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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.

Mixed stands of European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst.) frequently over-yield, when compared to respective monospecific stands. Over-yielding is attributed to enhanced resource uptake efficiency through niche complementarity alleviating species competition, for example through enhanced root stratification in mixture. Under severe and frequent summer drought, however, water limitation may become crucial in modifying the prevailing competitive interaction in mixed beech-spruce forests. We hypothesize, therefore, that under drought (H I) inter-specific interaction with beech reduces water accessibility for spruce more than intra-specific conditions, thus (H II) exacerbating drought susceptibility of spruce in terms of reduced photosynthesis and stem growth. Reactions at the organ (leaf, fine root), tree and stand scale were analysed in a mature forest with beech-spruce group mixture. Under inter-specific conditions spruce’s fine-root production and depth of water uptake (assessed via δ18O of xylem water) shifted to shallow, drought-prone soil horizons, in agreement with H I. Overall, lowered fine root production and ramification along with a reduction in long-distance explorative ectomycorrhizal types resulted in decreased soil exploitation in spruce when growing together with beech. Spruce’s drought sensitivity was exemplified by a distinct decrease in stomatal conductance, net CO2 uptake rate and stem growth during periods of water limitation. Notwithstanding, species interaction effects were absent in leaf gas exchange and stem diameter growth, during a six-week summer drought period in 2013 as well as in the extremely dry year of 2003, hence rejecting H II. Based on results from soil moisture measurements and water uptake depth, we interpret the conflicting findings for H I and H II to result from: (i) seasonal shifts between positive (during spring drought) and negative (during summer drought) effects of beech neighbourhood on soil water availability for spruce, possibly overriding each other in their effect on annual stem diameter growth and (ii) the group-wise mixture pattern, where spruce is exposed to competition with beech only along group edges, i.e. laterally only, so that the putatively adverse beech effect on water accessibility stays limited. Our results suggest, compared to single tree mixture, group-wise mixture of beech and spruce to be a favourable silvicultural option in the face of climate change.

Root trenching of Norway spruce and European beech caused no long-term decrease of stem growth and only temporal growth reductions in European beech. Root volume reduction is a common occurrence in silvicultural, urban, and scientific environments, often the direct result of trenching applications. Although this interference is widespread near forest roads and in rural and urban areas, trees’ growth reactions to these limitations have rarely been analysed. Here, we compare the growth of 81 mature Norway spruce (Picea abies [L.] Karst) and European beech (Fagus sylvatica [L.]) trees, trenched for an experiment down to 1 m depth in a Luvisol soil (Parabraunerde) in 2010, and compared with 69 untrenched trees. The trenching was conducted prior to a long-term precipitation-exclusion treatment in a temperate mixed forest in Southern Germany. Stem and crown sizes, tree positions, distance to the trench, and annual stem growth of the 150 trees were measured from 2006 through 2013. Statistical analysis revealed that (1) stem growth reduction does not correlate with the loss of rooting area by trenching, (2) trenching does not significantly decrease the stem growth in the long-term post-trenching application, and (3) European beech showed a greater reaction in the first years after trenching as indicated by a greater decrease in stem growth. The relationship between stem growth reduction and the loss of rooting area arises during the first year post-trenching, but diminishes by the third year. The trenching effect on stem growth was strongest in trees at the plot edge and diminished towards the centre of the plots, where the actual study trees of the precipitation-exclusion experiment were located. The relevance of trenching for growth reduction is discussed, including potential underlying mechanisms and conclusions about experimental interferences.

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.

Climate models predict increasing frequency and intensity of summer drought events for Central Europe. In a field experiment, we investigated the response of young beech (Fagus sylvatica L.) to extreme and repeated summer drought and the modulation of drought response patterns along the natural gradient of light availability at the study site. In autumn 2008, two-year-old, nursery derived beech – as used for forest conversion practices – was planted under a Norway spruce stand primarily opened through winter storm. Precipitation was manipulated in the growing seasons of 2009 through 2011, inducing a pronounced gradient of water availability. Individual drought-stress doses (DSD) and light doses (LD) were calculated for each beech sapling during the three growing seasons. Plant growth, CO2-assimilation rate and stomatal conductance were reduced with increasing drought stress, but facilitated by increasing light availability. Progressive acclimation to water and light limitation during the three years of the experiment led to a decreased drought and shade sensitivity of diameter growth. Water-use efficiency, root/shoot ratio and rooting depth, were increased with decreasing water availability. Mean fine root diameter and specific fine root length correlated positively with both DSD and LD. Proceeding low-light acclimation was indicated by progressively increasing specific leaf area and reduced leaf dark-respiration. Present results suggest that nursery-induced high-light acclimation of the beech saplings, exacerbated light limitation upon transplant and hence productivity decline under co-occurring water limitation.

We hypothesize that reductions in stand density through thinning improve the recovery of radial stem growth in Norway spruce trees ( Picea abies ) from severe drought. However, thinning may not lead to higher relative radial growth during drought. Annual stem growth and stable carbon and oxygen isotopes in early- and latewood were assessed in trees from heavily thinned (HT), moderately thinned (MT) and un-thinned control stands at two sites in southern Germany. Physiological performance of trees as inferred from stable isotope analysis was used to interpret annual stem growth in response to the drought events in 1976 and 2003. Only in recently thinned stands, trees maintained growth probably through higher soil water availability during the drought year when compared to controls. In contrast, thinning improved the growth recovery in the years following the drought irrespective of the time span between thinning and drought. We conclude that thinning improves drought recovery response in the short and long term and should be considered as an effective management strategy to increase drought tolerance of Norway spruce stands.

Soil drought influences the C turnover as well as the fine-root system of tree saplings. Particularly during the period of establishment, the susceptibility to drought stress of saplings is increased because of incompletely developed root systems and reduced access to soil water. Here, we subjected beech saplings (Fagus sylvatica L.) to different levels of drought stress. Beech saplings were planted in rhizotrons, which were installed in the soil of a Norway spruce forest before bud burst. Soil moisture was manipulated in the following year during May to September. We measured photosynthetic net CO2 uptake, volume production of fine roots, and rhizosphere respiration during the growing season. Biometric parameters of the fine-root system, biomass, and nonstructural carbohydrates were analyzed upon harvest in October. Photosynthesis and rhizosphere respiration decreased with increasing drought-stress dose (cumulated soil water potential), and cumulative rhizosphere respiration was significantly negatively correlated with drought-stress dose. Fine-root length and volume production were highest at moderate soil drought, but decreased at severe soil drought. The proportion of fine-roots diameter < 0.2 mm and the root-to-shoot ratio increased whereas the live-to-dead ratio of fine roots decreased with increasing drought-stress dose. We conclude that the belowground C allocation as well as the relative water-uptake efficiency of beech saplings is increased under drought.

• 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.

Since the year 2000 mature beech and spruce trees were treated in a field experiment with double ambient ozone concentrations. Elevated ozone had no influence on average single leaf biomass and there were also no ozone effects on leaf nutrient concentrations in climatic normal years. However, the extraordinary dry summer 2003 triggered significant differences between the fumigated and control trees. For beech in the year after the drought event the control trees surprisingly had significantly lower foliar levels of K and P than in former years, whereas the ozone exposed trees showed no significant nutritional effects. There are indications, that the trees exposed to double ambient ozone were already adapted to higher ozone values, whereas the control trees experienced extraordinary high ambient ozone concentrations in the dry and sunny summer 2003. For spruce in autumn 2003 and 2004 ozone treated trees had significantly higher foliar levels of K in current year needles than control trees, an effect which cannot be thoroughly interpreted yet on the basis of the dataset available.

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.

We present a field study on the drought effects on total soil respiration (SRt) and its components, i.e., “autotrophic” (SRa: by roots/mycorrhizosphere) and “heterotrophic” respiration (SRh: by microorganisms and soil fauna in bulk soil), in a mature European beech/Norway spruce forest. SRa and SRh were distinguished underneath groups of beech and spruce trees using the root exclusion method. Seasonal courses of SRa and SRh were studied from 2002 to 2004, with the summer of 2003 being extraordinarily warm and dry in Central Europe. We (1) analyzed the soil temperature (T s) and moisture sensitivity of SRa and SRh underneath both tree species, and (2) examined whether drought caused differential decline of SRa between spruce and beech. Throughout the study period, SRa of beech accounted for 45–55% of SRt, independent of the soil water regime; in contrast, SRa was significantly reduced during drought in spruce, and amounted then to only 25% of SRt. In parallel, fine-root production was decreased during 2003 by a factor of six in spruce (from 750 to 130 mg l−1 a−1), but remained at levels similar to those in 2002 in beech (about 470 mg l−1 a−1). This species-specific root response to drought was related to a stronger decline of SRa in spruce (by about 70%) compared to beech (by about 50%). The sensitivity of SRa and SRh to changing T s and available soil water was stronger in SRa than SRh in spruce, but not so in beech. It is concluded that SRa determines the effect of prolonged drought on the C efflux from soil to a larger extent in spruce than beech, having potential implications for respective forest types.

The extraordinary drought during the summer of 2003 in Central Europe allowed to examine responses of adult beech trees (Fagus sylvatica) to co-occurring stress by soil moisture deficit and elevated O3 levels under forest conditions in southern Germany. The study comprised tree exposure to the ambient O3 regime at the site and to a twice-ambient O3 regime as released into the canopy through a free-air O3 fumigation system. Annual courses of photosynthesis (Amax), stomatal conductance (gs), electron transport rate (ETR) and chlorophyll levels were compared between 2003 and 2004, the latter year representing the humid long-term climate at the site. ETR, Amax and gs were lowered during 2003 by drought rather than ozone, whereas chlorophyll levels did not differ between the years. Radial stem increment was reduced in 2003 by drought but fully recovered during the subsequent, humid year. Comparison of AOT40, an O3 exposure-based risk index of O3 stress, and cumulative ozone uptake (COU) yielded a linear relationship throughout humid growth conditions, but a changing slope during 2003. Our findings support the hypothesis that drought protects plants from O3 injury by stomatal closure, which restricts O3 influx into leaves and decouples COU from high external ozone levels. High AOT40 erroneously suggested high O3 risk under drought. Enhanced ozone levels did not aggravate drought effects in leaves and stem.

Seasonal fluxes of CO2 from soil and the contribution of autotrophic (root + mycorrhizal) to total soil respiration (SR) were estimated for a mixed stand of European beech (Fagus sylvatica) and Norway spruce (Picea abies) in Central Europe. Mature trees of each species were girdled in August 2002 to eliminate carbohydrate allocation to roots. SR was measured at distances of 0.5, 1.0, and 1.5/2.0 m from the bole of each tree at 1–2 weeks intervals throughout the fall of 2002 and monthly during the spring and summer of 2003. The contribution of roots and mycorrhizae to total SR was estimated by the decrease in SR compared to ungirdled control trees to account for seasonal patterns evident in controls. SR decreased with soil temperature in the fall 2002 and increased again in 2003 as soil warmed. During most of the study period, SR was strongly related to soil temperature. During the dry summer of 2003, however, SR appeared to be uncoupled from temperature and was strongly related to soil water content (SWC). Mean rates of SR in beech and spruce control plots as well as root densities did not show a clear pattern with distance from the bole. SR decreased to levels below controls in beech within a few days after girdling, whereas spruce did not show a significant decrease until October 2002, 6 weeks after girdling. In both beech and spruce, decreased SR in response to girdling was greatest closest to the bole, possibly reflecting increased mycorrhizal activity close to the bole. Autotrophic respiration was estimated in beech to be as much as 50% of the total SR in the stand. The contribution of autotrophic respiration was less certain for spruce, although close to the bole, the autotrophic fraction may contribute to total SR as much as in beech. The large fraction of autotrophic respiration in total SR requires better understanding of tree level stresses that affect carbon allocation below ground.

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.

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.

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.

To study the single and combined effects of elevated carbon dioxide (CO2), ozone (O3), nitrogen nutrition, and water supply on photosynthetic gas exchange and biomass accumulation of Norway spruce, a four-factorial experiment was conducted in closed environmental chambers. Each factor was applied at two levels: (i) ambient and elevated (ambient + 200 μl 1-1) CO2, (ii) 20 and 80 nl 1-1 O3, (iii) low and high nitrogen fertilization, and (iv) a well watered and a drought treatment. Neither elevated O3 nor CO2 significantly changed stomatal conductances of spruce needles. Adverse effects of elevated O3 on photosynthetic parameters such as net assimilation rate and carboxylation efficiency occurred only when the plants were well watered and in a good nutritional status. After 6 weeks enhanced atmospheric CO2 resulted in increased net assimilation rates provided that nutrition was well balanced and plants were well watered. Acclimation processes became apparent and are interpreted as a consequence of sink regulation. While O3-effects were apparent only in biomass of 1-year-old plant material, elevated CO2 resulted in higher biomass of the buds expanding during the exposure and increased root biomass significantly. Above and below-ground biomass were strongly influenced by the water and nutrition treatments.

Summary The aim of the present study was to address the complex interactions of environmental constraints, ozone and drought stress, with elevated atmospheric CO 2 on the activities of antioxidative enzymes and soluble protein contents in needles of Norway spruce trees ( Picea abies L.). Five-year-old spruce trees were kept from bud break in June until January of the following year in phytochambers under climatic conditions similar to those of a natural site in the Bavarian forest. The trees were well-supplied with nitrogen and exposed to either elevated CO 2 (ambient + 200 μLL -1 ), elevated ozone (80 nLL -1 , from June to October) or to a combination of both factors. Controls were grown with 20 nLL -1 O 3 and ambient CO 2 levels. In each chamber, a subset of trees was subjected to episodical drought stress in summer. Needles from controls investigated in October (summer conditions) and January (winter conditions) showed little seasonal variation of superoxide dismutase (SOD), an approximately 2-fold reduction in catalase (CAT), and a 2-fold increase in guaiacol peroxidase (POD) activity. Exposure to elevated CO 2 did not affect the activities of any of these enzymes in October and January, respectively, but caused a significant reduction in soluble protein. Ozone had no significant effect. Drought stress caused memory effects. In January, needles from trees drought-stressed in summer contained higher activities of defence enzymes and soluble protein contents than needles from well-watered trees. Three weeks after the end of a drought episode in summer, needles from spruce trees grown at elevated CO 2 contained increased CAT and POD activities as compared to needles from trees grown at ambient CO 2 . This response was increased, if elevated ozone was present as an additional stress factor. These observations suggest that Norway spruce trees grown under elevated atmospheric CO 2 concentrations might better be able to compensate environmental stresses than trees grown at ambient atmospheric CO 2 concentrations.

An zwei ausgewachsenen Fichten im Freiland wurden die Stammdicken mit Dendrometern und das Matrixpotential des Bodens mit Tensiometern kontinuierlich gemessen. In einer frostfreien Periode ohne Zuwachs korrelieren Dickenanderungen der Freilandfichten eng mit der Anderung des Matrixpotentials. Eine funfjahrige Fichte unter kontrollierten Bedingungen in Klimakammern andert ihren Radius ebenfalls entsprechend der Wasserversorgung im Boden. Selbst nach extremer Bodentrockenheit (Matrixpotential ± −1000 hPa) sind die Radiusanderungen reversibel. Es scheint daher moglich, mit Hilfe von Stammdickenmessungen das Matrixpotential auch unterhalb von − 1000 hPa abschatzen zu konnen. Effects of soil water potential on stem radius changes of Norway spruce (Picea abies (L.) Karst.) The stem radius of 2 mature Norway spruce trees and the soil water potential were continuously measured in situ with dendrometers and tensiometers, respectively. In a period without growth and frost events the stem radius and the soil water potentials correlated closely. A young spruce tree grown under controlled conditions changed its radius in response to the soil water supply likewise. The shrinkage of the stem was reversible, even if the soil was very dry (soil water potential ± − 1000 hPa). An estimate of the soil water potential seems to be possible even beyond − 1000 hPa by measurements of stem radius changes.

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.

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.

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 × O 3 ) or an experimentally elevated twice-ambient O 3 regime (2 × O 3 ). The elevated 2 × 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 × O 3 compared with the ambient O 3 regime (1 × 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 × O 3 than 1 × 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.

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.