits of amorphous wax. Proportion of secondary alcohols varied due to year (P=0.004) and O(3) treatment (P=0.029). Secondary alcohols were reduced by 9.1% under 2x O(3). Exposure to 2x O(3) increased (P=0.037) proportions of fatty acids by 29%. Opposing trends in secondary alcohols and fatty acids indicate a direct action of O(3) on wax biosynthesis. These results demonstrate O(3)-induced changes in biologically important needle surface characteristics of 50-year-old field-grown trees.

Combined delta(13)C and delta(18)O analyses of leaf material were used to infer changes in photosynthetic capacity (A(max)) and stomatal conductance (g(l)) in Fagus sylvatica and Picea abies trees growing under natural and controlled conditions. Correlation between g(l) and delta(18)O in leaf cellulose (delta(18)O(cel)) allowed us to apply a semi-quantitative model to infer g(l) from delta(18)O(cel) and also interpret variation in delta(13)C as reflecting variation in A(max). Extraction of leaf cellulose was necessary, because delta(18)O from leaf organic matter (delta(18)O(LOM)) and delta(18)O(cel) was not reliably correlated. In juvenile trees, the model predicted elevated carbon dioxide (CO(2)) to reduce A(max) in both species, whereas ozone (O(3)) only affected beech by reducing CO(2) uptake via lowered g(l). In adult trees, A(max) declined with decreasing light level as g(l) was unchanged. O(3) did not significantly affect isotopic signatures in leaves of adult trees, reflecting the higher O(3) susceptibility of juvenile trees under controlled conditions. The isotopic analysis compared favourably to the performance of leaf gas exchange, underlining that the semi-quantitative model approach provides a robust way to gather time-integrated information on photosynthetic performance of trees under multi-faced ecological scenarios, in particular when information needed for quantitative modelling is only scarcely available.

Whole-tree O3 uptake was exemplified for Picea abies, Fagus sylvatica and Larix decidua in stands at high and low altitude and contrasting water availability through sap flow measurement in tree trunks, intrinsically accounting for drought and boundary layer effects on O3 flux. O3 uptake of evergreen spruce per unit foliage area was enhanced by 100% at high relative to low elevation, whereas deciduous beech and larch showed similar uptake regardless of altitude. The responsiveness of the canopy conductance to water vapor and, as a consequence, O3 uptake to soil moisture and air humidity did not differ between species. Unifying findings at the whole-tree level will promote cause–effect based O3 risk assessment and modeling.

In Central Europe, Fagus sylvatica and Picea abies represent contrasting extremes in foliage type, crown structure and length of growing season. In order to examine the competitive strategies of these two co-occurring species, we tested the following hypotheses: (1) the space occupied by the foliage of sun branches is characterized by greater foliar mass investment compared to shade branches, (2) the carbon (C) gain per unit of occupied space is greater in sun than in shade branches, and (3) annual C and water costs of the foliage for sustaining the occupied space are low, wherever C gain per unit of occupied space is low. These were investigated in a mature forest in Southern Germany. The examination was based on the annual assessment of space-related resource investments and gains of the foliage. The foliated space around branches was regarded as the relevant volume with respect to aboveground resource availability. Occupied crown space per standing foliage mass was higher in shade compared to sun branches of beech, whereas no difference existed in crown volume per foliage mass between sun and shade branches of spruce (hypothesis 1 accepted for beech but rejected for spruce). However, beech occupied more space per foliage mass than spruce. The C gain per occupied crown volume was greater in sun than in shade branches (hypothesis 2 accepted) but did not differ between species. The amount of occupied space per respiratory and transpiratory costs did not differ between species or between sun and shade branches. In beech and spruce, the proportion of foliage investment in the annual C balance of sun and shade branches remained rather stable, whereas respiratory costs distinctly increased in shade foliage. Hence, shade branches were costly structures to occupy space, achieving only low and even negative C balances (rejection of hypothesis 3), which conflicts with the claimed C autonomy of branches. Our findings suggest that competitiveness is determined by the standing foliage mass and the annual branch volume increment rather than annual investments in foliage. Expressing competitiveness in terms of space-related resource investments versus returns, as demonstrated here, has the potential of promoting mechanistic understanding of plant–plant interactions.

Cumulative ozone (O3) uptake and O3 flux were related to physiological, morphological and biochemical characteristics of Norway spruce (Picea abies (L.) Karst.) trees of different ages. Under ambient CO2 conditions, photosynthetic capacity (Amax) declined in mature trees when cumulative O3 uptake into needles, which provides a measure of effective O3 dose, exceeded 21 mmol m-2 of total needle surface area. A comparable decline in Amax of seedlings occurred when cumulative O(3) uptake was only 4.5 mmol m-2. The threshold O3 flux causing a significant decline in Amax ranged between 2.14 and 2.45 nmol m-2 s-1 in mature trees and seedlings subjected to exposure periods of > or = 70 and > or = 23 days, respectively. The greater O3 sensitivity of young trees compared with mature trees was associated with needle morphology. Biomass of a 100-needle sample increased significantly with tree age, whereas a negative correlation was found for specific leaf area, these changes parallel those observed during differentiation from shade-type to sun-type needles with tree ontogeny. Age-dependent changes in leaf morphology were related to changes in detoxification capacity, with area-based concentrations of ascorbate increasing during tree ontogeny. These findings indicate that the extent of O3-induced injury is related to the ratio of potentially available antioxidants to O3 influx. Because this ratio, when calculated for ascorbate, increased with tree age, we conclude that the ratio may serve as an empirical basis for characterizing age-related differences in tree responses to O3.

The total solar eclipse of August 11, 1999 provided a unique opportunity to observe the input of fast day-night and night-day transitions, under high solar elevation around noon, on the earth-atmosphere-biosphere system. Within the interdisciplinary e eld campaign BAYSOFI, measurements of radiation, boundary layer micrometeorology and photochemistry, photosynthesis and transpiration were carried out at Freising-Weihenstephan and several locations nearby focusing on short-term effects of the eclipse. Although the overall grosswetterlage on August 11 was not favourable for viewing the eclipse, with clouds covering most of central Europe, observational conditions at Weihenstephan were fair due to a large hole in the cloud layer which appeared just half hour before totality lasting for more than one hour. Thus signie cant effects of the eclipse on radiation, photolysis rates, OH, the temperature, wind, turbulence structure and stratie cation, ozone and CO 2 e uxes, photosynthesis, transpiration and sap e ow of trees could be observed which are reported and discussed in the following sequence of papers.