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The relationship between stomatal conductance (gₛ) and leaf water potential (Ψₗ) is key to the understanding of plant function under changing climate. The variability among tree species gave rise to selection towards either of two contrasting water management types: isohydric or anisohydric. This study explores the variability of gₛ to Ψₗ across tree species. Curves of gₛ(Ψₗ) were collected from the scientific literature for 70 woody plant species. The data set is comprised of angiosperm and gymnosperm species from all major forest biomes. The hypothesis that curves from different tree species diverge between isohydric and anisohydric behaviours was tested. Species‐specific curves formed a continuum, rather than dichotomy between isohydric and anisohydric, as confirmed by distribution models. Alternatively, the water potential at 50% of the maximum gₛ (Ψgₛ50) was used to quantitatively compare between species. A major difference emerged among xylem anatomy classes whereby ring‐porous species had higher absolute gₛ at Ψₗ < −2 MPa than diffuse‐porous and coniferous species. A positive, linear correlation was shown between Ψgₛ50 and Ψₗ at 50% loss of xylem conductivity. The results suggest that stomatal sensitivity to leaf water potential strongly relates to xylem characteristics. The use of Ψgₛ50 offers a quantitative alternative to the current, yet biased, distinction between isohydric and anisohydric species.

Plant function requires effective mechanisms to regulate water transport at a variety of scales. Here, we develop a new theoretical framework describing plant responses to drying soil, based on the relationship between midday and predawn leaf water potentials. The intercept of the relationship (Λ) characterizes the maximum transpiration rate per unit of hydraulic transport capacity, whereas the slope (σ) measures the relative sensitivity of the transpiration rate and plant hydraulic conductance to declining water availability. This framework was applied to a newly compiled global database of leaf water potentials to estimate the values of Λ and σ for 102 plant species. Our results show that our characterization of drought responses is largely consistent within species, and that the parameters Λ and σ show meaningful associations with climate across species. Parameter σ was ≤1 in most species, indicating a tight coordination between the gas and liquid phases of water transport, in which canopy transpiration tended to decline faster than hydraulic conductance during drought, thus reducing the pressure drop through the plant. The quantitative framework presented here offers a new way of characterizing water transport regulation in plants that can be used to assess their vulnerability to drought under current and future climatic conditions.

Climatic stresses limit plant growth and productivity. In the past decade, tree improvement programs were mainly focused on yield but it is obvious that enhanced stress resistance is also required. In this review we highlight important drought avoidance and tolerance mechanisms in forest trees. Genomes of economically important trees species with divergent resistance mechanisms can now be exploited to uncover the mechanistic basis of long-term drought adaptation at the whole plant level. Molecular tree physiology indicates that osmotic adjustment, antioxidative defense and increased water use efficiency are important targets for enhanced drought tolerance at the cellular and tissue level. Recent biotechnological approaches focused on overexpression of genes involved in stress sensing and signaling, such as the abscisic acid core pathway, and down-stream transcription factors. By this strategy, a suite of defense systems was recruited, generally enhancing drought and salt stress tolerance under laboratory conditions. However, field studies are still scarce. Under field conditions trees are exposed to combinations of stresses that vary in duration and magnitude. Variable stresses may overrule the positive effect achieved by engineering an individual defense pathway. To assess the usability of distinct modifications, large-scale experimental field studies in different environments are necessary. To optimize the balance between growth and defense, the use of stress-inducible promoters may be useful. Future improvement programs for drought resistance will benefit from a better understanding of the intricate networks that ameliorate molecular and ecological traits of forest trees.

Species-specific responses of plant intrinsic water-use efficiency (iWUE) to multiple environmental drivers associated with climate change, including soil moisture (θ), vapor pressure deficit (D), and atmospheric CO2 concentration (c a), are poorly understood. We assessed how the iWUE and growth of several species of deciduous trees that span a gradient of isohydric to anisohydric water-use strategies respond to key environmental drivers (θ, D and c a). iWUE was calculated for individual tree species using leaf-level gas exchange and tree-ring δ13C in wood measurements, and for the whole forest using the eddy covariance method. The iWUE of the isohydric species was generally more sensitive to environmental change than the anisohydric species was, and increased significantly with rising D during the periods of water stress. At longer timescales, the influence of c a was pronounced for isohydric tulip poplar but not for others. Trees’ physiological responses to changing environmental drivers can be interpreted differently depending on the observational scale. Care should be also taken in interpreting observed or modeled trends in iWUE that do not explicitly account for the influence of D.

Deciduous and evergreen trees are usually considered the main coexisting functional groups in seasonally dry tropical forests (SDTF). We compared leaf and stem traits of 22 woody species in the Brazilian Caatinga to investigate whether deciduous (DC) and evergreen (EV) species have divergent water-use strategies. Our hypothesis was that DC trees compensate for their short leaf longevity by being less conservative in water use and showing higher variation in the seasonal water potential after leaf shedding. Evergreen species should exhibit a highly conservative water use strategy, which reduces variations in seasonal water potential and the negative effects of desiccation. Our leaf dynamics results indicate that the crown area of DC trees is more sensitive to air and soil drought, whereas EV trees are only sensitive to soil drought. Deciduous species exhibit differences in a set of leaf traits confirming their acquisitive strategy, which contrasts with evergreen species. However, when stomatal traits are considered, we found that DC and EV have similar stomatal regulation strategies (partially isohydric). We also found divergent physiological strategies within DC. For high wood density DC, the xylem water potential (Ψxylem) continued to drop during the dry season. We also found a negative linear relationship between leaf life span (LL) and the transpiration rate per unit of hydraulic conductivity (Λ), indicating that species with high LL are less vulnerable to hydraulic conductivity loss than early-deciduous species. Collectively, our results indicate divergence in the physiology of deciduous species, which suggests that categorizing species based solely on their leaf phenology may be an oversimplification.

BACKGROUND: A large proportion of vineyards are located in regions with seasonal drought (e.g. Mediterranean-type climates) where soil and atmospheric water deficits, together with high temperatures, exert large constraints on yield and quality. The increasing demand for vineyard irrigation requires an improvement in the efficiency of water use. Deficit irrigation has emerged as a potential strategy to allow crops to withstand mild water stress with little or no decreases of yield, and potentially a positive impact on fruit quality. Understanding the physiological and molecular bases of grapevine responses to mild to moderate water deficits is fundamental to optimize deficit irrigation management and identify the most suitable varieties to those conditions. SCOPE: How the whole plant acclimatizes to water scarcity and how short- and long-distance chemical and hydraulic signals intervene are reviewed. Chemical compounds synthesized in drying roots are shown to act as long-distance signals inducing leaf stomatal closure and/or restricting leaf growth. This explains why some plants endure soil drying without significant changes in shoot water status. The control of plant water potential by stomatal aperture via feed-forward mechanisms is associated with 'isohydric' behaviour in contrast to 'anysohydric' behaviour in which lower plant water potentials are attained. This review discusses differences in this respect between grapevines varieties and experimental conditions. Mild water deficits also exert direct and/or indirect (via the light environment around grape clusters) effects on berry development and composition; a higher content of skin-based constituents (e.g. tannins and anthocyanins) has generally being reported. Regulation under water deficit of genes and proteins of the various metabolic pathways responsible for berry composition and therefore wine quality are reviewed.

The classical view that the drought-related hormone ABA simply acts locally at the guard cell level to induce stomatal closure is questioned by differences between isolated epidermis and intact leaves in stomatal response to several stimuli. We tested the hypothesis that ABA mediates, in addition to a local effect, a remote effect in planta by changing hydraulic regulation in the leaf upstream of the stomata. By gravimetry, porometry to water vapour and argon, and psychrometry, we investigated the effect of exogenous ABA on transpiration, stomatal conductance and leaf hydraulic conductance of mutants described as ABA-insensitive at the guard cell level. We show that foliar transpiration of several ABA-insensitive mutants decreases in response to ABA. We demonstrate that ABA decreases stomatal conductance and down-regulates leaf hydraulic conductance in both the wildtype Col-0 and the ABA-insensitive mutant ost2-2. We propose that ABA promotes stomatal closure in a dual way via its already known biochemical effect on guard cells and a novel, indirect hydraulic effect through a decrease in water permeability within leaf vascular tissues. Variability in sensitivity of leaf hydraulic conductance to ABA among species could provide a physiological basis to the isohydric or anisohydric behaviour.

Hydraulic theory suggests that tall trees are at greater risk of drought-triggered death caused by hydraulic failure than small trees. In addition the drop in growth, observed in several tree species prior to death, is often interpreted as an early-warning signal of impending death. We test these hypotheses by comparing size, growth, and wood-anatomy patterns of living and now-dead trees in two Italian oak forests showing recent mortality episodes. The mortality probability of trees is modeled as a function of recent growth and tree size. Drift-diffusion-jump (DDJ) metrics are used to detect early-warning signals. We found that the tallest trees of the anisohydric Italian oak better survived drought contrary to what was predicted by the theory. Dead trees were characterized by a lower height and radial-growth trend than living trees in both study sites. The growth reduction of now-dead trees started about 10 years prior to their death and after two severe spring droughts during the early 2000s. This critical transition in growth was detected by DDJ metrics in the most affected site. Dead trees were also more sensitive to drought stress in this site indicating different susceptibility to water shortage between trees. Dead trees did not form earlywood vessels with smaller lumen diameter than surviving trees but tended to form wider latewood vessels with a higher percentage of vessel area. Since living and dead trees showed similar competition we did not expect that moderate thinning and a reduction in tree density would increase the short-term survival probability of trees.

In the face of water stress, plants evolved with different abilities to limit the decrease in leaf water potential, notably in the daytime (ΨM). So-called isohydric species efficiently maintain high ΨM, whereas anisohydric species cannot prevent ΨM from dropping as soil water deficit develops. The genetic and physiological origins of these differences in (an)isohydric behaviours remain to be clarified. This is of particular interest within species such as Vitis vinifera L. where continuous variation in the level of isohydry has been observed among cultivars. With this objective, a 2 year experiment was conducted on the pseudo-F1 progeny from a cross between the two widespread cultivars Syrah and Grenache using a phenotyping platform coupled to a controlled-environment chamber. Potted plants of all the progeny were analysed for ΨM, transpiration rate, and soil-to-leaf hydraulic conductance, under both well-watered and water deficit conditions. A high genetic variability was found for all the above traits. Four quantitative trait loci (QTLs) were detected for ΨM under water deficit conditions, and 28 other QTLs were detected for the different traits in either condition. Genetic variation in ΨM maintenance under water deficit weakly correlated with drought-induced reduction in transpiration rate in the progeny, and QTLs for both traits did not completely co-localize. This indicates that genetic variation in the control of ΨM under water deficit was not due simply to variation in transpiration sensitivity to soil drying. Possible origins of the diversity in (an)isohydric behaviours in grapevine are discussed on the basis of concurrent variations in soil-to-leaf hydraulic conductance and stomatal control of transpiration. This study on a grapevine mapping population shows that isohydric or anisohydric behaviour is under genetic control and is not simply controlled by transpiration response to soil drought.

Key messageNorway spruce operates with larger hydraulic safety margins (HSM) than beech and Douglas-fir despite the known drought sensitivity of spruce, questioning a pivotal role of HSM in drought tolerance.The exceptional 2018/2019 drought exposed Central Europe’s forests to severe stress, highlighting the need to better understand stomatal regulation strategies and their relationship to xylem safety under extreme drought. We studied diurnal, seasonal, and inter-annual variation in stomatal conductance (gs) and leaf water potential (ΨLeaf) in co-occurring European beech (F. sylvatica), Norway spruce (P. abies), and Douglas-fir (P. menziesii) trees in the two summers and related them to hydraulic traits characterizing drought resistance. In 2018, F. sylvatica exhibited a continuous ΨLeaf decline from June to September, as is characteristic for an anisohydric strategy, while P. abies closed stomata early and reached the least negative ΨLeaf-values at the end of summer. P. menziesii showed low ΨLeaf-values close to P12 (the xylem pressure at onset of embolism) already in July. Both conifers closed stomata when approaching P12 and maintained low gs-levels throughout summer, indicative for isohydric regulation. In 2019, all three species showed a linear decline in ΨLeaf, but F. sylvatica crossed P12 in contrast to the conifers. The three species exhibited similar water potentials at turgor loss point (− 2.44 to − 2.51 MPa) and branch P50 (xylem pressure at 50% loss of hydraulic conductance; − 3.3 to − 3.8 MPa). Yet, F. sylvatica and P. menziesii operated with smaller hydraulic safety margins (HSM means: 0.79 and 0.77 MPa) than P. abies (1.28 MPa). F. sylvatica reduced leaf size and specific leaf area in 2019 and increased Huber value. Our species comparison during extreme drought contradicts the general assumption that conifers operate with larger HSMs than angiosperm trees. Contrary to expectation, P. abies appeared as hydraulically less vulnerable than Douglas-fir.