Relationships between xylem embolism and tree functioning during drought, recovery, and recurring drought in Aleppo pine.

Recent findings suggest that trees can survive high levels of drought-induced xylem embolism. In many cases, the embolism is irreversible and, therefore, can potentially affect post-drought recovery and tree function under recurring droughts. We examined the development of embolism in potted Aleppo pines, a common species in hot, dry Mediterranean habitats. We asked (1) how post-drought recovery is affected by different levels of embolism and (2) what consequences this drought-induced damage has under a recurring drought scenario. Young trees were dehydrated to target water potential (Ψx ) values of -3.5, -5.2 and -9.5 MPa (which corresponded to ~6%, ~41% and ~76% embolism), and recovery of the surviving trees was measured over an 8-months period (i.e., embolism, leaf gas-exchange, Ψx ). An additional group of trees was exposed to Ψx of -6.0 MPa, either with or without preceding drought (Ψx of -5.2 MPa) to test the effect of hydraulic damage during repeated drought. Trees that reached -9.5 MPa died, but none from the other groups. Embolism levels in dying trees were on average 76% of conductive xylem and no tree was dying below 62% embolism. Stomatal recovery was negatively proportional to the level of hydraulic damage sustained during drought, for at least a month after drought relief. Trees that experienced drought for the second time took longer to reach fatal Ψx levels than first-time dehydrating trees. Decreased stomatal conductance following drought can be seen as "drought legacy," impeding recovery of tree functioning, but also as a safety mechanism during a consecutive drought.

Anatomical adjustments of the tree hydraulic pathway decrease canopy conductance under long-term elevated CO2.

The cause of reduced leaf-level transpiration under elevated CO2 remains largely elusive. Here, we assessed stomatal, hydraulic, and morphological adjustments in a long-term experiment on Aleppo pine (Pinus halepensis) seedlings germinated and grown for 22-40 months under elevated (eCO2; c. 860 ppm) or ambient (aCO2; c. 410 ppm) CO2. We assessed if eCO2-triggered reductions in canopy conductance (gc) alter the response to soil or atmospheric drought and are reversible or lasting due to anatomical adjustments by exposing eCO2 seedlings to decreasing [CO2]. To quantify underlying mechanisms, we analyzed leaf abscisic acid (ABA) level, stomatal and leaf morphology, xylem structure, hydraulic efficiency, and hydraulic safety. Effects of eCO2 manifested in a strong reduction in leaf-level gc (-55%) not caused by ABA and not reversible under low CO2 (c. 200 ppm). Stomatal development and size were unchanged, while stomatal density increased (+18%). An increased vein-to-epidermis distance (+65%) suggested a larger leaf resistance to water flow. This was supported by anatomical adjustments of branch xylem having smaller conduits (-8%) and lower conduit lumen fraction (-11%), which resulted in a lower specific conductivity (-19%) and leaf-specific conductivity (-34%). These adaptations to CO2 did not change stomatal sensitivity to soil or atmospheric drought, consistent with similar xylem safety thresholds. In summary, we found reductions of gc under elevated CO2 to be reflected in anatomical adjustments and decreases in hydraulic conductivity. As these water savings were largely annulled by increases in leaf biomass, we do not expect alleviation of drought stress in a high CO2 atmosphere.

Accuracy, realism and general applicability of European forest models.

Forest models are instrumental for understanding and projecting the impact of climate change on forests. A considerable number of forest models have been developed in the last decades. However, few systematic and comprehensive model comparisons have been performed in Europe that combine an evaluation of modelled carbon and water fluxes and forest structure. We evaluate 13 widely used, state-of-the-art, stand-scale forest models against field measurements of forest structure and eddy-covariance data of carbon and water fluxes over multiple decades across an environmental gradient at nine typical European forest stands. We test the models' performance in three dimensions: accuracy of local predictions (agreement of modelled and observed annual data), realism of environmental responses (agreement of modelled and observed responses of daily gross primary productivity to temperature, radiation and vapour pressure deficit) and general applicability (proportion of European tree species covered). We find that multiple models are available that excel according to our three dimensions of model performance. For the accuracy of local predictions, variables related to forest structure have lower random and systematic errors than annual carbon and water flux variables. Moreover, the multi-model ensemble mean provided overall more realistic daily productivity responses to environmental drivers across all sites than any single individual model. The general applicability of the models is high, as almost all models are currently able to cover Europe's common tree species. We show that forest models complement each other in their response to environmental drivers and that there are several cases in which individual models outperform the model ensemble. Our framework provides a first step to capturing essential differences between forest models that go beyond the most commonly used accuracy of predictions. Overall, this study provides a point of reference for future model work aimed at predicting climate impacts and supporting climate mitigation and adaptation measures in forests.

Leaf Shedding and Non-Stomatal Limitations of Photosynthesis Mitigate Hydraulic Conductance Losses in Scots Pine Saplings During Severe Drought Stress.

During drought, trees reduce water loss and hydraulic failure by closing their stomata, which also limits photosynthesis. Under severe drought stress, other acclimation mechanisms are trigged to further reduce transpiration to prevent irreversible conductance loss. Here, we investigate two of them: the reversible impacts on the photosynthetic apparatus, lumped as non-stomatal limitations (NSL) of photosynthesis, and the irreversible effect of premature leaf shedding. We integrate NSL and leaf shedding with a state-of-the-art tree hydraulic simulation model (SOX+) and parameterize them with example field measurements to demonstrate the stress-mitigating impact of these processes. We measured xylem vulnerability, transpiration, and leaf litter fall dynamics in Pinus sylvestris (L.) saplings grown for 54 days under severe dry-down. The observations showed that, once transpiration stopped, the rate of leaf shedding strongly increased until about 30% of leaf area was lost on average. We trained the SOX+ model with the observations and simulated changes in root-to-canopy conductance with and without including NSL and leaf shedding. Accounting for NSL improved model representation of transpiration, while model projections about root-to-canopy conductance loss were reduced by an overall 6%. Together, NSL and observed leaf shedding reduced projected losses in conductance by about 13%. In summary, the results highlight the importance of other than purely stomatal conductance-driven adjustments of drought resistance in Scots pine. Accounting for acclimation responses to drought, such as morphological (leaf shedding) and physiological (NSL) adjustments, has the potential to improve tree hydraulic simulation models, particularly when applied in predicting drought-induced tree mortality.

Heatwave frequency and seedling death alter stress-specific emissions of volatile organic compounds in Aleppo pine.

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.

Increasing aridity will not offset CO2 fertilization in fast-growing eucalypts with access to deep soil water.

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.

Assessing model performance via the most limiting environmental driver in two differently stressed pine stands.

Climate change will impact 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.

Dying by drying: Timing of physiological stress thresholds related to tree death is not significantly altered by highly elevated CO2.

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 [CO2 ] 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 eCO2 , the seedlings were grown with ample of water and nutrient supply under either highly elevated [CO2 ] (eCO2 , c. 936 ppm) or ambient (aCO2 , 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 eCO2 than aCO2 . 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 eCO2 . Overall, our study provides for the first-time insights into the chronological interplay of physiological drought thresholds under long-term acclimation to elevated [CO2 ].

Tree mortality submodels drive simulated long-term forest dynamics: assessing 15 models from the stand to global scale.

Models are pivotal for assessing future forest dynamics under the impacts of changing climate and management practices, incorporating representations of tree growth, mortality, and regeneration. Quantitative studies on the importance of mortality submodels are scarce. We evaluated 15 dynamic vegetation models (DVMs) regarding their sensitivity to different formulations of tree mortality under different degrees of climate change. The set of models comprised eight DVMs at the stand scale, three at the landscape scale, and four typically applied at the continental to global scale. Some incorporate empirically derived mortality models, and others are based on experimental data, whereas still others are based on theoretical reasoning. Each DVM was run with at least two alternative mortality submodels. Model behavior was evaluated against empirical time series data, and then, the models were subjected to different scenarios of climate change. Most DVMs matched empirical data quite well, irrespective of the mortality submodel that was used. However, mortality submodels that performed in a very similar manner against past data often led to sharply different trajectories of forest dynamics under future climate change. Most DVMs featured high sensitivity to the mortality submodel, with deviations of basal area and stem numbers on the order of 10-40% per century under current climate and 20-170% under climate change. The sensitivity of a given DVM to scenarios of climate change, however, was typically lower by a factor of two to three. We conclude that (1) mortality is one of the most uncertain processes when it comes to assessing forest response to climate change, and (2) more data and a better process understanding of tree mortality are needed to improve the robustness of simulated future forest dynamics. Our study highlights that comparing several alternative mortality formulations in DVMs provides valuable insights into the effects of process uncertainties on simulated future forest dynamics.

Short-term effects of spring prescribed burning on the understory vegetation of a Pinushalepensis forest in Northeastern Spain.

Since the 1970s, fire regimes have been modified in the Northern Mediterranean region due to profound landscape changes mostly driven by socioeconomic factors, such as rural abandonment and large-scale plantations. Both fuel accumulation and the increasing vegetation spatial continuity, combined with the expansion of the wildland-urban interface, have enhanced fire risk and the occurrence of large wildfires. This situation will likely worsen under the projected aridity increase resulting from climate change. Higher fire recurrences, in particular, are expected to cause changes in vegetation composition or structure and affect ecosystems' resilience to fire, which may lead to further land degradation. Prescribed burning is a common fuel reduction technique used for fire prevention, but for conservation and restoration purposes as well. It is still poorly accepted in the Mediterranean region since constrained by critical knowledge gaps about, in particular, its effects on the ecosystems (soil, vegetation). We studied the short-term (10months) effects on the understory vegetation of a spring prescribed burning conducted in a Pinushalepensis forest in Mediterranean climate (Northeastern Spain). Our results show that the understory plant community recovered after the burning without short term significant changes in either species richness, diversity, or floristic composition. Most vegetation structural characteristics were modified though. The burning strongly reduced shrub height, shrub and herbaceous percentage covers, and aerial shrub phytomass; especially its living fine fraction, thus resulting in a less flammable community. The treatment proved to be particularly effective for the short term control of Ulexparviflorus, a highly flammable seeder species. Moreover, the strong reduction of seeder shrubs frequency in relation to resprouters' likely promoted the resilience to fire of this plant community. From a fuel-oriented perspective, the burning caused a strong reduction of spatial continuity and surface fuel loads, leading to a less fire-prone fuel complex.