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Xylem embolism resistance has been identified as a key trait with a causal relation to drought-induced tree mortality, but not much is known about its intra-specific trait variability (ITV) in dependence on environmental variation. We measured xylem safety and efficiency in 300 European beech (Fagus sylvatica L.) trees across 30 sites in Central Europe, covering a precipitation reduction from 886 to 522 mm year−1. A broad range of variables that might affect embolism resistance in mature trees, including climatic and soil water availability, competition, and branch age, were examined. The average P50 value varied by up to 1 MPa between sites. Neither climatic aridity nor structural variables had a significant influence on P50. However, P50 was less negative for trees with a higher soil water storage capacity, and positively related to branch age, while specific conductivity (Ks) was not significantly associated with either of these variables. The greatest part of the ITV for xylem safety and efficiency was attributed to random variability within populations. We conclude that the influence of site water availability on P50 and Ks is low in European beech, and that the high degree of within-population variability for P50, partly due to variation in branch age, hampers the identification of a clear environmental signal.

Leaf morphological and physiological traits control the carbon and water relations of mature trees and are determinants of drought tolerance, but it is not well understood how they are modified in response to water deficits. We analysed five sun-canopy leaf traits (mean leaf size (LS), specific leaf area (SLA), Huber value (HV), water potential at turgor loss point (Ψtlp ) and foliar carbon isotope signature (δ13 C)) in European beech (Fagus sylvatica L.) across three precipitation gradients sampled in moist (2010), dry (2019) and very dry (2018) summers, and tested their response to short-term water deficits (climatic water balance (CWB) preceding sample collection) and long-term water availability (mean annual precipitation (MAP), plant-available soil water capacity (AWC) and neighbourhood competition). Across the 34 sites, LS varied seven-fold (3.9-27.0 cm2 ), SLA four-fold (77.1-306.9 cm²·g-1 ) and HV six-fold (1.0-6.65 cm2 ·m-2 ). In the 2018 dataset, LS showed a negative and HV a positive relationship to MAP, which contradicts relations found in multi-species samples. Average Ψtlp ranged from -1.90 to -2.62 MPa and decreased across the sites with decreasing CWB in the month prior to measurement, as well as with decreasing MAP and AWC in 2019. Studied leaf traits varied considerably between years, suggesting that mast fruiting and the severe 2018 drought caused the formation of smaller leaves. We conclude that sun-canopy leaf traits of European beech exhibit considerable plasticity in response to climatic and edaphic aridity, and that osmotic adjustment may be an important element in the drought response strategy of this anisohydric tree species.

Key message By combining dendrochronological time-series analysis with radial vessel features, we show that the reconstruction of hydraulic properties improves our understanding of tree species’ acclimation potential to climate change. Abstract The vascular architecture plays a crucial role in the productivity and drought tolerance of broadleaf trees, but it is not yet fully understood how the hydraulic system is acclimating to a warmer and drier climate. Because vessel features may record temporal and spatial variability in climatic signals of the past better than tree-ring width, we combined dendrochronological time-series analysis with the calculation of stem hydraulic properties derived from radial vessel features. We aimed to reconstruct the development and sensitivity of the hydraulic system over six decades and to identify climatic control of xylem anatomy for five co-existing broad-leaved diffuse- and ring-porous tree species (genera Acer, Fagus, Fraxinus and Quercus) across three sites covering a precipitation gradient from 548 to 793 mm. We observed a significant influence of the climatic water balance (CWB) on the vessel features of all species, but the time lag, magnitude and direction of the response was highly species-specific. All diffuse-porous species suffered a decline in vessel diameter in dry years, and increase in vessel density in dry years and the year following. However, F. sylvatica was the only species with a significant long-term change in anatomical traits and a significant reduction in potential hydraulic conductivity (Kp) after dry winters and in dry summers, accompanied with the largest long-term decline in tree-ring width and the largest growth reduction in and after years with a more negative CWB. In contrast, the comparison across the precipitation gradient did not reveal any significant vessel-climate relationships. Our results revealed considerable plasticity in the hydraulic system especially of F. sylvatica, but also evidence of the drought-sensitivity of this species in accordance with earlier dendroecological and physiological studies. We conclude that the long-term reconstruction of hydraulic properties can add substantially to the understanding of the acclimation potential of different tree species to climate change.

Farmland tree cultivation is considered an important option for enhancing wood production. In South India, the native leaf-deciduous tree species Melia dubia is popular for short-rotation plantations. Across a rainfall gradient from 420 to 2170 mm year–1, we studied 186 farmland woodlots between one and nine years in age. The objectives were to identify the main factors controlling aboveground biomass (AGB) and growth rates. A power-law growth model predicts an average stand-level AGB of 93.8 Mg ha–1 for nine-year-old woodlots. The resulting average annual AGB increment over the length of the rotation cycle is 10.4 Mg ha–1 year–1, which falls within the range reported for other tropical tree plantations. When expressing the parameters of the growth model as functions of management, climate and soil variables, it explains 65% of the variance in AGB. The results indicate that water availability is the main driver of the growth of M. dubia. Compared to the effects of water availability, the effects of soil nutrients are 26% to 60% smaller. We conclude that because of its high biomass accumulation rates in farm forestry, M. dubia is a promising candidate for short-rotation plantations in South India and beyond.

Unprecedented tree dieback across Central Europe caused by recent global change-type drought events highlights the need for a better mechanistic understanding of drought-induced tree mortality. Although numerous physiological risk factors have been identified, the importance of two principal mechanisms, hydraulic failure and carbon starvation, is still debated. It further remains largely unresolved how the local neighborhood composition affects individual mortality risk. We studied 9435 young trees of 12 temperate species planted in a diversity experiment in 2013 to assess how hydraulic traits, carbon dynamics, pest infestation, tree height and neighborhood competition influence individual mortality risk. Following the most extreme global change-type drought since record in 2018, one third of these trees died. Across species, hydraulic safety margins (HSMs) were negatively and a shift towards a higher sugar fraction in the non-structural carbohydrate (NSC) pool positively associated with mortality risk. Moreover, trees infested by bark beetles had a higher mortality risk, and taller trees a lower mortality risk. Most neighborhood interactions were beneficial, although neighborhood effects were highly species-specific. Species that suffered more from drought, especially Larix spp. and Betula spp., tended to increase the survival probability of their neighbors and vice versa. While severe tissue dehydration marks the final stage of drought-induced tree mortality, we show that hydraulic failure is interrelated with a series of other, mutually inclusive processes. These include shifts in NSC pools driven by osmotic adjustment and/or starch depletion as well as pest infestation and are modulated by the size and species identity of a tree and its neighbors. A more holistic view that accounts for multiple causes of drought-induced tree mortality is required to improve predictions of trends in global forest dynamics and to identify mutually beneficial species combinations.

Understanding the vulnerability of trees to drought-induced mortality is key to predicting the fate of forests in a future climate with more frequent and intense droughts, although the underlying mechanisms are difficult to study in adult trees. Here, we explored the dynamic changes of water relations and limits of hydraulic function in dying adults of Norway spruce (Picea abies L.) during the progression of the record-breaking 2018 Central European drought. In trees on the trajectory to drought-induced mortality, we observed rapid, nonlinear declines of xylem pressure that commenced at the early onset of xylem cavitation and caused a complete loss of xylem hydraulic conductance within a very short time. We also observed severe depletions of nonstructural carbohydrates, though carbon starvation could be ruled out as the cause of the observed tree death, as both dying and surviving trees showed these metabolic limitations. Our observations provide striking field-based evidence for fast dehydration and hydraulic collapse as the cause of drought-induced mortality in adult Norway spruce. The nonlinear decline of tree water relations suggests that considering the temporal dynamics of dehydration is critical for predicting tree death. The collapse of the hydraulic system within a short time demonstrates that trees can rapidly be pushed out of the zone of hydraulic safety during the progression of a severe drought. In summary, our findings point toward a higher mortality risk for Norway spruce than previously assumed, which is in line with current reports of unprecedented levels of drought-induced mortality in this major European tree species.

SummaryThe increasing frequency of global change-type droughts has created a need for fast, accurate and widely applicable techniques for estimating xylem embolism resistance to improve forecasts of future forest changes.We used data from 12 diffuse-porous temperate tree species covering a wide range of xylem safety to compare the pneumatic and flow-centrifuge method for constructing xylem vulnerability curves. We evaluated the agreement between parameters estimated with both methods and the sensitivity of pneumatic measurements to the measurement duration.The agreement between xylem water potentials at 50% air discharged (PAD) estimated with the Pneumatron and 50% loss of hydraulic conductivity (PLC) estimated with the flow-centrifuge method was high (mean signed deviation: 0.12 MPa, Pearson correlation: 0.96 after 15 sec of gas extraction). However, the relation between the estimated slopes was more variable, resulting in lower agreement in xylem water potential at 12% and 88% PAD/PLC. All parameters were sensitive to the duration of the pneumatic measurement, with highest overall agreement between methods after 16 sec.We conclude that, if applied correctly, the pneumatic method enables fast and inexpensive estimations of embolism resistance for a wide range of temperate, diffuse-porous species, which makes it attractive for predicting plant performance under climate change.

Farmland tree cultivation is considered an important option for enhancing wood production. In South India, the native leaf-deciduous tree species Melia dubia is popular for short-rotation plantations. Across a rainfall gradient from 420 to 2170 mm year–1, we studied 186 farmland woodlots between one and nine years in age. The objectives were to identify the main factors controlling aboveground biomass (AGB) and growth rates. A power-law growth model predicts an average stand-level AGB of 93.8 Mg ha–1 for nine-year-old woodlots. The resulting average annual AGB increment over the length of the rotation cycle is 10.4 Mg ha–1 year–1, which falls within the range reported for other tropical tree plantations. When expressing the parameters of the growth model as functions of management, climate and soil variables, it explains 65% of the variance in AGB. The results indicate that water availability is the main driver of the growth of M. dubia. Compared to the effects of water availability, the effects of soil nutrients are 26% to 60% smaller. We conclude that because of its high biomass accumulation rates in farm forestry, M. dubia is a promising candidate for short-rotation plantations in South India and beyond.

An extreme summer drought caused unprecedented tree dieback across Central Europe in 2018, highlighting the need for a better mechanistic understanding of drought-induced tree mortality. While numerous physiological risk factors have been identified, the principal mechanisms, hydraulic failure and carbon starvation, are still debated. We studied 9,435 trees from 12 temperate species planted in a diversity experiment in 2013 to assess how hydraulic traits, carbon dynamics, pest infestation, tree height and neighbourhood competition influence individual mortality risk. We observed a reduced mortality risk for trees with wider hydraulic safety margins, while a rising sugar fraction of the non-structural carbohydrate pool and bark beetle infestation were associated with higher risk. Taller trees had a lower mortality risk. The sign and magnitude of neighbourhood effects on mortality risk depended on the species-identity of the involved trees, with most species having beneficial and some having deleterious effects on their neighbours. While severe tissue dehydration causing hydraulic failure precedes drought-induced tree mortality, we show that the probability of this event depends on a series of mutually inclusive processes including pest infestation and starch depletion for osmotic adjustment, and is modulated by the size and species identity of a tree and its neighbours.

Plant hydraulic traits are key for understanding and predicting tree drought responses. Information about the degree of the traits' intra-specific variability may guide the selection of drought-resistant genotypes and is crucial for trait-based modelling approaches. For the three temperate minor broadleaf tree species Acer platanoides, Carpinus betulus and Tilia cordata, we measured xylem embolism resistance (P50 ), leaf turgor loss point (PTLP ), specific hydraulic conductivity (KS ), Huber values (HVs), and hydraulic safety margins in adult trees across a precipitation gradient. We further quantified trait variability on different organizational levels (inter-specific to within-canopy variation), and analysed its relationship to climatic and soil water availability. Although we observed a certain intra-specific trait variability (ITV) in safety-related traits (P50 , PTLP ) with higher within-tree and between-tree than between populations variability, the magnitude was small compared to inter-specific differences, which explained 78.4% and 58.3% of the variance in P50 and PTLP , respectively. In contrast, efficiency-related traits (KS , HV) showed a high ITV both within populations and within the crowns of single trees. Surprisingly, the observed ITV of all traits was neither driven by climatic nor soil water availability. In conclusion, the high degree of conservatism in safety-related traits highlights their potential for trait-based modelling approaches.

Vessel length is an important functional trait for plant hydraulics, because it determines the ratio of flow resistances posed by lumen and pit membranes and hence controls xylem hydraulic efficiency. The most commonly applied methods to estimate vessel lengths are based on the injection of silicon or paint into cut-off stem segments. The number of stained vessels in a series of cross-sections in increasing distance from the injection point is then counted. The resulting infusion profiles are used to estimate the vessel length distribution using one of several statistical algorithms. However, the basis of these algorithms has not been systematically analysed using probability theory. We derive a general mathematical expression for the expected shape of the infusion profile for a given vessel length distribution, provide analytic solutions for five candidate distributions (exponential, Erlang(2), gamma, Weibull, and log-normal), and present maximum likelihood estimators for the parameters of these distributions including implementations in R based on two potential sampling schemes (counting all injected vessels or counting the injected and empty vessels in a random subset of each cross-section). We then explore the performance of these estimators relative to other methods with Monte Carlo experiments. Our analysis demonstrates that most published methods estimate the conditional length distribution of vessels that cross an injection point, which is a size-biased version of the overall length distribution in the stem. We show the mathematical relationship between these distributions and provide methods to estimate either of them. According to our simulation experiments, vessel length distribution was best described by the more flexible models, especially the Weibull distribution. In simulations, the estimators were able to recover the parameters of the vessel length distribution if its functional form was known, achieving an overlap of 90% or more between the true and predicted length distribution when counting no more than 500 injected vessels in 10 cross-sections. This sample size nowadays can easily be reached with the help of automated image analysis.

Sap flow probes are routinely used in forest and horticulture hydrology for estimating tree water use. This requires unbiased measurements when upscaling from tree to stand level, but accuracy and comparability of different thermometric methods have been questioned. Three sap flow measuring techniques were compared against gravimetric flow measurement in cut stem segments: ‘Granier-type’ thermal dissipation probes (TDP; three different sensor types), the heat field deformation method (HFD), and the heat ratio method (HRM). For the empirical methods (TDP and HFD), new calibration parameters were estimated using a nonlinear hierarchical modelling approach. 66 stem segments from five temperate, diffuse-porous tree species (9–16 cm stem diameter, 100 cm stem length) were exposed to a wide range of flux densities by applying subatmospheric pressure (−50 to −650 hPa) analogous to natural flow conditions in the field. All TDP probes underestimated flux density by 23–45% when calculated with Granier's original calibration parameters, with the deviation increasing with flux rate. The accuracy was significantly improved by estimating new calibration parameters, especially for probes differing from Granier's original sensor design. Species-specific parameters further improved accuracy, although the species differences might partially be explained by variation in the observed ranges of sap flux. The HFD sensor overestimated gravimetric flow by ∼11%; empirical calibration did not improve its accuracy compared to the manufacturer's equation. At low to medium flow rates, the HRM system achieved higher accuracy than the other probes (0.8% underestimation), while performing poorly at high flux rates under our measurement settings (energy input of 25 J). Both for TDP and HFD sensors, we observed a surprisingly large variability in calibration parameters between different stems of the same species. We conclude that (i) TDP and HFD sensors require species-specific calibration to measure sap flux with high accuracy, (ii) the original Granier equation cannot be used for TDP probes with deviating design, and (iii), at low to medium flow rates, the highest accuracy can be achieved with HRM sensors. Our results help to increase the accuracy of tree sap flow measurements with thermal dissipation probes, and to assess various levels of errors related to the different thermometric methods. This is important when synthesizing forest transpiration data on regional and global scales.

Studying how cambial age and axial height affects wood anatomical traits may improve our understanding of xylem hydraulics, heartwood formation and axial growth. Radial strips were collected from six different heights (0&ndash, 11.3 m) along the main trunk of three Manchurian catalpa (Catalpa bungei) trees, yielding 88 samples. In total, thirteen wood anatomical vessel and fiber traits were observed usinglight microscopy (LM) and scanning electron microscopy (SEM), and linear models were used to analyse the combined effect of axial height, cambial age and their interaction. Vessel diameter differed by about one order of magnitude between early- and latewood, and increased significantly with both cambial age and axial height in latewood, while it was positively affected by cambial age and independent of height in earlywood. Vertical position further had a positive effect on earlywood vessel density, and negative effects on fibre wall thickness, wall thickness to diameter ratio and length. Cambial age had positive effects on the pit membrane diameter and vessel element length, while the annual diameter growth decreased with both cambial age and axial position. In contrast, early- and latewood fiber diameter were unaffected by both cambial age and axial height. We further observed an increasing amount of tyloses from sapwood to heartwood, accompanied by an increase of warty layers and amorphous deposits on cell walls, bordered pit membranes and pit apertures. This study highlights the significant effects of cambial age and vertical position on xylem anatomical traits, and confirms earlier work that cautions to take into account xylem spatial position when interpreting wood anatomical structures, and thus, xylem hydraulic functioning.

Sap flow measurements are a crucial tool for studies of plant water relations, but upscaling to tree or stand transpiration requires accounting for the radial distribution of sap flow across the stem. Because radial sap flow profiles cannot be measured with single-point measuring methods, predicting their shape based on easier to measure structural or functional traits is a potential route to cost-efficiently improve water-use estimates. We used heat field deformation (HFD) sensors to measure sap flux density at different sapwood depths on 38 trees of eight tropical tree species in secondary dry forest patches in north-west Costa Rica. Based on a Bayesian hierarchical modeling framework, we developed a flexible model for radial profiles that expresses the average profile depth and the spread of the profile around this depth as functions of tree height, wood density and stem growth rate, while allowing for random tree and species effects. From the model output, we scale up to approximate whole-tree daily water use (DWU). Our model revealed pronounced tree height effects, with taller trees having narrower sap flow profiles peaking closer to the cambium. While it explained 96% of the variance in the dataset, large fractions were attributed to random species and tree individual effects. Including functional traits as predictors improved the accuracy of predictions considerably both for new trees and new species. DWU responded positively to tree height, but not to annual stem increment and wood density. Extrapolating the average sap flux in the outermost 2 cm over the entire sapwood area overestimated DWU on average by 26% compared to estimates based on radial profiles. Our model provides a starting point for future studies aiming to improve landscape-scale water-use estimates integrating single-point sap flow measurements and radial profiles from a subset of trees.

With the prospect of climate change and more frequent El Niño-related dry spells, the drought tolerance of oil palm (Elaeis guineensis Jacq.), one of the most important tropical crop species, is of major concern. We studied the influence of soil water availability and palm height on the plasticity of xylem anatomy of oil palm fronds and their embolism resistance at well-drained and seasonally flooded riparian sites in lowland Sumatra, Indonesia. We found overall mean P12 and P50 values, i.e., the xylem pressures at 12% or 50% loss of hydraulic conductance, of −1.05 and − 1.86 MPa, respectively, indicating a rather vulnerable frond xylem of oil palm. This matches diurnal courses of stomatal conductance, which in combination with the observed low xylem safety evidence a sensitive water loss regulation. While the xylem anatomical traits vessel diameter (Dh), vessel density and potential hydraulic conductivity (Kp) were not different between the sites, palms in the moister riparian plots had on average by 0.4 MPa higher P50 values than plants in the well-drained plots. This could largely be attributed to differences in palm height between systems. As a consequence, palms of equal height had 1.3 MPa less negative P50 values in the moister riparian plots than in the well-drained plots. While palm height was positively related to P50, Dh and Kp decreased with height. The high plasticity in embolism resistance may be an element of the drought response strategy of oil palm, which, as a monocot, has a relatively deterministic hydraulic architecture. We conclude that oil palm fronds develop a vulnerable water transport system, which may expose the palms to increasing drought stress in a warmer and drier climate. However, the risk of hydraulic failure may be reduced by considerable plasticity in the hydraulic system and the environmental control of embolism resistance, and a presumably large stem capacitance.