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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 past decade with the drought years of 2015, 2018 and 2019 in Central Europe revealed strikingly the devastating consequences of severe and repeated drought/heat events on forest ecosystems. Nevertheless, responses of the water balance of trees and forest stands to such repeated drought events and subsequent recovery are poorly understood. The estimation of the water consumption of trees and forest ecosystems is a crucial part of many process-based models, especially under severe drought events. One major factor for the calculation of the water consumption per tree by xylem sap flow measurements is the radial profile of the xylem sap flow density decreasing towards the inner part of the sapwood. However such profiles are very scarce, especially under repeated and severe drought events.Here, we present the changes of the profile of xylem sap flow density of European beech and Norway spruce within a five-year throughfall-exclusion (TE) experiment and a subsequent recovery. Two different methods were used to measure the xylem sap flow density up to 8cm sapwood depth, i.e. the heat dissipation and heat field deformation method. In beech, there was no difference in the linear radial profile between the TE and the CO (control) trees. However, under drought, for spruce the xylem sap flow density was strongly reduced along the profile by about 48 ± 16 % and the profile changed to an exponential decrease compared to the linear decrease for CO trees. Even two years upon drought release, the profile has not recovered. The reduction in the xylem sap flow density profile of drought stressed spruce was accompanied by heavy loss of fine roots and a reduction of the leaf area.The use of standardized xylem sap flow profiles without the consideration of drought induced changes would lead to an overestimation of the water consumption of more than 30%. These results stress the importance of the radial profile measurements for the calculation of water balance of trees and thus forest ecosystems.

Acclimation processes in a changing climate require the ability to tolerate and survive abiotic stress events (e.g. drought), challenging especially immobile and long-living species and ecosystems, such as forests. The drought years 2018/2019 in Central Europe have laid bare the vulnerability of temperate forest ecosystems to drought and heat. The recovery phase after the stress event represents a crucial phase, especially after intense and repeated drought periods, and might be different for anisohydric and isohydric species.The second phase of the Kranzberg Forest Roof (KROOF) experiment in southeast Germany focused on the watering of mature more anisohydric European beech (Fagus sylvatica L.) and more isohydric Norway spruce (Picea abies (L.) H. Karst.) after five years of repeated experimental summer drought. Both treatments, the former throughfall-exclusion (TE, recovering trees) and control (CO) plots were labeled with 2H enriched water by controlled watering, to end the experimental drought. Pre-dawn leaf water potential, stomatal conductance, xylem sap flow density (at breast height and crown base) and leaf osmoregulation were recorded for two growing seasons after drought release and the resilience and recovery times were calculated. All measured parameters were strongly reduced by on average 30% in both species due to the drought treatment. While the distribution of the labeled water upon irrigation across the soil profile occurred within a few days in both treatments, the water uptake and distribution within the trees was delayed by several days in recovering trees compared to control trees and in recovering spruce compared to recovering beech. Additionally, upon drought release recovering beech reached full resilience (i.e. same level as control trees) earlier than recovering spruce in water potential, stomatal conductance and xylem sap flow density and even showed signs of overcompensation by surpassing the control trees. No differences were found between the two species in the recovery of leaf osmoregulation.The “opposing” drought mitigation strategies seem to be responsible for the differences detected between more anisohydric beech and spruce during the recovery period. For example, the lack of recovery of xylem sap flow density at crown base in TE spruce indicates a re-filling of the stem water reservoirs upon watering. Additionally, we found fast responding parameters as water potential (hours to days) and stomatal conductance (days to weeks) compared to slow responding parameters such as osmoregulation (weeks to months) and full hydraulic recovery, i.e. xylem sap flow density, may even take years. Rapid physiological recovery after drought events, which are expected to increase in frequency and intensity with ongoing climate change, will be beneficial for overall recovery and might put faster-reacting trees in favor over slower responding species.

Due to climate change suitable forest management measures are required to mitigate the proposed exacerbating drought events as already observed in central Europe in summer 2003, 2015, 2018, and 2019. This contribution summarizes the findings of two long-term field experiments studying different forest management measures aiming at mitigating drought in forest stands.The first study investigated the potential of mixing tree composition for mitigating drought stress in Norway spruce (Picea abies [L.] Karst.). To this end, a five-year study with repeated summer droughts, experimentally induced via throughfall-exclusion (TE), was performed. The study objects were mature (60-80 years) old stands of Norway spruce in monoculture and mixture with European beech (Fagus sylvatica (L.). The impacts of repeated summer droughts were assessed on about 100 trees distributed on 12 plots and accessible via canopy crane (Kranzberg forest ROOF experiment in southern Germany). Predawn leaf water potentials of Norway spruce reached minima of -1.8 MPa, but were not affected by species mixture. Nevertheless, daily xylem sapflow density was increased up to 40% in mixture compared to pure strands. Likewise, stem growth, i.e. relative basal area increment, showed significantly higher drought resistance in mixture compared to monocultures.While altering forest stand composition seems to be promising on the long-term, the conversion from monocultures is often economically not suitable for young stands in the short- to medium-term. Therefore, reducing intraspecific competition via thinning is a frequently discussed option investigated in the second, eight-year-long study. In a 26-year old Norway spruce monoculture, three thinning intensities, i.e. unchanged (control), moderate thinning (MT with reduced basal area by 43%) and heavy thinning (HT, reduced basal area by 67%) were applied, potentially mitigating drought by reducing intraspecific competition and increasing soil water availability. Indeed, in both thinning intensities the duration of drought stress for the trees (soil water content below critical value) was reduced compared to controls for up to 5-7 years following the event. However, increased radiation and higher growth rates of the individual trees accompanied by the fast establishment of a vital ground vegetation diminished the difference in stand transpiration between MT and HT within two years. Moreover, belowground competition with the understorey vegetation suppressed fine root recovery under HT compared to MT in contradiction to increased leaf area on HT on tree-level and therefore increased transpirational demand.The presented studies suggest admixing of broadleaved beech into monocultures of Norway spruce to be a promising management measure in the long-term. In juvenile monocultures of Norway spruce frequent and intense thinning interventions while preventing the establishment of a vital understorey vegetation appears to be a promising forest measure, mitigating drought without losing sight of economic needs.

Every drought period will eventually end and plants will have access to water again. This phase of “re-watering” is a critical point that will either ensure survival or collapse of ecosystems. The drought years 2018 and 2019 have laid bare how vulnerable Central European forest-systems are, even under short-term water scarcity. To understand the effects of repeated summer drought and its release on mature forest stands we investigated the recovery of a mixed forest stand. After 5 years of repeated experimental summer drought on roughly 100 trees (with n = 6 plots) the second phase of the Kranzberg Forest Roof (k.roof) experiment was started, which focuses on the re-watering with Deuterium labeled water (2H2O) of the mature stand composed of European beech (Fagus sylvatica (L.)) and Norway spruce (Picea abies (L.)H.Karst.). According to our hypotheses the water household of the more anisohydric beech will recover faster and “stronger” (higher resilience) than the more isohydric spruce, due to the differences in stomatal control (hypothetical hydraulic regulation in beech vs. hormonal (ABA) control in spruce). We simulated a rainfall event to end our experimental drought and labeled the throughfall-exclusion (TE) and control (CO) plots of the k.roof experiment with roughly 13000 L for TE and 2000 L for CO of 2H2O enriched water, i.e. δ2H 1500 and 400 ‰ respectively. We traced the 2H2O signal along the soil-plant-atmosphere continuum (SPAC) from the soil through the stems and branches up to the leaves with conventional and real-time techniques (xylem sensors connected to CRDS system). Additionally, we measured leaf water potential and pressure-volume (PV) curves to assess the release of the drought stress. The distribution of the “new” water within the soil happened within a few days and we could not find any differences between the beech, mix or spruce dominated sites. However, the water uptake of the trees was significantly delayed in spruce compared to beech, evident from both the deuterium tracer signal (in stems and leaves) and leaf water potential. However, release of osmotic adjust was not different in the two species. The data allow for estimating the drought resilience of the water household of a mature forest stand after five-years of repeated summer drought and subsequent re-watering. While both species recovered their water household after several months to the same level as the control trees, we found beech to react faster and stronger than spruce.

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.

Vegetation in urban areas provides benefits to people, which are increasingly assessed and valued as ecosystem services (ESS). The regulation of the urban microclimate is one of these services, since trees have the potential to reduce urban heat loads by evapotranspiration and shading. Simultaneously, it has been suggested that trees from dry habitats should be used to cope with the increasing risks of drought under climate change in Central Europe. The underlying properties that enable those trees to reduce dependence on the environment, however, are assumed to come at the expense of biomass production and water loss. In the potentially conflicted area between drought tolerance and ESS we compared water consumption, water-use efficiency (WUE), crown structure and growth of six roadside species/cultivars, which are assumed to vary in drought tolerance, due to differing resource supplies in their native habitats. Acer platanoides, Carpinus betulus ‘Fastigiata’, and Tilia cordata ‘Greenspire’ were compared with Acer campestre, Ostrya carpinifolia, and Tilia tomentosa ‘Brabant’, the latter presumably being less water-demanding. Measurements took place in the municipal nursery of Munich during summer 2016. Surprisingly, the less water-demanding species/cultivars exhibited on average 1.24 times higher values of maximum daily sap flux density and up to seven times higher growth rates, both contributing to higher WUE. Scaled to leaf level, however, their mean daily transpiration rates were on average lower (0.21 and 0.31 kg H2O m−2 d−1, respectively). They also showed higher flexibility in response to changing weather with increased growth and transpiration under favorable conditions in early summer, but a more conservative water use in dry late summer. The results suggest that these species/cultivars tend to regulate their water use stronger under increasing dryness, whereas canopy size and leaf amount are still the main important determinants of species and cultivar differences in water use.

European beech (Fagus sylvatica L.) is a common tree species in Central European forests but the fitness of this species under future climate conditions is still a matter of debate. Here, we study the response of transplanted beech saplings in terms of their chemical and morphological traits to different levels of soil drought in a mountainous, partially open, storm damaged Norway spruce forest. Throughfall was excluded by roof constructions in three consecutive growing seasons. Spatial soil drought gradients resulted from soil water consumption by heterogeneously distributed mature spruce trees. Individual drought stress dose (DSD) was assessed for each beech sapling using cumulative soil matric potential in the rooting zone. After three years of drought treatment, we analyzed biomass components, traits of leaves and fine roots of sixty beech saplings. In the third year, the classical permanent wilting point in the soil was exceeded for some beech saplings, however, all saplings survived the drought treatments. Several chemical plant traits responded to DSD sum of the last two years, while morphological traits and biomass components were mostly not or weakly correlated with DSD. Potassium concentration and associated C:K and N:K ratios in fine roots were most sensitive to drought stress. We assume that K and other nutrients were hardly available in the extremely dry forest floor and that the undersupply could not be compensated by nutrient uptake from the mineral soil. Increasing C concentration in leaves indicates an acclimation towards sclerophylly to maintain the hydraulic function of leaves. Increasing specific root length (SRL), specific root tip density (SRTD) and decreasing fine root diameter also indicate a morphological acclimation of saplings to increasing DSD. Weak correlations between morphological traits and DSD partly resulted from non-linear responses to increasing DSD. Maximum number of mycorrhizal root tips and maximum fine root biomass occurred at medium DSD. Our results suggest that the acclimation of chemical and morphological traits fortifies the resilience of beech saplings against soil drought in the years after transplantation. However, reduced uptake of soil nutrients caused by drought can only be partially compensated by redistribution within the sapling. An expansion of the rooting zone in the moist, nutrient-poor subsoil can improve the water uptake of beech saplings during drought periods, but this strategy hardly improves their nutrient supply.

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.

By using a growth model, a simulation study was done to detect differing transpiration sums of an evergreen and a deciduous tree species within a mixed stand. The dependence of summer drought and transpiration on tree size and species, and the relationship of water use efficiency and tree growth was analyzed. The process-based growth model BALANCE was used to simulate the water balance and the growth of individual trees for the isohydric species Picea abies and the anisohydric species Fagus sylvatica within a mixed forest stand. The individual tree based model was able to realistically simulate the water balances at tree and stand level. Actual evapotranspiration and soil water content differed in species and was size-dependent. Spruce was more affected by drought than beech. Drought stress increased with tree size, an effect which was more pronounced for spruce than for beech. Wood productivity was positively correlated with water-use efficiency being more distinct in beech than spruce. Using individual tree based growth models effects of tree individuals in structured forest stands on water consumption, growth and productivity can be analyzed. The simulation results, i.e. the information of species-specific water consumption, growth rates and dependencies between water consumption and tree growth in stands of Norway spruce and European beech can help to mitigate effects of climate change on forest stand productivity and preserve an appropriate proportion of high quality timber mainly provided by spruce.

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.

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.

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.

A more detailed understanding of the micro-climatic thermal benefits of different urban tree species and the retrospective species characteristics is necessary to guide management decisions. In this review, we focused specifically on empirical data collected at ground level for below-canopy surface temperature (ST) and transpiration cooling (AT), using a meta-analysis method. Tree canopy density was clearly identified as the most influential driver of different mechanisms of cooling benefits. Secondly, climate of the cities where the trees were grown showed significant impacts on cooling potentials: trees grown in Oceanic and Continental climates showed a higher cooling potential compared to trees grown in Mediterranean climate for AT and sub-tropical climate for ST. Thirdly, tree growth in size and ground surface cover showed significant impact. ST decreases almost linearly with the increase of canopy density; however, the rate is significantly lower over transpiring grass surfaces. Transpiration of trees planted over grass was ten times higher (4.15 g m−2 min−1) compared to a tree planted in paved cut-out pits (0.44 g m−2 min−1). Moreover, diffuse porous wood anatomy and trees originating from temperate and resource-rich forests showed better cooling potentials. Among the leaf traits, dark green leaves

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.

Predicted intense and prolonged drought events challenge forest management. Thinning is debated as a silvicultural measure for reducing drought risk in densely established forest stands. We report on a thinning experiment in a 26-year-old Norway spruce stand (Picea abies), comprising of two thinning intensities and one unthinnned control. The removal of 43% (moderate thinning, MT) and 67% (heavy thinning, HT) of the initial basal area led to increased water availability during the entire three year observation period. Stand-level transpiration (Es) was decreased by about 25% upon moderate, and by about 50% upon heavy thinning during the first year after the interventions had been carried out. However, differences in Es across the treatments decreased within three years after thinning mainly due to increased single-tree transpiration and additional understory evapotranspiration at HT. Nevertheless, due to lower interception and transpiration on the thinned plots three years after treatment MT and HT still showed a substantial surplus in extractable soil water. The results showed that the main determinants concerning the extent of the mitigation effect with increasing thinning intensity were the available soil water storage capacity and the emerging understory vegetation. We conclude that repeated moderate thinning, through enhancing the water availability to the remaining trees, can mitigate drought risk in young spruce stands and thus, represent a viable silvicultural measure in anticipating possible water limitations due to climate change.

We review causes of synergies in mixed-species stands, develop guiding hypotheses for revealing their mechanisms and present a rainfall exclusion experiment along with a transect (KROOF) for exploring drought effects. While monocultures have dominated forest research and practice in the past, in face of growing resource scarcity and climate change, mixed-species stands are on the advance. Long-term observations show that mixed-species stands frequently over-yield monocultures, and they further suggest that the over-yielding is often higher on poor than on fertile sites and in low-growth than in high-growth years. However, the underlying causes have not yet been clarified. We start with a review of, among others, hydraulic redistribution, complementary eco-physiological traits, and ectomycorrhizal networks as possible causes behind the observed productivity gains in mixed-species stands. Then, we develop guiding hypotheses for further exploration of synergies in mixed-species stands. Finally, we introduce into the concept and model approach of the rainfall exclusion experiment for examining the role of water supply in mixed versus pure forest stands of spruce and beech. At the Kranzberg site, six plots are equipped with water retention roofs, which only close during rain events. The remaining six plots serve as non-roofed control. Together with the rainfall exclusion experiment, an ecological gradient with five sites extending through SE-Germany forms the “Kranzberg Roof Experiment” (in short KROOF). Kranzberg Forest is a part of this gradient from moist to dry conditions, with each site providing mixed and pure associations each of spruce and beech. The ecophysiological approach will be complemented by tree ring analysis and modelling of productivity of the tree associations under intense drought.

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.

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.

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 (O3) is a critical threat to forest ecosystems. A stomatal flux-based risk evaluation methodology at the leaf level was established recently in the context of the Convention on Long-Range Transboundary Air Pollution (LRTAP). This study demonstrates improvement and validation of the stomatal flux-effect approach for adult beech with results from the 8-year free-air O3 enrichment experiment at “Kranzberger Forst” (Germany). The risk assessment module of the SVAT model FO3REST, being under development for local scale O3-risk assessment of adult beech stands, was parameterized according to the LRTAP Convention’s Mapping Manual. Mean maximum stomatal conductance for water vapour of 245 mmol H2O m−2 PLA s−1, as suggested in the LRTAP Convention’s Mapping Manual for beech, was affirmed by assessment at “Kranzberger Forst”, resulting in 162 mmol O3 m−2 PLA s−1 upon recommended adjustment of the O3/water vapour diffusivity ratio to 0.663. Based on this ratio, a provisional corrected flux-effect function was deduced. Modelled Phytotoxic O3 Doses (POD 1) and potential O3-caused losses in biomass formation estimated with a site-specific stomatal conductance algorithm differed slightly only from estimates by the original LRTAP parameterisation. Analysis-derived POD 1 target value within the meaning of Article 2 of the European Council Directive 2008/50/EC of 10 mmol O3 m−2 corresponded to potential loss in biomass formation of about 10 % in ambient air relative to “pre-industrial” conditions. However, exceedance occurred by about a factor of two during the study period, indicating high risk at “Kranzberger Forst” under ambient air. Assessment for doubled O3 exposure indicated potential underestimation even of the O3 risk because modelled losses in biomass formation are in the lower range of the standard deviation of the observed ones.

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.

Compared to lowland trees and forests, evidence on water relations at mountainous and sub-alpine sites with their climatic and topographic peculiarities is scarce. On such limited grounds, the outcome of three pilot studies is reviewed, intended to launch working hypotheses for initiating integrative research across the altitudinal gradient of the European Alps. Highlighted are tree transpiration and structural differentiation as well as stand water balance of mono-specific and/or mixed forests with Picea abies, Pinus cembra, Larix decidua and Fagus sylvatica at sub-alpine (timberline ecotone), mountainous and colline elevation. Given the preliminary evidence, tree-allometric structures of relevancy for water transport appeared to be independent of elevation and forest type, although timberline trees under open-canopy conditions did not buffer transpiration by internal water storage and had enhanced foliage/sapwood area ratios. Transpiration appeared to depend on site conditions rather than site-dependent adaptation. Canopy transpiration approached 20% of the high seasonal precipitation at the mountainous site, with about 60% being ascribed to run-off, whereas the water budget of the colline site was balanced during summer. The water balance of the subalpine site resembled, at lower precipitation, that of the mountainous site. The derived hypotheses focus on mixed-stand transpiration under altitude-specific topography and moisture regimes, hydraulic adaptation and water demand versus uptake capacity, as this latter ratio is crucial at high altitude in view of expected warming. The clarification of consistencies relative to contrasts indicated by the pilot studies will be challenging across altitudes in view of tree species, forest types and topography.

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.

To prevent carbon (C) loss to consumers, trees need to defend their primary production. The tree-internal conflict in resource allocation between growth and defence demands has been the subject of various hypotheses but still requires quantification. A conceptual approach to approximating the C amount dispensable in favour of primary production at the expense of defence is demonstrated which is based on nine defence-related metabolite groups. Quantification is exemplified at the level of sun and shade leaves of adult Fagus sylvatica and Picea abies trees, two species contrasting in foliage type, under oxidative stress as induced by ozone exposure. The difference between maximum and minimum metabolite levels sampled several times throughout four consecutive growing seasons were conceived as dispensable between growth and defence-related metabolism and expressed in proportion of the mean annual gross primary production (GPP) of the foliage. In both species, this proportion amounted to between 2 and 5% of GPP (on a molar C basis). Remaining uncertainties are discussed as concerning functional overlap of substances between growth and defence-related metabolism, estimated classification of metabolite turnover rates and “third-party” trade-offs across C demands. Given the complexity of plant defence, simplification is needed for modelling allocation trade-offs in plants. The presented conceptual approach meets this need in approximating C transfer capacities between competing physiological demands and in stimulating empirical assessments towards mechanistic understanding.

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.

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

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.

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.