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Understanding the physiological and biochemical responses of tree seedlings under extreme drought stress, along with recovery during rewatering, and potential intra-species differences, will allow us to more accurately predict forest responses under future climate change. Here, we selected seedlings from four provenances (AH (Anhui), JX (Jiangxi), HN (Hunan) and GX (Guangxi)) of Schima superba and carried out a simulated drought-rewatering experiment in a field-based rain-out shelter. Seedlings were progressively dried until they reached 50% and 88% loss of xylem hydraulic conductivity (PLC) (i.e. P 50 and P 88 ), respectively, before they were rehydrated and maintained at field capacity for 30 days. Leaf photosynthesis (A sat ), water status, activity of superoxide dismutase (SOD), and proline (Pro) concentration were monitored and their associations were determined. Increasing drought significantly reduced A sat , relative water content (RWC) and SOD activity in all provenances, and Pro concentration was increased to improve water retention; all four provenances exhibited similar response patterns, associated with similar leaf ultrastructure at pre-drought. Upon rewatering, physiological and biochemical traits were restored to well-watered control values in P 50 -stressed seedlings. In P 88 -stressed seedlings, Pro was restored to control values, while SOD was not fully recovered. The recovery pattern differed partially among provenances. There was a progression of recovery following watering, with RWC firstly recovered, followed by SOD and Pro, and then A sat , but with significant associations among these traits. Collectively, the intra-specific differences of S. superba seedlings in recovery of physiology and biochemistry following rewatering highlight the need to consider variations within a given tree species coping with future more frequent drought stress., (© 2024. The Author(s).)

Drought-induced tree mortality may increase with ongoing climate change. Unraveling the links between stem hydraulics and mortality thresholds, and the effects of intraspecific variation, remain important unresolved issues. We conducted a water manipulation experiment in a rain-out shelter, using four provenances of Schima superba originating from a gradient of annual precipitation (1124-1796 mm) and temperature (16.4-22.4°C). Seedlings were droughted to three levels of percentage loss of hydraulic conductivity (i.e., P 50 , P 88  and P 99) and subsequently rewatered to field capacity for 30 days; traits related to water and carbon relations were measured. The lethal water potential associated with incipient mortality was between P 50 and P 88 . Seedlings exhibited similar drought responses in xylem water potential, hydraulic conductivity and gas exchange. Upon rehydration, patterns of gas exchange differed among provenances but were not related to the climate at the origin. The four provenances exhibited a similar degree of stem hydraulic recovery, which was correlated with the magnitude of antecedent drought and stem soluble sugar at the end of the drought. Results suggest that there were intraspecific differences in the capacity of S. superba seedlings for carbon assimilation during recovery, indicating a decoupling between gas exchange recovery and stem hydraulics across provenances., (© 2022 John Wiley & Sons Ltd.)

Stomatal response to environmental conditions forms the backbone of all ecosystem and carbon cycle models, but is largely based on empirical relationships. Evolutionary theories of stomatal behaviour are critical for guarding against prediction errors of empirical models under future climates. Longstanding theory holds that stomata maximise fitness by acting to maintain constant marginal water use efficiency over a given time horizon, but a recent evolutionary theory proposes that stomata instead maximise carbon gain minus carbon costs/risk of hydraulic damage. Using data from 34 species that span global forest biomes, we find that the recent carbon-maximisation optimisation theory is widely supported, revealing that the evolution of stomatal regulation has not been primarily driven by attainment of constant marginal water use efficiency. Optimal control of stomata to manage hydraulic risk is likely to have significant consequences for ecosystem fluxes during drought, which is critical given projected intensification of the global hydrological cycle., (© 2018 The Authors. Ecology Letters published by CNRS and John Wiley & Sons Ltd.)

Adaptation to future climates characterized by more frequent severe droughts requires enhanced mechanistic understanding of tree mortality. However, our knowledge of the physiological limits to withstand extreme drought, and how the coordination between water and carbon traits enhances survival, is still limited. Potted seedlings of Pinus massoniana were dehydrated to three target droughts (percentage loss of stem hydraulic conductivity of ca. 50%, 85%, and 100%; PLC50, PLC85 and PLC100) and then relieved from these target droughts by fully rewatering. Predawn and midday water potentials (Ψ), relative water content (RWC), PLC and nonstructural carbohydrates (NSC) were monitored. During drought, Ψ and RWC declined as PLC increased. Root RWC declined more rapidly than other organ RWCs, particularly after PLC50 stress. All organ NSC concentrations were above predrought values. During rewatering, water trait recovery declined as drought increased, with no mortality at PLC50 but 75% mortality at PLC85. The observed stem hydraulic recovery at PLC50 following rewatering was not correlated to NSC dynamics. Collectively, our results highlighted the primary role of hydraulic failure in Pinus massoniana seedling mortality by assessing mortality threshold and links among water status and water supply. Root RWC can be considered as a potential warning signal of P. massoniana mortality. Summary statement: Hydraulic failure and multiorgan dehydration triggered seedling mortality of Pinus massoniana, with root relative water content (RWC) exhibiting faster decline than other organ RWCs in extreme drought. [ABSTRACT FROM AUTHOR]

The live fuel moisture content (LFMC) is an important precondition for wildfire activity, yet it remains challenging to predict LFMC due to the dynamic interplay between atmospheric and hydrological conditions that determine the plant's access to, and loss of water.We monitored LFMC and a range of plant water‐use traits (predawn and midday leaf water potentials [Ψleaf]), leaf traits (specific leaf area [SLA]), hydrological status (soil water content [SWC] in the shallow layer and full profile) and atmospheric variables (air temperature, vapour pressure deficit [VPD], CO2 concentrations) in a mature eucalypt woodland at the Eucalyptus Free‐Air CO2 Enrichment (EucFACE) facility during a drought.We combined plant traits, hydrological status and atmospheric variables into a biophysical model to predict LFMC dynamics, and compared these with predictions of LFMC based on a satellite model and established relationships between Ψleaf and LFMC from pressure–volume curves.Predawn Ψleaf could be well predicted from changes in SWC, but variation in midday Ψleaf and LFMC were more responsive to atmospheric than hydrological variables. The biophysical model explained up to 89% of variability in LFMC and outperformed established approaches to predict LFMC. SLA was the single most important variable to predict LFMC, followed by VPD, which explained 33% of the remaining variability in LFMC.Our study demonstrates that the co‐variation of plant traits and atmospheric and hydrological conditions affect LFMC during drought, suggesting a new way forward for predicting LFMC by combining biophysical and satellite‐based models of LFMC with seasonal forecasts of meteorological and hydrological variables. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Subtropical tree species may experience severe drought stress due to variable rainfall under future climates. However, the capacity to restore hydraulic function post-drought might differ among co-occurring species with contrasting leaf habits (e.g., evergreen and deciduous) and have implications for future forest composition. Moreover, the links between hydraulic recovery and physiological and morphological traits related to water-carbon availability are still not well understood. Here, potted seedlings of six tree species (four evergreen and two deciduous) were grown outdoors under a rainout shelter. They grew under favorable water conditions until they were experimentally subjected to a soil water deficit leading to losses of ca. 50% of hydraulic conductivity, and then soils were re-watered to field capacity. Traits related to carbon and water relations were measured. There were differences in drought responses and recovery between species, but not as a function of evergreen or deciduous groups. Sapindus mukorossi exhibited the most rapid drought response, which was associated with a suite of physiological and morphological traits (larger plant size, the lowest hydraulic capacitance (Cbranch), higher minimum conductance (gmin) and lower HV (Huber value)). Upon re-watering, xylem water potential exhibited fast recovery in 1-3 days among species, while photosynthesis at saturating light (Asat) and stomatal conductance (gs) recovery lagged behind water potential recovery depending on species, with gs recovery being more delayed than Asat in most species. Furthermore, none of the six species exhibited significant hydraulic recovery during the 7 days re-watering period, indicating that xylem refilling was apparently limited; in addition, NSC availability had a minimal role in facilitating hydraulic recovery during this short-term period. Collectively, if water supply is limited by insignificant hydraulic recovery post-drought, the observed carbon assimilation recovery of seedlings may not be sustained over the longer term, potentially altering seedling regeneration and shifting forest species composition in subtropical China under climate change. [ABSTRACT FROM AUTHOR]

Background and Aim: Drought is the main abiotic stress affecting Mediterranean forests. Root systems are responsible for water uptake, but intraspecific variability in tree root morphology is poorly understood mainly owing to sampling difficulties. The aim of this study was to gain knowledge on the adaptive relevance of rooting traits for a widespread pine using a non-invasive, high-throughput phenotyping technique. Methods: Ground-Penetrating Radar (GPR) was used to characterize variability in coarse root features (depth, diameter and frequency) among populations of the Mediterranean conifer Pinus halepensis evaluated in a common garden. GPR records were examined in relation to aboveground growth and climate variables at origin of populations. Results: Variability was detected for root traits among 56 range-wide populations categorized into 16 ecotypes. Root diameter decreased eastward within the Mediterranean basin. In turn, root frequency, but not depth and diameter, decreased following a northward gradient. Root traits also varied with climatic variables at origin such as the ratio of summer to annual precipitation, summer temperature or solar radiation. Particularly, root frequency increased with aridity, whereas root depth and diameter were maximum for ecotypes occupying the thermal midpoint of the species distribution range. Conclusion: GPR is a high-throughput phenotyping tool that allows detection of intraspecific variation in root traits of P. halepensis and its dependencies on eco-geographic characteristics at origin, thereby informing on the adaptive relevance of root systems for the species. It is also potentially suited for inferring population divergence in resource allocation above- and belowground in forest genetic trials. [ABSTRACT FROM AUTHOR]

Stomatal closure is one of the earliest responses to water stress but residual water losses may continue through the cuticle and incomplete stomatal closure. Residual conductance (g res ) plays a large role in determining time to mortality but we currently do not understand how do drought and shade interact to alter g res because the underlying drivers are largely unknown. Furthermore, g res may play an important role in models of water use, but the exact form in which g res should be incorporated into modeling schemes is currently being discussed. Here we report the results of a study where two different oak species were experimentally subjected to highly contrasting levels of drought (resulting in 0, 50 and 80% losses of hydraulic conductivity) and radiation (photosynthetic photon flux density at 1,500 μmol m–2 s–1 or 35–45 μmol m–2 s–1). We observed that the effects of radiation and drought were interactive and species-specific and g res correlated positively with concentrations of leaf non-structural carbohydrates and negatively with leaf nitrogen. We observed that different forms of measuring g res , based on either nocturnal conductance under high atmospheric water demand or on the water mass loss of detached leaves, exerted only a small influence on a model of stomatal conductance and also on a coupled leaf gas exchange model. Our results indicate that, while understanding the drivers of g res and the effects of different stressors may be important to better understand mortality, small differences in g res across treatments and measurements exert only a minor impact on stomatal models in two closely related species. [ABSTRACT FROM AUTHOR]

Background: Understanding the mechanisms that explain the spatial distribution of conifers across biogeographical gradients is important for anticipating potential range shifts owing to global change. Classical explanations have involved trade-offs between shade and drought tolerances, but more recent studies observed that trade-offs between fire and drought tolerances could also be important. Aims: Here we propose that a contributing mechanism to explain how conifer species are distributed across productivity gradients - with marked variation in the incidence of fire - involves a trade-off between allocation to bark, which serves to protect against fire, or to embolism resistance, which serves to protect against drought. Methods: We compiled information from different datasets and performed regression analyses. Results: We observed a trade-off between bark thickness and embolism resistance in conifer species such that species show either large investments of carbon to the bark or have thinner barks but xylem resistant to embolism; we did not observe conifer species concomitantly showing high fire tolerance and embolism resistance. Conclusions: This study serves as a starting point for a novel framework on how fire and drought adaptations affect conifer biogeography. Additional studies will be necessary to discover the generality of our findings by including other species of conifers, for example those in the Southern Hemisphere. [ABSTRACT FROM AUTHOR]

Abstract: Over the past 15 years, 3 million hectares of forests have been converted into shrublands or grasslands in the Mediterranean countries of the European Union. Fire and drought are the main drivers underlying this deforestation. Here we present a conceptual framework for the process of fire‐induced deforestation based on the interactive effects of fire and drought across three hierarchical scales: resistance in individuals, resilience in populations, and transitions to a new state. At the individual plant level, we review the traits that confer structural and physiological resistance, as well as allow for resprouting capacity: deforestation can be initiated when established individuals succumb to fire. After individuals perish, the second step toward deforestation requires a limited resilience from the population, that is, a reduced ability of that species to regenerate after fire. If individuals die after fire and the population fails to recover, then a transition to a new state will occur. We document trade‐offs between drought survival and fire survival, as embolism resistance is negatively correlated with fire tolerance in conifers and leaf shedding or drought deciduousness, a process that decreases water consumption at the peak of the dry season, temporally increases crown flammability. Propagule availability and establishment control resilience after mortality, but different hypotheses make contrasting predictions on the drivers of post‐fire establishment. Mycorrhizae play an additional role in modulating the response by favoring recovery through amelioration of the nutritional and water status of resprouts and new germinants. So far, resprouter species such as oaks have provided a buffer against deforestation in forests dominated by obligate seeder trees, when present in high enough density in the understory. While diversifying stands with resprouters is often reported as advantageous for building resilience, important knowledge gaps exist on how floristic composition interacts with stand flammability and on the “resprouter exhaustion syndrome,” a condition where pre‐fire drought stress, or short fire return intervals, seriously restrict post‐fire resprouting. Additional attention should be paid to the onset of novel fire environments in previously fire‐free environments, such as high altitude forests, and management actions need to accommodate this complexity to sustain Mediterranean forests under a changing climate. [ABSTRACT FROM AUTHOR]

Understanding the mechanisms underlying the response of different plant functional types to current and projected changes in rainfall is particularly important in drought-prone areas like the Mediterranean. Here, we report the responses of two species with contrasting leaf characteristics and post-fire regeneration strategies (Cistus ladanifer L., malacophyllous, seeder; Erica arborea L., sclerophyllous, resprouter) to a manipulative field experiment that simulated a severe drought (45% reduction of historical average rainfall). We measured monthly changes in relative growth rate (RGR), specific leaf area (SLA), bulk leaf carbon isotope composition (Δ[sup 13]C), predawn water potential (Ψ[sub pd]), photosynthetic gas exchange, bulk modulus of elasticity and osmotic potential at maximum turgor (?). Temporal (monthly) changes in RGR of C. ladanifer were correlated with all measured leaf traits (except Π) and followed Ψ[sub pd] variation. However, the temporal pattern of RGR in E. arborea was largely unrelated to water availability. SLA monthly variation reflected RGR variation reasonably well in C. ladanifer, but not in E. arborea, in which shoot growth and Δ[sup 13]C increased at the time of maximum water stress in late summer. The relationship between water availability, and RGR and carbon assimilation in C. ladanifer, and the lack of any relationship in E. arborea suggest that the former has an enhanced capacity to harness unpredictable rainfall pulses compared with the latter. These contrasting responses to water availability indicate that the projected changes in rainfall with global warming could alter the competitive ability of these two species, and contribute to changes in plant dominance in Mediterranean shrublands. Global warming will cause an increase in the intensity and frequency of droughts in Mediterranean areas. We report the responses of two species with contrasting leaf characteristics and post-fire regeneration strategies during a severe simulated drought. We found a contrasting response to water availability, indicating that the projected changes could alter the competitive ability of these species and contribute to changes in plant dominance in Mediterranean shrublands. [ABSTRACT FROM AUTHOR]

Highlights • Live fuel moisture content is an important predictor of forest flammability. • Fuel moisture varies due to differences in plant access to water (rooting depth). • Physiological responses to soil dryness also influence fuel moisture content. • Predawn leaf water potential provides a good predictor of live fuel moisture. • Regional models of live fuel moisture content may be improved with physiology. Abstract The moisture content of live fuels is an important determinant of forest flammability. Current approaches for modelling live fuel moisture content typically focus on the use of drought indices. However, these have mixed success partly because of species-specific differences in drought responses. Here we seek to understand the physiological mechanisms driving changes in live fuel moisture content, and to investigate the potential for incorporating plant physiological traits into live fuel moisture models. We measured the dynamics of leaf moisture content, access to water resources (through stable isotope analyses) and physiological traits (including leaf water potential, stomatal conductance, and cellular osmotic and elastic adjustments) across a fire season in a Mediterranean mixed forest in Catalonia, NE Spain. We found that differences in both seasonal variation and minimum values of live fuel moisture content were a function of access to water resources and plant physiological traits. Specifically, those species with the lowest minimum moisture content and largest seasonal variation in moisture (Cistus albidus : 49–137% and Rosmarinus officinalis : 47–144%) were most reliant on shallow soil water and had the lowest values of predawn leaf water potential. Species with the smallest variation in live fuel moisture content (Pinus nigra : 96–116% and Quercus ilex : 56–91%) exhibited isohydric behaviour (little variation in midday leaf water potential, and relatively tight regulation of stomata in response to soil drying). Of the traits measured, predawn leaf water potential provided the strongest predictor of live fuel moisture content (R 2 = 0.63, AIC = 249), outperforming two commonly used drought indices (both with R 2 = 0.49, AIC = 258). This is the first study to explicitly link fuel moisture with plant physiology and our findings demonstrate the potential and importance of incorporating ecophysiological plant traits to investigating seasonal changes in fuel moisture and, more broadly, forest flammability. [ABSTRACT FROM AUTHOR]

• Live Fuel Moisture Content (LFMC) is an essential constraint of wildfire activity. • LFMC estimated from weather/vegetation indices lacks species-specific predictions. • We use a water balance model to predict species-specific LFMC from water potential. • Our model shows an improvement in predictive performance over existing approaches. • Our approach can be implemented within large-scale fire danger forecast systems. Live Fuel Moisture Content (LFMC) is one of the main factors affecting forest ignitability as it determines the availability of existing live fuel to burn. Currently, LFMC is monitored through spectral vegetation indices or inferred from meteorological drought indices. While useful, neither approach provides mechanistic insights into species-specific LFMC variation and they are limited in the ability to forecast LFMC under altered future climates. Here, we developed a semi-mechanistic model to predict daily variation in LFMC across woody species from different functional types by adjusting a soil water balance model which estimates predawn leaf water potential (Ψ pd). Our overarching goal was to balance the trade-off between biological realism, which enhances model applicability, and parameterization complexity, which may limit its value within operational settings. After calibration, model predictions were validated against a dataset comprising 1659 LFMC observations across peninsular Spain, belonging to different functional types and from contrasting climates. The overall goodness of fit for our model (R 2 = 0.5) was better than that obtained by an existing models based on drought indices (R 2 = 0.3) or spectral vegetation indices (R 2 = 0.1). We observed the best predictive performance for seeding shrubs (R 2 = 0.6) followed by trees (R 2 = 0.5) and resprouting shrubs (R 2 = 0.4). Through its relatively simple parameterization, the approach developed here may pave the way for a new generation of process-based models that can be used for operational purposes within fire risk mitigation scenarios. [ABSTRACT FROM AUTHOR]