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In recent years, important processes controlling ecosystem carbon dynamics have been connected to fine-root exudation of soluble carbon compounds. Root exudation patterns may change depending on plant interactions and plant susceptibility to and recovery from drought. Recent investigations suggest that root exudation tends to increase with stress events, but quantification of the amount of carbon released from roots across soil depths with differing water availability and species interactions are missing.We tested if root exudation rates were negatively correlated with soil water content across soil depths during and after drought. We further tested if species in mixture, often considered to be less stressed under drought, exuded less carbon than species in monospecific environments. Exudates were sampled in a mature Fagus sylvatica L. and Picea abies (L.) Karst. forest at the end of a five-year throughfall exclusion period and again one year after the drought ended. We quantified root exudates and their variation with soil depth for both tree species in monospecific and mixed species zones.Carbon exudation significantly increased in fine roots exposed to dry soils (

Forests are increasingly affected by heatwaves, often co-occurring with drought, with consequences for water and carbon (C) cycling. However, our ability to project tree resilience to more intense hot droughts remains limited. Here, we used single tree chambers (n = 18) to investigate transpiration (E), net assimilation (Anet), root respiration (Rroot) and stem diameter change in Scots pine seedlings in a control treatment and during gradually intensifying heat or drought-heat stress (max. 42 °C), including recovery. Alongside this, we assessed indicators of stress impacts and recovery capacities. In the heat treatment, excessive leaf heating was mitigated via increased E, while under drought-heat, E ceased and leaf temperatures reached 46 °C. However, leaf electrolyte leakage was negligible, while light-adapted quantum yield of photosystem II (F′v/F′m) declined alongside Anet moderately in heat, but strongly in drought-heat seedlings, in which respiration exceeded C uptake. Drought-heat largely affected the hydraulic system as apparent in stem diameter shrinkage, declining relative needle water content (RWCNeedle) and water potential (ΨNeedle) reaching −2.7 MPa, alongside a 90% decline of leaf hydraulic conductance (KLeaf). Heat alone resulted in low functional impairment and all measured parameters recovered quickly. Contrary, following drought-heat, the recovery of KLeaf was incomplete and stem hydraulic conductivity (KS) was 25% lower than the control. However, F′v/F′m recovered and the tree net C balance reached control values 2 days post-stress, with stem increment rates accelerating during the second recovery week. This indicates a new equilibrium of C uptake and release in drought-heat seedlings independent of hydraulic impairment, which may slowly contribute to the repair of damaged tissues. In summary, Scots pine recovered rapidly following moderate heat stress, while combined with drought, hydraulic and thermal stress intensified, resulting in functional damage and slow recovery of hydraulic conductance. This incomplete hydraulic recovery could critically limit evaporative cooling capacities and C uptake under repeated heatwaves.

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.

Heatwaves combined with drought affect tree functioning with as yet undetermined legacy effects on carbon (C) and nitrogen (N) allocation. - We continuously monitored shoot and root gas exchange, δ13CO2 of respiration and stem growth in well-watered and drought-treated Pinus sylvestris (Scots pine) seedlings exposed to increasing daytime temperatures (max. 42°C) and evaporative demand. Following stress release, we used 13CO2 canopy pulse-labeling, supplemented by soil-applied 15N, to determine allocation to plant compartments, respiration and soil microbial biomass (SMB) over 2.5 wk. - Previously heat-treated seedlings rapidly translocated 13C along the long-distance transport path, to root respiration (Rroot; 7.1 h) and SMB (3 d). Furthermore, 13C accumulated in branch cellulose, suggesting secondary growth enhancement. However, in recovering drought-heat seedlings, the mean residence time of 13C in needles increased, whereas C translocation to Rroot was delayed (13.8 h) and 13C incorporated into starch rather than cellulose. Concurrently, we observed stress-induced low N uptake and aboveground allocation. - C and N allocation during early recovery were affected by stress type and impact. Although C uptake increased quickly in both treatments, drought-heat in combination reduced the above–belowground coupling and starch accumulated in leaves at the expense of growth. Accordingly, C allocation during recovery depends on phloem translocation capacity., New Phytologist, 233 (2), ISSN:0028-646X, ISSN:1469-8137

After drought events, tree recovery depends on sufficient carbon (C) allocation to the sink organs. The present study aimed to elucidate dynamics of tree-level C sink activity and allocation of recent photoassimilates (C$_{new}$) and stored C in c. 70-year-old Norway spruce (Picea abies) trees during a 4-week period after drought release. We conducted a continuous, whole-tree $^{13}$C labeling in parallel with controlled watering after 5 years of experimental summer drought. The fate of C$_{new}$ to growth and CO$_{2}$ efflux was tracked along branches, stems, coarse- and fine roots, ectomycorrhizae and root exudates to soil CO$_{2}$ efflux after drought release. Compared with control trees, drought recovering trees showed an overall 6% lower C sink activity and 19% less allocation of C$_{new}$ to aboveground sinks, indicating a low priority for aboveground sinks during recovery. In contrast, fine-root growth in recovering trees was seven times greater than that of controls. However, only half of the C used for new fine-root growth was comprised of C$_{new}$ while the other half was supplied by stored C. For drought recovery of mature spruce trees, in addition to C$_{new}$, stored C appears to be critical for the regeneration of the fine-root system and the associated water uptake capacity.

Under ongoing global climate change, drought periods are predicted to increase in frequency and intensity in the future. Under these circumstances, it is crucial for tree's survival to recover their restricted functionalities quickly after drought release. To elucidate the recovery of carbon (C) transport rates in c. 70-year-old Norway spruce (Picea abies [L.] KARST.) after 5 years of recurrent summer droughts, we conducted a continuous whole-tree $^{13}$C labeling experiment in parallel with watering. We determined the arrival time of current photoassimilates in major C sinks by tracing the $^{13}$C label in stem and soil CO$_{2}$ efflux, and tips of living fine roots. In the first week after watering, aboveground C transport rates (CTR) from crown to trunk base were still 50% lower in previously drought-stressed trees (0.16 ± 0.01 m h$^{-1}$) compared to controls (0.30 ± 0.06 m h$^{-1}$). Conversely, CTR below ground, that is, from the trunk base to soil CO$_{2}$ efflux were already similar between treatments (c. 0.03 m h$^{-1}$). Two weeks after watering, aboveground C transport of previously drought-stressed trees recovered to the level of the controls. Furthermore, regrowth of water-absorbing fine roots upon watering was supported by faster incorporation of $^{13}$C label in previously drought-stressed (within 12 ± 10 h upon arrival at trunk base) compared to control trees (73 ± 10 h). Thus, the whole-tree C transport system from the crown to soil CO$_{2}$ efflux fully recovered within 2 weeks after drought release, and hence showed high resilience to recurrent summer droughts in mature Norway spruce forests. This high resilience of the C transport system is an important prerequisite for the recovery of other tree functionalities and productivity.

Carbon (C) exuded via roots is proposed to increase under drought and facilitate important ecosystem functions. However, it is unknown how exudate quantities relate to the total C budget of a drought-stressed tree, that is, how much of net-C assimilation is allocated to exudation at the tree level. - We calculated the proportion of daily C assimilation allocated to root exudation during early summer by collecting root exudates from mature Fagus sylvatica and Picea abies exposed to experimental drought, and combining above- and belowground C fluxes with leaf, stem and fine-root surface area. - Exudation from individual roots increased exponentially with decreasing soil moisture, with the highest increase at the wilting point. Despite c. 50% reduced C assimilation under drought, exudation from fine-root systems was maintained and trees exuded 1.0% (F. sylvatica) to 2.5% (P. abies) of net C into the rhizosphere, increasing the proportion of C allocation to exudates two- to three-fold. Water-limited P. abies released two-thirds of its exudate C into the surface soil, whereas in droughted F. sylvatica it was only one-third. - Across the entire root system, droughted trees maintained exudation similar to controls, suggesting drought-imposed belowground C investment, which could be beneficial for ecosystem resilience.

Plant responses to drought and heat stress have been extensively studied, whereas post-stress recovery, which is fundamental to understanding stress resilience, has received much less attention. Here, we present a conceptual stress-recovery framework with respect to hydraulic and metabolic functioning in woody plants. We further synthesize results from controlled experimental studies following heat or drought events and highlight underlying mechanisms that drive post-stress recovery. We find that the pace of recovery differs among physiological processes. Leaf water potential and abscisic acid concentration typically recover within few days upon rewetting, while leaf gas exchange-related variables lag behind. Under increased drought severity as indicated by a loss in xylem hydraulic conductance, the time for stomatal conductance recovery increases markedly. Following heat stress release, a similar delay in leaf gas exchange recovery has been observed, but the reasons are most likely a slow reversal of photosynthetic impairment and other temperature-related leaf damages, which typically manifest at temperatures above 40 °C. Based thereon, we suggest that recovery of gas exchange is fast following mild stress, while recovery is slow and reliant on the efficiency of repair and regrowth when stress results in functional impairment and damage to critical plant processes. We further propose that increasing stress severity, particular after critical stress levels have been reached, increases the carbon cost involved in reestablishing functionality. This concept can guide future experimental research and provides a base for modeling post-stress recovery of carbon and water relations in trees.

Forests are increasingly affected by heatwaves, often co-occurring with drought, with consequences for water and carbon (C) cycling. However, our ability to project the resilience of trees to an intensification of hot droughts remains limited. Here, we used single tree cuvettes (n=18) allowing us to investigate transpiration (E), net assimilation (Anet), root respiration (Rroot) and stem diameter change in Scots pine seedlings during gradually intensifying heat or drought-heat stress (max. 42°C), and post-stress. Further, we assessed indicators of stress impacts and recovery capacities.Under heat stress, well-watered seedlings prevented overheating of leaves effectively via increased E, while under drought-heat leaf temperatures increased to 46°C. However, leaf electrolyte leakage was negligible, but F’v/F’m declined alongside Anet moderately in heat but strongly in drought-heat seedlings, in which respiration exceeded C uptake. Further, the decrease of needle water potential (ψNeedle) to −2.7 MPa and relative needle water content (RWCNeedle) under drought-heat reflected a decline of leaf hydraulic conductance (KLeaf) by 90% and stem hydraulic conductivity (KS) by 25%. Alongside, we observed pronounced stem diameter shrinkage.Heat stress alone resulted in low functional impairment and all measured parameters recovered fast. In contrast, larger impacts following combined heat and drought led to the incomplete recovery of KLeaf and KS. Despite Anet tended to be reduced albeit F’v/F’m had recovered, the seedlings’ net C balance reached control values 2 d after stress release and stem growth rates exceeded control rates in the 2nd week post-stress. This indicates that a new equilibrium of C uptake and release was maintained at the tree level, slowly supporting regaining of stress-induced losses.In summary, we highlight that under moderate heatwaves with low functional impairment, recovery is fast in Scots pine, while in combination with drought hydraulic and thermal stress are intensified, resulting in functional damage and delayed recovery processes. The incomplete recovery of hydraulic conductance indicates limited water transport capacities that could become critical under repeated heat events.

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.

This contribution presents the result of a free-air 13C labeling experiment on mature Norway spruce (P. abies [L.] KARST.) upon watering after five years of recurrent summer drought in southern Germany, focusing on whole tree allocation processes. Mature spruce trees had been exposed to recurrent summer drought from 2014 to 2018 through complete exclusion of precipitation throughfall from spring to late fall (i.e., March to November). In early summer 2019, the drought stressed spruce trees were watered to investigate their recovery processes. In parallel with the watering, we conducted a whole-tree 13C labeling in canopies and traced the signal in various C sinks, i.e. stem phloem and CO2 efflux, tree rings at different heights, coarse roots, fine root tips, mycorrhiza, root exudates, and soil CO2 efflux.We hypothesize that drought stressed spruce preferentially allocates newly assimilated C to belowground sinks upon drought release. Conversely to our expectations, allocation to belowground C sinks was not stimulated in drought stressed compared to control spruce. Likewise, the relative amount of recently fixed C allocated to aboveground sinks did not differ between treatments. Our findings suggest that the belowground C sinks are not of higher priority for the allocation of newly assimilated C upon watering after long-term drought. The observed allocation pattern is discussed taking total above- and belowground biomass as well as C source/sink relations into account.

Rising atmospheric [CO

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.

The resistance of trees to stress events has been studied intensively, however we know little on underlying processes affecting the recovery of trees following stress release. Hence, this clearly impairs our ability to project the resilience of trees and forests to an intensification of heatwaves and drought spells.Here we studied the legacy effects of heat and heat-drought stress on carbon (C) allocation dynamics in Scots pine. We were particularly interested in how C allocation changes post heat and heat-drought stress and how this change in allocation affects tree growth. We exposed Pinus sylvestris seedlings to increasing temperatures from 30 to 40°C within 18 days either under well-watered or drought conditions and measured stem growth, leaf water potential and above- and belowground gas exchange. Two days after stress release, we conducted a 13CO2 pulse-labelling experiment in custom build single tree cuvettes (n=18) allowing us to continuously monitor 13CO2 shoot and root gas exchange. We then chased the fate of the newly assimilated C from leaves to roots via soluble sugars, starch and cellulose.Our results showed that Pinus sylvestris is able to recover gas exchange following heat release immediately in the well-watered trees, while drought-treated trees recovered slightly slower. We found indications for a stress compensatory response of the previously heat-treated trees, which tended to translocate recent assimilates faster compared to the control trees as identified in the dynamics of water-soluble carbon in the phloem and root 13CO2 efflux. In addition, we found larger stem growth rates in the heat-treated trees which was also reflected by a larger investment of new assimilates to cellulose. In the trees that experienced both, heat and drought stress, C allocation differed strongly from the control trees as apparent in a half as fast C translocation from leaves to root respiration and large investments of new assimilates into starch. This delayed translocation but enhanced allocation towards C storage in needle tissues was reflected in a delayed recovery of stem growth and very low detection of the 13C signal in twig, root and stem cellulose. We can conclude that heatwaves of 40°C have relatively moderate responses on C allocation post-stress, whereas hot drought stress clearly affects C allocation as indicated by a delayed C transport capacity and a preferential allocation towards C storage in needle tissues. This could indicate that C allocation following hot drought stress is affected by an impaired phloem functionality, which only slowly recovers post-stress.

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

Trees are increasingly exposed to hot droughts due to CO

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.

Predicting the net effects on the carbon and water balance of semi-arid forests under future conditions depends on ecosystem processes responding to changes in soil and atmospheric drought. Here we apply a combination of field observations and soil–plant–atmosphere modeling (SPA) to study carbon and water dynamics in a regenerating ponderosa pine forest. The effects of soil and atmospheric drought were quantified based on a field irrigation experiment combined with model simulations. To assess future effects of intensifying drought on ecosystem processes, the SPA model was run using temperature and precipitation scenarios for 2040 and 2080. Experimentally increased summer water availability clearly affected tree hydraulics and enhanced C uptake in both the observations and the model. Simulation results showed that irrigation was sufficient to eliminate soil water limitation and maintaining transpiration rates, but gross primary productivity (GPP) continued to decrease. Observations of stomatal conductance indicated a dominant role of vapor pressure deficit (VPD) in limiting C uptake. This was confirmed by running the simulation under reduced atmospheric drought (VPD of 1 kPa), which largely maintained GPP rates at pre-drought conditions. The importance of VPD as a dominant driver was underlined by simulations of extreme summer conditions. We found GPP to be affected more by summer temperatures and VPD as predicted for 2080 (−17%) than by reductions in summer precipitation (−9%). Because heterotrophic respiration responded less to heat (−1%) than to reductions in precipitation (−10%), net ecosystem C uptake declined strongest under hotter (−38%) compared to drier summer conditions (−8%). Considering warming trends across all seasons (September–May: +3 °C and June–August: +4.5 °C), the negative drought effects were largely compensated by an earlier initiation of favorable growing conditions and bud break, enhancing early season GPP and needle biomass. An adverse effect, triggered by changes in early season allocation patterns, was the decline of wood and root biomass. This imbalance may increase water stress over the long term to a threshold at which ponderosa pine may not survive, and highlights the need for an integrated process understanding of the combined effects of trends and extremes.

© 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.

ScaleX is a collaborative measurement campaign, collocated with a long-term environmental observatory of the German Terrestrial Environmental Observatories (TERENO) network in the mountainous terrain of the Bavarian Prealps, Germany. The aims of both TERENO and ScaleX include the measurement and modeling of land surface–atmosphere interactions of energy, water, and greenhouse gases. ScaleX is motivated by the recognition that long-term intensive observational research over years or decades must be based on well-proven, mostly automated measurement systems, concentrated in a small number of locations. In contrast, short-term intensive campaigns offer the opportunity to assess spatial distributions and gradients by concentrated instrument deployments, and by mobile sensors (ground and/or airborne) to obtain transects and three-dimensional patterns of atmospheric, surface, or soil variables and processes. Moreover, intensive campaigns are ideal proving grounds for innovative instruments, methods, and techniques to measure quantities that cannot (yet) be automated or deployed over long time periods. ScaleX is distinctive in its design, which combines the benefits of a long-term environmental-monitoring approach (TERENO) with the versatility and innovative power of a series of intensive campaigns, to bridge across a wide span of spatial and temporal scales. This contribution presents the concept and first data products of ScaleX-2015, which occurred in June–July 2015. The second installment of ScaleX took place in summer 2016 and periodic further ScaleX campaigns are planned throughout the lifetime of TERENO. This paper calls for collaboration in future ScaleX campaigns or to use our data in modelling studies. It is also an invitation to emulate the ScaleX concept at other long-term observatories.

Extreme weather conditions, like heat waves and drought, can substantially affect tree physiology and the emissions of biogenic volatile organic compounds (BVOC), including isoprene. To date, however, there is only limited understanding of BVOC emission patterns during prolonged heat and coupled heat–drought stress as well as post-stress recovery. To assess the impacts of heat and heat–drought stress on BVOC emissions, we studied gas exchange and isoprene emissions of black locust trees under controlled environmental conditions. Leaf gas exchange of isoprene, CO2 and H2O was quantified using branch chambers connected to a protontransfer-reaction mass spectrometer and an infrared gas analyzer. Heat and heat–drought stress resulted in a sharp decline of photosynthesis and stomatal conductance. Simultaneously, isoprene emissions increased six- to eight-fold in the heat and heat–drought treatment and resulted in a carbon loss that was equivalent to 12 % and 20 % of assimilated carbon at the time of measurement. Once temperature stress was released at the end of two 15 days long heat waves, stomatal conductance remained reduced, while isoprene emissions and photosynthesis recovered quickly to values of the control trees. Further, we found isoprene emissions to co-vary with net photosynthesis during non-stressful conditions, while during the heat waves, isoprene emissions could be solely described by non-linear functions of light and temperature. However, when isoprene emissions betweentreatments were compared under the same temperature and light conditions (e.g., T = 30° C, PAR = 500 µmol m−2 s−1), heat and heat–drought stressed trees would emit less isoprene than control trees. Ourfindings suggest that different parameterizations of light and temperature functions are needed in order to predict tree isoprene emissions under heat and combined heat–drought stress.

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.

a b s t r a c t Changes in the hydrological cycle, as predicted and currently observed, are expected to significantly impact the water and carbon balance of water-limited forest ecosystems. However, differences in the water-sensitivity of component processes make carbon balance predictions challenging. To exam- ine responses of ecosystem components to water limitations, we conducted a study of tree, soil and ecosystem-level processes in a young ponderosa pine stand under natural summer drought (control) and increased soil water conditions (watered). Weekly-averaged tree transpiration (Ttree), gross ecosystem photosynthesis (GPP) and soil CO2 efflux (Rstree; nearby trees) were related with soil water content (SWC; polynomial form: Ttree R2 = 0.98 and Rstree R2 = 0.91, logarithmic form: GPP R2 = 0.86) and declined rapidly when relative extractable soil water (REW) was

Although soil respiration, a major CO(2) flux in terrestrial ecosystems, is known to be highly variable with time, the response of its component fluxes to temperature and phenology is less clear. Therefore, we partitioned soil respiration (SR) into microbial (MR) and root-rhizosphere respiration (RR) using small root exclusion treatments in a mixed mountain forest in Switzerland. In addition, fine root respiration (FRR) was determined with measurements of excised roots. RR and FRR were strongly related to each other (R(2) = 0.92, n = 7), with RR contributing about 46% and FRR about 32% to total SR. RR rates increased more strongly with temperature (Q(10) = 3.2) than MR rates (Q(10) = 2.3). Since the contribution of RR to SR was found to be higher during growing (50%) than during dormant periods (40%), we separated the 2-year data set into phenophases. During the growing period of 2007, the temperature sensitivity of RR (Q(10) = 2.5, R(2) = 0.62) was similar to that of MR (Q(10) = 2.2, R(2) = 0.57). However, during the dormant period of 2006/2007, RR was not related to soil temperature (R(2) = 0.44, n.s.), in contrast to MR (Q(10) = 7.2; R(2) = 0.92). To better understand the influence of plant activity on root respiration, we related RR and FRR rates to photosynthetic active radiation (both R(2) = 0.67, n = 7, P = 0.025), suggesting increased root respiration rates during times with high photosynthesis. During foliage green-up in spring 2008, i.e., from bud break to full leaf expansion, RR increased by a factor of 5, while soil temperature increased only by about 5 degrees C, leading to an extraordinary high Q(10) of 10.6; meanwhile, the contribution of RR to SR increased from 29 to 47%. This clearly shows that root respiration and its apparent temperature sensitivity highly depend on plant phenology and thus on canopy assimilation and carbon allocation belowground.

Studies on soil respiration in mountain forests are rather scarce compared to their broad distribution. Therefore, we investigated daily, seasonal and annual soil respiration rates in a mixed forest (Lageren), located at about 700 m in the Swiss Jura mountains, during 2 years (2006 and 2007). Soil respiration (SR) was measured continuously with high temporal resolution (half-hourly) at one single point (SRautomated) and periodically with high spatial resolution (SRmanual) at 16 plots within the study site. Both, SRautomated and SRmanual showed a similar seasonal cycle. SR strongly depended on soil temperature in 2007 (R 2 = 0.82–0.92), but less so in 2006 (R 2 = 0.56–0.76) when SR was water limited during a summer drought. Including soil moisture improved the fit of the 2006 model significantly (R 2 = 0.78–0.97). Total annual SR for the study site was estimated as 869 g C m−2 year−1 for 2006 and as 907 g C m−2 year−1 for 2007 (uncertainty

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.