Isohydricity and hydraulic isolation explain reduced hydraulic failure risk in an experimental tree species mixture.

Species mixture is promoted as a crucial management option to adapt forests to climate change. However, there is little consensus on how tree diversity affects tree water stress, and the underlying mechanisms remain elusive. By using a greenhouse experiment and a soil-plant-atmosphere hydraulic model, we explored whether and why mixing the isohydric Aleppo pine (Pinus halepensis, drought avoidant) and the anisohydric holm oak (Quercus ilex, drought tolerant) affects tree water stress during extreme drought. Our experiment showed that the intimate mixture strongly alleviated Q. ilex water stress while it marginally impacted P. halepensis water stress. Three mechanistic explanations for this pattern are supported by our modelling analysis. First, the difference in stomatal regulation between species allowed Q. ilex trees to benefit from additional soil water in mixture, thereby maintaining higher water potentials and sustaining gas exchange. By contrast, P. halepensis exhibited earlier water stress and stomatal regulation. Second, P. halepensis trees showed stable water potential during drought, although soil water potential strongly decreased, even when grown in a mixture. Model simulations suggested that hydraulic isolation of the root from the soil associated with decreased leaf cuticular conductance was a plausible explanation for this pattern. Third, the higher predawn water potentials for a given soil water potential observed for Q. ilex in mixture can - according to model simulations - be explained by increased soil-to-root conductance, resulting from higher fine root length. This study brings insights into the mechanisms involved in improved drought resistance of mixed species forests.

Plant hydraulics at the heart of plant, crops and ecosystem functions in the face of climate change.

Plant hydraulics is crucial for assessing the plants' capacity to extract and transport water from the soil up to their aerial organs. Along with their capacity to exchange water between plant compartments and regulate evaporation, hydraulic properties determine plant water relations, water status and susceptibility to pathogen attacks. Consequently, any variation in the hydraulic characteristics of plants is likely to significantly impact various mechanisms and processes related to plant growth, survival and production, as well as the risk of biotic attacks and forest fire behaviour. However, the integration of hydraulic traits into disciplines such as plant pathology, entomology, fire ecology or agriculture can be significantly improved. This review examines how plant hydraulics can provide new insights into our understanding of these processes, including modelling processes of vegetation dynamics, illuminating numerous perspectives for assessing the consequences of climate change on forest and agronomic systems, and addressing unanswered questions across multiple areas of knowledge.

The geophysical toolbox applied to forest ecosystems - A review.

Studying the forest subsurface is a challenge because of its heterogeneous nature and difficult access. Traditional approaches used by ecologists to characterize the subsurface have a low spatial representativity. This review article illustrates how geophysical techniques can and have been used to get new insights into forest ecology. Near-surface geophysics offers a wide range of methods to characterize the spatial and temporal variability of subsurface properties in a non-destructive and integrative way, each with its own advantages and disadvantages. These techniques can be used alone or combined to take advantage of their complementarity. Our review led us to define three topics how near-surface geophysics can support forest ecology studies: 1) detection of root systems, 2) monitoring of water quantity and dynamics, and 3) characterisation of spatial heterogeneity in subsurface properties at the stand level. The number of forest ecology studies using near-surface geophysics is increasing and this multidisciplinary approach opens new opportunities and perspectives for improving quantitative assessment of biophysical properties and exploring forest response to the environment and adaptation to climate change.

Small and slow is safe: On the drought tolerance of tropical tree species.

Understanding how evolutionary history and the coordination between trait trade-off axes shape the drought tolerance of trees is crucial to predict forest dynamics under climate change. Here, we compiled traits related to drought tolerance and the fast-slow and stature-recruitment trade-off axes in 601 tropical woody species to explore their covariations and phylogenetic signals. We found that xylem resistance to embolism (P50) determines the risk of hydraulic failure, while the functional significance of leaf turgor loss point (TLP) relies on its coordination with water use strategies. P50 and TLP exhibit weak phylogenetic signals and substantial variation within genera. TLP is closely associated with the fast-slow trait axis: slow species maintain leaf functioning under higher water stress. P50 is associated with both the fast-slow and stature-recruitment trait axes: slow and small species exhibit more resistant xylem. Lower leaf phosphorus concentration is associated with more resistant xylem, which suggests a (nutrient and drought) stress-tolerance syndrome in the tropics. Overall, our results imply that (1) drought tolerance is under strong selective pressure in tropical forests, and TLP and P50 result from the repeated evolutionary adaptation of closely related taxa, and (2) drought tolerance is coordinated with the ecological strategies governing tropical forest demography. These findings provide a physiological basis to interpret the drought-induced shift toward slow-growing, smaller, denser-wooded trees observed in the tropics, with implications for forest restoration programmes.

Coordination of stem and leaf traits define different strategies to regulate water loss and tolerance ranges to aridity.

Adaptation to drought involves complex interactions of traits that vary within and among species. To date, few data are available to quantify within-species variation in functional traits and they are rarely integrated into mechanistic models to improve predictions of species response to climate change. We quantified intraspecific variation in functional traits of two Hakea species growing along an aridity gradient in southeastern Australia. Measured traits were later used to parameterise the model SurEau to simulate a transplantation experiment to identify the limits of drought tolerance. Embolism resistance varied between species but not across populations. Instead, populations adjusted to drier conditions via contrasting sets of trait trade-offs that facilitated homeostasis of plant water status. The species from relatively mesic climate, Hakea dactyloides, relied on tight stomatal control whereas the species from xeric climate, Hakea leucoptera dramatically increased Huber value and leaf mass per area, while leaf area index (LAI) and epidermal conductance (gmin ) decreased. With trait variability, SurEau predicts the plasticity of LAI and gmin buffers the impact of increasing aridity on population persistence. Knowledge of within-species variability in multiple drought tolerance traits will be crucial to accurately predict species distributional limits.

Tree xylem water isotope analysis by Isotope Ratio Mass Spectrometry and laser spectrometry: A dataset to explore tree response to drought.

Water isotopes from plant xylem and surrounding environment are increasingly used in eco-hydrological studies. Carrière et al. [1] analyzed a dataset of water isotopes in (i) the xylem of three different tree species, (ii) the surrounding soil and drainage water and (iii) the underlying karst groundwater, to understand tree water uptake during drought in two different Mediterranean forests on karst setting. The xylem and soil water were extracted by cryogenic distillation. The full dataset was obtained with Isotope Ratio Mass Spectrometry (IRMS) and Isotope Ratio Infrared Spectrometer (IRIS), and included 219 measurements of δ2H and δ18O. Prompted by unexpected isotopic data characterized by a very negative deuterium excess, a subsample of 46 xylem samples and 9 soil water samples were double checked with both analytical techniques. IRMS and IRIS analyses yielded similar data. Therefore, the results reveal that laser spectrometry allows an accurate estimation of xylem and soil water isotopes. The dataset highlights a strong 2H depletion in xylem water for all species. Deuterium does not seem adequate to interpret ecological processes in this dataset given the strong fractionation.

The role of deep vadose zone water in tree transpiration during drought periods in karst settings - Insights from isotopic tracing and leaf water potential.

Karst environments are unusual because their dry, stony and shallow soils seem to be unfavorable to vegetation, and yet they are often covered with forests. How can trees survive in these environments? Where do they find the water that allows them to survive? This study uses midday and predawn water potentials and xylem water isotopes of branches to assess tree water status and the origin of transpired water. Monitoring was conducted during the summers of 2014 and 2015 in two dissimilar plots of Mediterranean forest located in karst environments. The results show that the three monitored tree species (Abies alba Mill, Fagus sylvatica L, and Quercus ilex L.) use deep water resources present in the karst vadose zone (unsaturated zone) more intensively during drier years. Quercus ilex, a species well- adapted to water stress, which grows at the drier site, uses the deep water resource very early in the summer season. Conversely, the two other species exploit the deep water resource only during severe drought. These results open up new perspectives to a better understanding of ecohydrological equilibrium and to improved water balance modeling in karst forest settings.

On the minimum leaf conductance: its role in models of plant water use, and ecological and environmental controls.

Contents Summary 693 I. Introduction 693 II. Comparison of various definitions and measurement techniques of minimum conductance 694 III. Cuticular conductance 695 IV. Contribution of stomata 696 V. Environmental and ecological variation in minimum conductance 696 VI. Use of minimum conductance in models 698 VII. Conclusions 703 Acknowledgements 703 References 703 SUMMARY: When the rate of photosynthesis is greatly diminished, such as during severe drought, extreme temperature or low light, it seems advantageous for plants to close stomata and completely halt water loss. However, water loss continues through the cuticle and incompletely closed stomata, together constituting the leaf minimum conductance (gmin ). In this review, we critically evaluate the sources of variation in gmin , quantitatively compare various methods for its estimation, and illustrate the role of gmin in models of leaf gas exchange. A literature compilation of gmin as measured by the weight loss of detached leaves is presented, which shows much variation in this trait, which is not clearly related to species groups, climate of origin or leaf type. Much evidence points to the idea that gmin is highly responsive to the growing conditions of the plant, including soil water availability, temperature and air humidity - as we further demonstrate with two case studies. We pay special attention to the role of the minimum conductance in the Ball-Berry model of stomatal conductance, and caution against the usual regression-based method for its estimation. The synthesis presented here provides guidelines for the use of gmin in ecosystem models, and points to clear research gaps for this drought tolerance trait.

Xylem resistance to embolism: presenting a simple diagnostic test for the open vessel artefact.

Xylem vulnerability to embolism represents an essential trait for the evaluation of the impact of hydraulics in plant function and ecology. The standard centrifuge technique is widely used for the construction of vulnerability curves, although its accuracy when applied to species with long vessels remains under debate. We developed a simple diagnostic test to determine whether the open-vessel artefact influences centrifuge estimates of embolism resistance. Xylem samples from three species with differing vessel lengths were exposed to less negative xylem pressures via centrifugation than the minimum pressure the sample had previously experienced. Additional calibration was obtained from non-invasive measurement of embolism on intact olive plants by X-ray microtomography. Results showed artefactual decreases in hydraulic conductance (k) for samples with open vessels when exposed to a less negative xylem pressure than the minimum pressure they had previously experienced. X-Ray microtomography indicated that most of the embolism formation in olive occurs at xylem pressures below -4.0 MPa, reaching 50% loss of hydraulic conductivity at -5.3 MPa. The artefactual reductions in k induced by centrifugation underestimate embolism resistance data of species with long vessels. A simple test is suggested to avoid this open vessel artefact and to ensure the reliability of this technique in future studies.

Environmental control of carbon allocation matters for modelling forest growth.

We aimed to evaluate the importance of modulations of within-tree carbon (C) allocation by water and low-temperature stress for the prediction of annual forest growth with a process-based model. A new C allocation scheme was implemented in the CASTANEA model that accounts for lagged and direct environmental controls of C allocation. Different approaches (static vs dynamic) to modelling C allocation were then compared in a model-data fusion procedure, using satellite-derived leaf production estimates and biometric measurements at c. 104 sites. The modelling of the environmental control of C allocation significantly improved the ability of CASTANEA to predict the spatial and year-to-year variability of aboveground forest growth along regional gradients. A significant effect of the previous year's water stress on the C allocation to leaves and wood was reported. Our results also are consistent with a prominent role of the environmental modulation of sink demand in the wood growth of the studied species. Data available at large scales can inform forest models about the processes driving annual and seasonal C allocation. Our results call for a greater consideration of C allocation drivers, especially sink-demand fluctuations, for the simulations of current and future forest productivity with process-based models.

Growth duration is a better predictor of stem increment than carbon supply in a Mediterranean oak forest: implications for assessing forest productivity under climate change.

Understanding whether tree growth is limited by carbon gain (source limitation) or by the direct effect of environmental factors such as water deficit or temperature (sink limitation) is crucial for improving projections of the effects of climate change on forest productivity. We studied the relationships between tree basal area (BA) variations, eddy covariance carbon fluxes, predawn water potential (Ψpd ) and temperature at different timescales using an 8-yr dataset and a rainfall exclusion experiment in a Quercus ilex Mediterranean coppice. At the daily timescale, during periods of low temperature (< 5°C) and high water deficit (< -1.1 MPa), gross primary productivity and net ecosystem productivity remained positive whereas the stem increment was nil. Thus, stem increment appeared limited by drought and temperature rather than by carbon input. Annual growth was accurately predicted by the duration of BA increment during spring (Δtt0-t1 ). The onset of growth (t0 ) was related to winter temperatures and the summer interruption of growth (t1 ) to a threshold Ψpd value of -1.1 MPa. We suggest that using environmental drivers (i.e. drought and temperature) to predict stem growth phenology can contribute to an improvement in vegetation models and may change the current projections of Mediterranean forest productivity under climate change scenarios.

Assessing the effects of management on forest growth across France: insights from a new functional-structural model.

BACKGROUND AND AIMS: The structure of a forest stand, i.e. the distribution of tree size features, has strong effects on its functioning. The management of the structure is therefore an important tool in mitigating the impact of predicted changes in climate on forests, especially with respect to drought. Here, a new functional-structural model is presented and is used to assess the effects of management on forest functioning at a national scale. METHODS: The stand process-based model (PBM) CASTANEA was coupled to a stand structure module (SSM) based on empirical tree-to-tree competition rules. The calibration of the SSM was based on a thorough analysis of intersite and interannual variability of competition asymmetry. The coupled CASTANEA-SSM model was evaluated across France using forest inventory data, and used to compare the effect of contrasted silvicultural practices on simulated stand carbon fluxes and growth. KEY RESULTS: The asymmetry of competition varied consistently with stand productivity at both spatial and temporal scales. The modelling of the competition rules enabled efficient prediction of changes in stand structure within the CASTANEA PBM. The coupled model predicted an increase in net primary productivity (NPP) with management intensity, resulting in higher growth. This positive effect of management was found to vary at a national scale across France: the highest increases in NPP were attained in forests facing moderate to high water stress; however, the absolute effect of management on simulated stand growth remained moderate to low because stand thinning involved changes in carbon allocation at the tree scale. CONCLUSIONS: This modelling approach helps to identify the areas where management efforts should be concentrated in order to mitigate near-future drought impact on national forest productivity. Around a quarter of the French temperate oak and beech forests are currently in zones of high vulnerability, where management could thus mitigate the influence of climate change on forest yield.

The temporal response to drought in a Mediterranean evergreen tree: comparing a regional precipitation gradient and a throughfall exclusion experiment.

Like many midlatitude ecosystems, Mediterranean forests will suffer longer and more intense droughts with the ongoing climate change. The responses to drought in long-lived trees differ depending on the time scale considered, and short-term responses are currently better understood than longer term acclimation. We assessed the temporal changes in trees facing a chronic reduction in water availability by comparing leaf-scale physiological traits, branch-scale hydraulic traits, and stand-scale biomass partitioning in the evergreen Quercus ilex across a regional precipitation gradient (long-term changes) and in a partial throughfall exclusion experiment (TEE, medium term changes). At the leaf scale, gas exchange, mass per unit area and nitrogen concentration showed homeostatic responses to drought as they did not change among the sites of the precipitation gradient or in the experimental treatments of the TEE. A similar homeostatic response was observed for the xylem vulnerability to cavitation at the branch scale. In contrast, the ratio of leaf area over sapwood area (LA/SA) in young branches exhibited a transient response to drought because it decreased in response to the TEE the first 4 years of treatment, but did not change among the sites of the gradient. At the stand scale, leaf area index (LAI) decreased, and the ratios of stem SA to LAI and of fine root area to LAI both increased in trees subjected to throughfall exclusion and from the wettest to the driest site of the gradient. Taken together, these results suggest that acclimation to chronic drought in long-lived Q. ilex is mediated by changes in hydraulic allometry that shift progressively from low (branch) to high (stand) organizational levels, and act to maintain the leaf water potential within the range of xylem hydraulic function and leaf photosynthetic assimilation.

Gas exchange and leaf aging in an evergreen oak: causes and consequences for leaf carbon balance and canopy respiration.

Leaves of Mediterranean evergreens experience large variations in gas exchange rates over their life span due to aging and seasonally changing environmental conditions. Accounting for the changing respiratory physiology of leaves over time will help improve estimations of leaf and whole-plant carbon balances. Here we examined seasonal variations in light-saturated net CO(2) assimilation (A(max)), dark respiration (R(d)) and the proportional change in R(d) per 10 °C change in temperature (Q(10) of R(d)) in previous-year (PY) and current-year (CY) leaves of the broadleaved evergreen tree Quercus ilex L. A(max) and R(d) were lower in PY than in CY leaves. Differences in nitrogen between cohorts only partly explained such differences, and rates of A(max) and R(d) expressed per unit of leaf nitrogen were still significantly different between cohorts. The decline in A(max) in PY leaves did not result in the depletion of total non-structural carbohydrates, whose concentration was in fact higher in PY than CY leaves. Leaf-level carbon balance modeled from gas exchange data was positive at all ages. Q(10) of R(d) did not differ significantly between leaf cohorts; however, failure to account for distinct R(d) between cohorts misestimated canopy leaf respiration by 13% across dates when scaling up leaf measurements to the canopy. In conclusion, the decline in A(max) in old leaves that are close to or exceed their mean life span does not limit the availability of carbohydrates, which are probably needed to sustain new growth, as well as R(d) and nutrient resorption during senescence. Accounting for leaf age as a source of variation of R(d) improves the estimation of foliar respiratory carbon release at the stand scale.

Photosynthetic sensitivity to drought varies among populations of Quercus ilex along a rainfall gradient.

Drought frequency and intensity are expected to increase in the Mediterranean as a consequence of global climate change. To understand how photosynthetic capacity responds to long-term water stress, we measured seasonal patterns of stomatal (SL), mesophyll (MCL) and biochemical limitations (BL) to net photosynthesis (Amax) in three Quercus ilex (L.) populations from sites differing in annual rainfall. In the absence of water stress, stomatal conductance (gs), maximum carboxylation capacity (Vcmax), photosynthetic electron transport rate (Jmax) and Amax were similar among populations. However, as leaf predawn water potential (Ψl,pd) declined, the population from the wettest site showed steeper declines in gs, Vcmax, Jmax and Amax than those from the drier sites. Consequently, SL, MCL and BL increased most steeply in response to decreasing Ψl,pd in the population from the wettest site. The higher sensitivity of Amax to drought was primarily the result of stronger stomatal regulation of water loss. Among-population differences were not observed when gs was used instead of Ψl,pd as a drought stress indicator. Given that higher growth rates, stature and leaf area index were observed at the wettest site, we speculate that hydraulic architecture may explain the greater drought sensitivity of this population. Collectively, these results highlight the importance of considering among-population differences in photosynthetic responses to seasonal drought in large scale process-based models of forest ecosystem function.