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Global warming and drying trends, as well as the increase in frequency and intensity of droughts, may have unprecedented impacts on various forest ecosystems. We assessed the role of internal water storage (WS) in drought resistance of mature pine trees in the semi-arid Yatir Forest. Transpiration (T), soil moisture and sap flow (SF) were measured continuously, accompanied by periodical measurements of leaf and branch water potential (Ψleaf) and water content (WC). The data were used to parameterize a tree hydraulics model to examine the impact of WS capacitance on the tree water relations. The results of the continuous measurements showed a 5-h time lag between T and SF in the dry season, which peaked in the early morning and early afternoon, respectively. A good fit between model results and observations was only obtained when the empirically estimated WS capacitance was included in the model. Without WS during the dry season, Ψleaf would drop below a threshold known to cause hydraulic failure and cessation of gas exchange in the studied tree species. Our results indicate that tree WS capacitance is a key drought resistance trait that could enhance tree survival in a drying climate, contributing up to 45% of the total daily transpiration during the dry season.

Rising atmospheric [CO2 ] (Ca ) generally enhances tree growth if nutrients are not limiting. However, reduced water availability and elevated evaporative demand may offset such fertilization. Trees with access to deep soil water may be able to mitigate such stresses and respond more positively to Ca . Here, we sought to evaluate how increased vapor pressure deficit and reduced precipitation are likely to modify the impact of elevated Ca (eCa ) on tree productivity in an Australian Eucalyptus saligna Sm. plantation with access to deep soil water. We parameterized a forest growth simulation model (GOTILWA+) using data from two field experiments on E. saligna: a 2-year whole-tree chamber experiment with factorial Ca (ambient =380, elevated =620 μmol mol-1 ) and watering treatments, and a 10-year stand-scale irrigation experiment. Model evaluation showed that GOTILWA+ can capture the responses of canopy C uptake to (1) rising vapor pressure deficit (D) under both Ca treatments; (2) alterations in tree water uptake from shallow and deep soil layers during soil dry-down; and (3) the impact of irrigation on tree growth. Simulations suggest that increasing Ca up to 700 μmol mol-1 alone would result in a 33% increase in annual gross primary production (GPP) and a 62% increase in biomass over 10 years. However, a combined 48% increase in D and a 20% reduction in precipitation would halve these values. Our simulations identify high D conditions as a key limiting factor for GPP. They also suggest that rising Ca will compensate for increasing aridity limitations in E. saligna trees with access to deep soil water under non-nutrient limiting conditions, thereby reducing the negative impacts of global warming upon this eucalypt species. Simulation models not accounting for water sources available to deep-rooting trees are likely to overestimate aridity impacts on forest productivity and C stocks.

Drought‐induced tree mortality is expected to occur more frequently under predicted climate change. However, the extent of a possibly mitigating effect of simultaneously rising atmospheric [CO$_{2}$] on stress thresholds leading to tree death is not fully understood, yet. Here, we studied the drought response, the time until critical stress thresholds were reached and mortality occurrence of Pinus halepensis (Miller). In order to observe a large potential benefit from eCO$_{2}$, the seedlings were grown with ample of water and nutrient supply under either highly elevated [CO$_{2}$] (eCO$_{2}$, c. 936 ppm) or ambient (aCO$_{2}$, c. 407 ppm) during 2 years. The subsequent exposure to a fast or a slow lethal drought was monitored using whole‐tree gas exchange chambers, measured leaf water potential and non‐structural carbohydrates. Using logistic regressions to derive probabilities for physiological parameters to reach critical drought stress thresholds, indicated a longer period for halving needle starch storage under eCO$_{2}$ than aCO$_{2}$. Stomatal closure, turgor loss, the duration until the daily tree C balance turned negative, leaf water potential at thresholds and time‐of‐death were unaffected by eCO$_{2}$. Overall, our study provides for the first‐time insights into the chronological interplay of physiological drought thresholds under long‐term acclimation to elevated [CO$_{2}$].

Drought alters carbon (C) allocation within trees, thereby impairing tree growth. Recovery of root and leaf functioning and prioritized C supply to sink tissues after drought may compensate for drought -induced reduction of assimilation and growth. It remains unclear if C allocation to sink tissues during and following drought is controlled by altered sink metabolic activities or by the availability of new assimilates. Understanding such mechanisms is required to predict forests' resilience to a changing climate. We investigated the impact of drought and drought release on C allocation in a 100-y-old Scots pine forest. We applied (CO2)-C-13 pulse labeling to naturally dry control and long-term irrigated trees and tracked the fate of the label in aboveand belowground C pools and fluxes. Allocation of new assimilates belowground was ca. 53% lower under nonirrigated conditions. A short rainfall event, which led to a temporary increase in the soil water content (SWC) in the topsoil, strongly increased the amounts of C transported belowground in the nonirrigated plots to values comparable to those in the irrigated plots. This switch in allocation patterns was congruent with a tipping point at around 15% SWC in the response of the respiratory activity of soil microbes. These results indicate that the metabolic sink activity in the rhizosphere and its modulation by soil moisture can drive C allocation within adult trees and ecosystems. Even a subtle increase in soil moisture can lead to a rapid recovery of belowground functions that in turn affects the direction of C transport in trees.

Trees are increasingly exposed to hot droughts due to CO2-induced climate change. However, the direct role of [CO2] in altering tree physiological responses to drought and heat stress remains ambiguous. - Pinus halepensis (Aleppo pine) trees were grown from seed under ambient (421 ppm) or elevated (867 ppm) [CO2]. The 1.5-yr-old trees, either well watered or drought treated for 1 month, were transferred to separate gas-exchange chambers and the temperature gradually increased from 25°C to 40°C over a 10 d period. Continuous whole-tree shoot and root gasexchange measurements were supplemented by primary metabolite analysis. - Elevated [CO2] reduced tree water loss, reflected in lower stomatal conductance, resulting in a higher water-use efficiency throughout amplifying heat stress. Net carbon uptake declined strongly, driven by increases in respiration peaking earlier in the well-watered (31– 32°C) than drought (33–34°C) treatments unaffected by growth [CO2]. Further, drought altered the primary metabolome, whereas the metabolic response to [CO2] was subtle and mainly reflected in enhanced root protein stability. - The impact of elevated [CO2] on tree stress responses was modest and largely vanished with progressing heat and drought. We therefore conclude that increases in atmospheric [CO2] cannot counterbalance the impacts of hot drought extremes in Aleppo pine.

Climate change will have a considerable impact on forest productivity worldwide. Forecasting the magnitude of such impact, with multiple environmental stressors changing simultaneously, is only possible with the help of process-based models. In order to assess their performance, such models require careful evaluation against measurements. However, direct comparison of model outputs against observational data is often not reliable, as models may provide the right answers due to the wrong reasons. This would severely hinder forecasting abilities under unprecedented climate conditions. Here, we present a methodology for model assessment, which supplements the traditional output-to-observation model validation. It evaluates model performance through its ability to reproduce observed seasonal changes of the most limiting environmental driver (MLED) for a given process, here daily gross primary productivity (GPP). We analyzed seasonal changes of the MLED for GPP in two contrasting pine forests, the Mediterranean Pinus halepensis Mill. Yatir (Israel) and the boreal Pinus sylvestris L. Hyytiälä (Finland) from three years of eddy-covariance flux data. Then, we simulated the same period with a state-of-the-art process-based simulation model (LandscapeDNDC). Finally, we assessed if the model was able to reproduce both GPP observations and MLED seasonality. We found that the model reproduced the seasonality of GPP in both stands, but it was slightly overestimated without site-specific fine-tuning. Interestingly, although LandscapeDNDC properly captured the main MLED in Hyytiälä (temperature) and in Yatir (soil water availability), it failed to reproduce high-temperature and high-vapor pressure limitations of GPP in Yatir during spring and summer. We deduced that the most likely reason for this divergence is an incomplete description of stomatal behavior. In summary, this study validates the MLED approach as a model evaluation tool, and opens up new possibilities for model improvement.

Globally, soils store two to three times as much carbon as currently resides in the atmosphere, and it is critical to understand how soil greenhouse gas (GHG) emissions and uptake will respond to ongoing climate change. In particular, the soil‐to‐atmosphere CO2 flux, commonly though imprecisely termed soil respiration (R S), is one of the largest carbon fluxes in the Earth system. An increasing number of high‐frequency R S measurements (typically, from an automated system with hourly sampling) have been made over the last two decades; an increasing number of methane measurements are being made with such systems as well. Such high frequency data are an invaluable resource for understanding GHG fluxes, but lack a central database or repository. Here we describe the lightweight, open‐source COSORE (COntinuous SOil REspiration) database and software, that focuses on automated, continuous and long‐term GHG flux datasets, and is intended to serve as a community resource for earth sciences, climate change syntheses and model evaluation. Contributed datasets are mapped to a single, consistent standard, with metadata on contributors, geographic location, measurement conditions and ancillary data. The design emphasizes the importance of reproducibility, scientific transparency and open access to data. While being oriented towards continuously measured R S, the database design accommodates other soil‐atmosphere measurements (e.g. ecosystem respiration, chamber‐measured net ecosystem exchange, methane fluxes) as well as experimental treatments (heterotrophic only, etc.). We give brief examples of the types of analyses possible using this new community resource and describe its accompanying R software package., Here we describe the lightweight, open source COSORE (COntinuous SOil REspiration) database and software. COSORE focuses on automated, continuous and long‐term greenhouse gas flux datasets, and is intended to serve as a community resource for earth sciences, climate change syntheses and model evaluation.

Biogenic volatile organic compounds (BVOC) play important roles in plant stress responses and can serve as stress indicators. While the impacts of gradual environmental changes on BVOCs have been studied extensively, insights in emission responses to repeated stress and recovery are widely absent. Therefore, we studied the dynamics of shoot gas exchange and BVOC emissions in Pinus halepensis seedlings during an induced moderate drought, two four-day-long heatwaves, and the combination of drought and heatwaves. We found clear stress-specific responses of BVOC emissions. Reductions in acetone emissions with declining soil water content and transpiration stood out as a clear drought indicator. All other measured BVOC emissions responded exponentially to rising temperatures during heat stress (maximum of 43 °C), but monoterpenes and methyl salicylate showed a reduced temperature sensitivity during the second heatwave. We found that these decreases in monoterpene emissions between heatwaves were not reflected by similar declines in their internal storage pools. Because stress intensity was extremely severe, most of the seedlings in the heat-drought treatment died at the end of the second heatwave (dark respiration ceased). Interestingly, BVOC emissions (methanol, monoterpenes, methyl salicylate, and acetaldehyde) differed between dying and surviving seedlings, already well before indications of a reduced vitality became visible in gas exchange dynamics. In summary, we could clearly show that the dynamics of BVOC emissions are sensitive to stress type, stress frequency, and stress severity. Moreover, we found indications that stress-induced seedling mortality was preceded by altered methanol, monoterpene, and acetaldehyde emission dynamics. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-04905-y.

Hydraulic redistribution (HR) of water from wet- to dry-soil zones is suggested as an important process in the resilience of forest ecosystems to drought stress in semiarid and tropical climates. Scenarios of future climate change predict an increase of severe drought conditions in temperate climate regions. This implies the need for adaptations of locally managed forest systems, such as European beech (Fagus sylvatica L.) monocultures, for instance, through the admixing of deep-rooting silver fir (Abies alba Mill.). We designed a stable-isotope-based split-root experiment under controlled conditions to test whether silver fir seedlings could perform HR and therefore reduce drought stress in neighboring beech seedlings. Our results showed that HR by silver fir does occur, but with a delayed onset of three weeks after isotopic labelling with 2H2O (&delta, 2H &asymp, +6000&permil, ), and at low rates. On average, 0.2% of added &sup2, H excess could be recovered via HR. Fir roots released water under dry-soil conditions that caused some European beech seedlings to permanently wilt. On the basis of these results, we concluded that HR by silver fir does occur, but the potential for mitigating drought stress in beech is limited. Admixing silver fir into beech stands as a climate change adaptation strategy needs to be assessed in field studies with sufficient monitoring time.

In 2018, Central Europe experienced one of the most severe and long-lasting summer drought and heat wave ever recorded. Before 2018, the 2003 millennial drought was often invoked as the example of a “hotter drought”, and was classified as the most severe event in Europe for the last 500 years. First insights now confirm that the 2018 drought event was climatically more extreme and had a greater impact on forest ecosystems of Austria, Germany and Switzerland than the 2003 drought. Across this region, mean growing season air temperature from April to October was more than 3.3°C above the long-term average, and 1.2 °C warmer than in 2003. Here, we present a first impact assessment of the severe 2018 summer drought and heatwave on Central European forests. In response to the 2018 event, most ecologically and economically important tree species in temperate forests of Austria, Germany and Switzerland showed severe signs of drought stress. These symptoms included exceptionally low foliar water potentials crossing the threshold for xylem hydraulic failure in many species and observations of widespread leaf discoloration and premature leaf shedding. As a result of the extreme drought stress, the 2018 event caused unprecedented drought-induced tree mortality in many species throughout the region. Moreover, unexpectedly strong drought-legacy effects were detected in 2019. This implies that the physiological recovery of trees was impaired after the 2018 drought event, leaving them highly vulnerable to secondary drought impacts such as insect or fungal pathogen attacks. As a consequence, mortality of trees triggered by the 2018 events is likely to continue for several years. Our assessment indicates that many common temperate European forest tree species are more vulnerable to extreme summer drought and heat waves than previously thought. As drought and heat events are likely to occur more frequently with the progression of climate change, temperate European forests might approach the point for a substantial ecological and economic transition. Our assessment also highlights the urgent need for a pan-European ground-based monitoring network suited to track individual tree mortality, supported by remote sensing products with high spatial and temporal resolution to track, analyse and forecast these transitions. ISSN:1439-1791 ISSN:1618-0089

Climate-induced tree mortality became a global phenomenon during the last century and it is expected to increase in many regions in the future along with a further increase in the frequency of drought and heat events. However, tree mortality at the ecosystem level remains challenging to quantify since long-term, tree-individual, reliable observations are scarce. Here, we present a unique data set of monitoring records from 276 permanent plots located in 95 forest stands across Switzerland, which include five major European tree species (Norway spruce, Scots pine, silver fir, European beech, and sessile and common oak) and cover a time span of over one century (1898–2013), with inventory periods of 5–10 years. The long-term average annual mortality rate of the investigated forest stands was 1.5%. In general, species-specific annual mortality rates did not consistently increase over the last decades, except for Scots pine forests at lower altitudes, which exhibited a clear increase of mortality since the 1960s. Temporal trends of tree mortality varied also depending on diameter at breast height (DBH), with large trees generally experiencing an increase in mortality, while mortality of small trees tended to decrease. Normalized mortality rates were remarkably similar between species and a modest, but a consistent and steady increasing trend was apparent throughout the study period. Mixed effects models revealed that gradually changing stand parameters (stand basal area and stand age) had the strongest impact on mortality rates, modulated by climate, which had increasing importance during the last decades. Hereby, recent climatic changes had highly variable effects on tree mortality rates, depending on the species in combination with abiotic and biotic stand and site conditions. This suggests that forest species composition and species ranges may change under future climate conditions. Our data set highlights the complexity of forest dynamical processes such as long-term, gradual changes of forest structure, demography and species composition, which together with climate determine mortality rates., Frontiers in Plant Science, 10, ISSN:1664-462X

Forest leaf area has enormous leverage on the carbon cycle because it mediates both forest productivity and resilience to climate extremes. Despite widespread evidence that trees are capable of adjusting to changes in environment across both space and time through modifying carbon allocation to leaves, many vegetation models use fixed carbon allocation schemes independent of environment, which introduces large uncertainties into predictions of future forest responses to atmospheric CO2 fertilization and anthropogenic climate change. Here, we develop an optimization-based model, whereby tree carbon allocation to leaves is an emergent property of environment and plant hydraulic traits. Using a combination of meta-analysis, observational datasets, and model predictions, we find strong evidence that optimal hydraulic-carbon coupling explains observed patterns in leaf allocation across large environmental and CO2 concentration gradients. Furthermore, testing the sensitivity of leaf allocation strategy to a diversity in hydraulic and economic spectrum physiological traits, we show that plant hydraulic traits in particular have an enormous impact on the global change response of forest leaf area. Our results provide a rigorous theoretical underpinning for improving carbon cycle predictions through advancing model predictions of leaf area, and underscore that tree-level carbon allocation to leaves should be derived from first principles using mechanistic plant hydraulic processes in the next generation of vegetation models.

© 2018, The Ecological Society of Japan. Understanding which species are able to recover from drought, under what conditions, and the mechanistic processes involved, will facilitate predictions of plant mortality in response to global change. In response to drought, some species die because of embolism-induced hydraulic failure, whilst others are able to avoid mortality and recover, following rehydration. Several tree species have evolved strategies to avoid embolism, whereas others tolerate high embolism rates but can recover their hydraulic functioning upon drought relief. Here, we focus on structures and processes that might allow some plants to recover from drought stress via embolism reversal. We provide insights into how embolism repair may have evolved, anatomical and physiological features that facilitate this process, and describe possible trade-offs and related costs. Recent controversies on methods used for estimating embolism formation/repair are also discussed, providing some methodological suggestions. Although controversial, embolism repair processes are apparently based on the activity of phloem and ray/axial parenchyma. The mechanism is energetically demanding, and the costs to plants include metabolism and transport of soluble sugars, water and inorganic ions. We propose that embolism repair should be considered as a possible component of a ‘hydraulic efficiency-safety’ spectrum. We also advance a framework for vegetation models, describing how vulnerability curves may change in hydrodynamic model formulations for plants that recover from embolism.

International audience; nitori Accumulating evidence highlights increased mortality risks for trees during severe drought, particularly under warmer temperatures and increasing vapour pressure deficit (VPD). Resulting forest die-off events have severe consequences for ecosystem services, biophysical and biogeochemical land-atmosphere processes. Despite advances in monitoring, modelling and experimental studies of the causes and consequences of tree death from individual tree to ecosystem and global scale, a general mechanistic understanding and realistic predictions of drought mortality under future climate conditions are still lacking. We update a global tree mortality map and present a roadmap to a more holistic understanding of forest mortality across scales. We highlight priority research frontiers that promote: (1) new avenues for research on key tree ecophysiological responses to drought; (2) scaling from the tree/plot level to the ecosystem and region; (3) improvements of mortality risk predictions based on both empirical and mechanistic insights; and (4) a global mong network of forest mortality. In light of recent and anticipated large forest die-off events such a research agenda is timely and needed to achieve scientific understanding for realistic predictions of drought-induced tree mortality. The implementation of a sustainable network will require support by stakeholders and political authorities at the international level.

Stem CO(2) concentrations (stem [CO(2)]) undergo large temporal variations that need to be understood to better link tree physiological processes to biosphere-atmosphere CO(2) exchange. During 19 months, stem [CO(2)] was continuously measured in mature subalpine Norway spruce trees (Picea abies) and jointly analysed with stem, soil and air temperatures, sap flow rates, stem radius changes and CO(2) efflux rates from stem and soil on different time scales. Stem [CO(2)] exhibited a strong seasonality, of which over 80% could be explained with stem and soil temperatures. Both physical equilibrium processes of CO(2) between water and air according to Henry's law as well as physiological effects, including sap flow and local respiration, concurrently contributed to these temporal variations. Moreover, the explanatory power of potential biological drivers (stem radius changes, sap flow and soil respiration) varied strongly with season and temporal resolution. We conclude that seasonal and daily courses of stem [CO(2)] in spruce trees are a combined effect of physical equilibrium and tree physiological processes. Furthermore, we emphasize the relevance of axial diffusion of CO(2) along air-filled spaces in the wood, and potential wound response processes owing to sensor installation.

The frequency and intensity of climatic extremes, such as heat waves, are predicted to increase globally, with severe implications for terrestrial carbon and water cycling. Temperatures may rise above critical thresholds that allow trees to function optimally, with unknown long-term consequences for forest ecosystems. In this context, we investigated how photosynthetic traits and the water balance in Douglas-fir are affected by exposure to three heat waves with temperatures about 12 degrees C above ambient. Photosynthetic carboxylation efficiency (V-cmax) was mostly unaffected, but electron transport (J(max)) and photosynthetic rates under saturating light (A(sat)) were strongly influenced by the heat waves, with lagging limitations on photosynthesis still being observed six weeks after the last heat wave. We also observed lingering heat-induced inhibitions on transpiration, minimum stomatal conductance, and night-time stomatal conductance (g(s-night)). Results from the stomatal models used to calculate minimum stomatal conductance were similar to g(s-night) and indicated changes in leaf morphology, e.g. stomatal occlusions and alterations in epicuticular wax. Our results show Douglas-fir's ability to restrict water loss following heat stress, but at the price of reduced photosynthetic performance. Such limitations indicate potential long-term restrictions that heat waves can impose on tree development and functioning under extreme climatic conditions. (C) 2016 Elsevier GmbH. All rights reserved.

From the 21st to the 23rd June 2017, the Herrenhausen castle in Hannover/Germany hosted a diverse and large crowd with more than 70 tree physiologists, forest ecologists, forest inventory experts, remote-sensing scientists, and vegetation modelers. Participants from six continents and from more than 20 countries gathered to discuss how to improve the scientific determination of global-scale patterns, drivers, and trends of a threatening phenomenon: the apparent emergence of recent widespread tree mortality events in diverse forests around the world. Continuing the theme of a workshop held at the Max-Planck Institute for Biogeochemistry in Jena (Germany) in 2014 (Hartmann et al., 2015), the Hanover meeting intended to develop approaches, tools and collaborative actions to accelerate progress in addressing regional patterns and trends of tree mortality (Williams et al., 2013). Over the last decade climate change related tree mortality events have been increasingly reported around the globe (van Mantgem et al., 2009; Carnicer et al., 2011; Peng et al., 2011; Brienen et al., 2015), but to what degree this is a global trend, amplifying under increasing climate change, remains uncertain.