Hydraulic Vulnerability Research Articles

Within-ring variability of wood structure and its relationship to drought sensitivity in Norway spruce trunks

ABSTRACTRelationships between hydraulic vulnerability expressed asP50(the air pressure causing 50% loss of hydraulic conductivity) and within-ring differences in wood density (WD) and anatomical features were investigated with the aim to find efficient proxies forP50relating to functional aspects. WD and tracheid dimensions were measured with SilviScan on Norway spruce (Picea abies(L.) Karst.) trunk wood.P50was strongly related to mean WD (r= -0.64) and conduit wall reinforcement ((t/b)2), the square of the ratio between the tracheid double wall thickness (t) and the lumen width (b), where use of tangential lumen width ((t/bt)2) gave better results (r= -0.54) than radial lumen width (r= -0.31). The correlations ofP50with earlywood (EW), transition wood (TW) and latewood (LW) traits were lower than with the specimen averages, both for WD (r= -0.60 for WDEW,r= -0.56 for WDTW,r= -0.23 for WDLW) and all anatomical traits. The loss of hydraulic conductivity was addressed as a dynamic process and was simulated by defining consecutive phases of 5% theoretical conductivity loss. WD and tracheid traits were calculated and correlated withP50values of each specimen. Tightest correlations were found for (t/bt)2, at relative cumulated theoretical conductivities until 45 to 50% (r= -0.75).We conclude that WD is one of the best available proxies forP50, but does not necessarily reflect the mechanism behind resistance to cavitation. The new trait, based on estimation of conductivity loss as a dynamic process, provided even stronger correlations.

Insights from invivo micro-CT analysis: testing the hydraulic vulnerability segmentation in Acer pseudoplatanus and Fagus sylvatica seedlings.

Summary The seedling stage is the most susceptible one during a tree′s life. Water relations may be crucial for seedlings due to their small roots, limited water buffers and the effects of drought on water transport. Despite obvious relevance, studies on seedling xylem hydraulics are scarce as respective methodical approaches are limited.Micro‐CT scans of intact Acer pseudoplatanus and Fagus sylvatica seedlings dehydrated to different water potentials (Ψ) allowed the simultaneous observation of gas‐filled versus water‐filled conduits and the calculation of percentage loss of conductivity (PLC) in stems, roots and leaves (petioles or main veins). Additionally, anatomical analyses were performed and stem PLC measured with hydraulic techniques.In A. pseudoplatanus, petioles showed a higher Ψ at 50% PLC (Ψ50 −1.13MPa) than stems (−2.51 MPa) and roots (−1.78 MPa). The main leaf veins of F. sylvatica had similar Ψ50 values (−2.26 MPa) to stems (−2.74 MPa) and roots (−2.75 MPa). In both species, no difference between root and stems was observed. Hydraulic measurements on stems closely matched the micro‐CT based PLC calculations.Micro‐CT analyses indicated a species‐specific hydraulic architecture. Vulnerability segmentation, enabling a disconnection of the hydraulic pathway upon drought, was observed in A. pseudoplatanus but not in the especially shade‐tolerant F. sylvatica. Hydraulic patterns could partly be related to xylem anatomical traits.

The Causes of Leaf Hydraulic Vulnerability and Its Influence on Gas Exchange in Arabidopsis thaliana.

The influence of the dynamics of leaf hydraulic conductance (K leaf) diurnally and during dehydration on stomatal conductance and photosynthesis remains unclear. Using the model species Arabidopsis (Arabidopsis thaliana ecotype Columbia-0), we applied a multitiered approach including physiological measurements, high-resolution x-ray microcomputed tomography, and modeling at a range of scales to characterize (1) K leaf decline during dehydration; (2) its basis in the hydraulic conductances of leaf xylem and outside-xylem pathways (K ox); (3) the dependence of its dynamics on irradiance; (4) its impact on diurnal patterns of stomatal conductance and photosynthetic rate; and (5) its influence on gas exchange and survival under simulated drought regimes. Arabidopsis leaves showed strong vulnerability to dehydration diurnally in both gas exchange and hydraulic conductance, despite lack of xylem embolism or conduit collapse above the turgor loss point, indicating a pronounced sensitivity of K ox to dehydration. K leaf increased under higher irradiance in well-hydrated leaves across the full range of water potential, but no shift in K leaf vulnerability was observed. Modeling indicated that responses to dehydration and irradiance are likely attributable to changes in membrane permeability and that a dynamic K ox would contribute strongly to stomatal closure, improving performance, survival, and efficient water use during drought. These findings for Columbia-0 provide a baseline for assessing variation across genotypes in hydraulic traits and their influence on gas exchange during dehydration.

Coordination between leaf, stem, and root hydraulics and gas exchange in three arid-zone angiosperms during severe drought and recovery.

The ability to resist hydraulic dysfunction in leaves, stems, and roots strongly influences whether plants survive and recover from drought. However, the coordination of hydraulic function among different organs within species and their links to gas exchange during drought and recovery remains understudied. Here, we examine the interaction between gas exchange and hydraulic function in the leaves, stems, and roots of three semiarid evergreen species exposed to a cycle of severe water stress (associated with substantial cavitation) and recovery. In all species, stomatal closure occurred at water potentials well before 50% loss of stem hydraulic conductance, while in two species, leaves and/or roots were more vulnerable than stems. Following soil rewetting, leaf-level photosynthesis (Anet ) returned to prestress levels within 2-4weeks, whereas stomatal conductance and canopy transpiration were slower to recover. The recovery of Anet was decoupled from the recovery of leaf, stem, and root hydraulics, which remained impaired throughout the recovery period. Our results suggest that in addition to high embolism resistance, early stomatal closure and hydraulic vulnerability segmentation confers drought tolerance in these arid zone species. The lack of substantial embolism refilling within all major organs suggests that vulnerability of the vascular system to drought-induced dysfunction is a defining trait for predicting postdrought recovery.

Prediction of hydraulic conductivity loss from relative water loss: new insights into water storage of tree stems and branches.

More frequently occurring, drought waves call for a deeper understanding of tree hydraulics and fast and easily applicable methods to measure drought stress. The aim of this study was to establish empirical relationships between the percent loss of hydraulic conductivity (PLC) and the relative water loss (RWL) in woody stem axes with different P 50, i.e. the water potential (Ψ) that causes 50% conductivity loss. Branches and saplings of temperate conifer (Picea abies, Larix decidua) and angiosperm species (Acer campestre, Fagus sylvatica , Populus x canescens, Populus tremula, Sorbus torminalis) and trunk wood of mature P. abies trees were analyzed. P 50 was calculated from hydraulic measurements following bench top dehydration or air injection. RWL and PLC were fitted by linear, quadratic or cubic equations. Species‐ or age‐specific RWLs at P 50 varied between 10 and 25% and P 88, the Ψ that causes 88% conductivity loss, between 18 and 44%. P 50 was predicted from the relationship between Ψ and the RWL. The predictive quality for P 50 across species was almost 1:1 (r2 = 0.99). The approach presented allows thus reliable and fast prediction of PLC from RWL. Branches and saplings with high hydraulic vulnerability tended to have lower RWLs at P 50 and at P 88. The results are discussed with regard to the different water storage capacities in sapwood and survival strategies under drought stress. Potential applications are screening trees for drought sensitivity and a fast interpretation of diurnal, seasonal or drought induced changes in xylem water content upon their impact on conductivity loss.

Is xylem of angiosperm leaves less resistant to embolism than branches? Insights from microCT, hydraulics, and anatomy.

According to the hydraulic vulnerability segmentation hypothesis, leaves are more vulnerable to decline of hydraulic conductivity than branches, but whether stem xylem is more embolism resistant than leaves remains unclear. Drought-induced embolism resistance of leaf xylem was investigated based on X-ray microcomputed tomography (microCT) for Betula pendula, Laurus nobilis, and Liriodendron tulipifera, excluding outside-xylem, and compared with hydraulic vulnerability curves for branch xylem. Moreover, bordered pit characters related to embolism resistance were investigated for both organs. Theoretical P50 values (i.e. the xylem pressure corresponding to 50% loss of hydraulic conductance) of leaves were generally within the same range as hydraulic P50 values of branches. P50 values of leaves were similar to branches for L. tulipifera (-2.01 versus -2.10 MPa, respectively), more negative for B. pendula (-2.87 versus -1.80 MPa), and less negative for L. nobilis (-6.4 versus -9.2 MPa). Despite more narrow conduits in leaves than branches, mean interconduit pit membrane thickness was similar in both organs, but significantly higher in leaves of B. pendula than in branches. This case study indicates that xylem shows a largely similar embolism resistance across leaves and branches, although differences both within and across organs may occur, suggesting interspecific variation with regard to the hydraulic vulnerability segmentation hypothesis.

Elucidating the hydraulic vulnerability of the longest-lived Southern Hemisphere conifer to aridification

Drier climatic conditions will be the future scenario in many regions worldwide, including southern South America. Few studies have characterized the ecophysiological vulnerability of the endemic tree species that inhabit this area, to climate change. In this study we assessed the hydraulic vulnerability of the longest-lived tree of the Southern Hemisphere, Fitzroya cupressoides, focusing on adult trees and saplings from two highly disturbed populations: the Coastal Range (AC) and Central Depression (FN) of southern Chile, which represent contrasting site conditions. This, as a basis for the design of conservation strategies to safeguard the persistence of these endangered forests in a drier future. We assessed water potentials (WP) throughout a growing season, their relationships with environmental conditions, as well as leaf and stem (branch) traits and hydraulic safety margins. Despite that the studied summer was the second driest in the last seven decades, minimum WP were not that negative (−1.3 to −1.5 MPa); which could be partly explained by a high leaf capacitance in this species. Adult trees and saplings from both sites did not significantly differ in their WP at turgor loss point, and their associated leaf safety margins, which were relatively low in all cases. However, they significantly differed in the xylem WP causing a 50% loss of stem conductivity (P50): adults AC: −5.14, saplings AC: −2.53, adults FN: −3.71, and saplings FN: −3.87 MPa. These values led to a relatively large stem safety margin (SSM) in most cases, and their variation was not explained neither by wood density, nor by tracheid size changes. Moreover, there appears to be an ontogenic adjustment in the more restrictive site AC, which was not seen in FN. Within the continuum of species strategies to cope with water stress, Fitzroya has features of the two ends of the continuum: tissues with large SSM, and tissues that sustain milder operation pressures through capacitance. Although Fitzroya appears to be relatively resistant to water scarcity, saplings from AC, seem to be the most vulnerable to the aridification trend in southern Chile. Moreover, future drying could become a significant extra threat to the highly endangered Central Depression population. Conservation actions are urgently needed to secure the future of Fitzroya forests in southern Chile.

No role for xylem embolism or carbohydrate shortage in temperate trees during the severe 2015 drought

AbstractTemperate forests are predicted to experience an increased frequency and intensity of climate change‐induced summer droughts and heat waves in the near future. Yet, while previous studies clearly showed a high drought sensitivity of different temperate tree species, the vulnerability of the physiological integrity of these trees remains unclear.Here, we assessed the sensitivity of six temperate tree species to severe water limitation during three consecutive growing seasons, including the exceptional 2015 central European summer drought and heat wave. Specifically, we assessed stem increment growth, sap flow, water potentials, hydraulic vulnerability, and nonstructural carbohydrate contents in leaves and branches to determine how mature temperate trees responded to this exceptional weather event and how the observed responses relate to variation in xylem embolism and carbohydrate economy.We found that the trees' predawn water potentials reached their minimum values during the 2015 summer drought and most species reduced their sap flow by up to 80%. Also, increment growth was strongly impaired with the onset of the drought in all species. Despite the strong responses in the trees' growth and water relations, all species exhibited minimum midday shoot water potentials well away from values associated with severe embolism (P50). In addition, we detected no distinct decrease in nonstructural carbohydrate contents in leaves, bark, and stems throughout the drought event.Synthesis. This study shows that mature individuals of six common central European forest tree species strongly reacted to a severe summer drought by reducing their water consumption and stopping growth. We found, however, no indications for xylem embolism or carbohydrate depletion in these trees. This suggests, that xylem embolism formation and carbohydrate reserve depletion are not routine in temperate trees during seasonal strong drought and reveals a low vulnerability of the physiological integrity of temperate trees during drought events as we describe here.

Plant height and hydraulic vulnerability to drought and cold.

Understanding how plants survive drought and cold is increasingly important as plants worldwide experience dieback with drought in moist places and grow taller with warming in cold ones. Crucial in plant climate adaptation are the diameters of water-transporting conduits. Sampling 537 species across climate zones dominated by angiosperms, we find that plant size is unambiguously the main driver of conduit diameter variation. And because taller plants have wider conduits, and wider conduits within species are more vulnerable to conduction-blocking embolisms, taller conspecifics should be more vulnerable than shorter ones, a prediction we confirm with a plantation experiment. As a result, maximum plant size should be short under drought and cold, which cause embolism, or increase if these pressures relax. That conduit diameter and embolism vulnerability are inseparably related to plant size helps explain why factors that interact with conduit diameter, such as drought or warming, are altering plant heights worldwide.

Leaf hydraulic vulnerability triggers the decline in stomatal and mesophyll conductance during drought in rice.

Understanding the physiological responses of crops to drought is important for ensuring sustained crop productivity under climate change, which is expected to exacerbate the frequency and intensity of periods of drought. Drought responses involve multiple traits, and the correlations between these traits are poorly understood. Using a variety of techniques, we estimated the changes in gas exchange, leaf hydraulic conductance, and leaf turgor in rice (Oryza sativa) in response to both short- and long-term soil drought. We performed a photosynthetic limitation analysis to quantify the contributions of each limiting factor to the resultant overall decrease in photosynthesis during drought. Biomass, leaf area, and leaf width significantly decreased during the 2-week drought treatment, but leaf mass per area and leaf vein density increased. Light-saturated photosynthetic rate declined dramatically during soil drought, mainly due to the decrease in stomatal conductance (gs) and mesophyll conductance (gm). Stomatal modeling suggested that the decline in leaf hydraulic conductance explained most of the decrease in stomatal closure during the drought treatment, and may also trigger the drought-related decrease of stomatal conductance and mesophyll conductance. The results of this study provide insight into the regulation of carbon assimilation under drought conditions.

The links between leaf hydraulic vulnerability to drought and key aspects of leaf venation and xylem anatomy among 26 Australian woody angiosperms from contrasting climates.

The structural properties of leaf venation and xylem anatomy strongly influence leaf hydraulics, including the ability of leaves to maintain hydraulic function during drought. Here we examined the strength of the links between different leaf venation traits and leaf hydraulic vulnerability to drought (expressed as P50leaf by rehydration kinetics) in a diverse group of 26 woody angiosperm species, representing a wide range of leaf vulnerabilities, from four low-nutrient sites with contrasting rainfall across eastern Australia. For each species we measured key aspects of leaf venation design, xylem anatomy and leaf morphology. We also assessed for the first time the scaling relationships between hydraulically weighted vessel wall thickness (th) and lumen breadth (bh) across vein orders and habitats. Across species, variation in P50leaf was strongly correlated with the ratio of vessel wall thickness (th) to lumen breadth (bh) [(t/b)h; an index of conduit reinforcement] at each leaf vein order. Concomitantly, the scaling relationship between th and bh was similar across vein orders, with a log-log slope less than 1 indicating greater xylem reinforcement in smaller vessels. In contrast, P50leaf was not related to th and bh individually, to major vein density (Dvmajor) or to leaf size. Principal components analysis revealed two largely orthogonal trait groupings linked to variation in leaf size and drought tolerance. Our results indicate that xylem conduit reinforcement occurs throughout leaf venation, and remains closely linked to leaf drought tolerance irrespective of leaf size.

Acclimation of branch and leaf hydraulics in adult Fagus sylvatica and Picea abies in a forest through-fall exclusion experiment.

Decreasing water availability due to climate change poses the question of whether and to what extent tree species are able to hydraulically acclimate and how hydraulic traits of stems and leaves are coordinated under drought. In a through-fall exclusion experiment, hydraulic acclimation was analyzed in a mixed forest stand of Fagus sylvatica L. and Picea abies (L.) Karst. In drought-stressed (TE, through-fall exclusion over 2 years) and control (CO) trees, hydraulic vulnerability was studied in branches as well as in leaves (F. sylvatica) and end-twigs (P. abies, entirely formed during the drought period) sampled at the same height in sun-exposed portions of the tree crown. In addition, relevant xylem anatomical traits and leaf pressure-volume relations were analyzed. The TE trees reached pre-dawn water potentials down to -1.6 MPa. In both species, water potentials at 50% loss of xylem hydraulic conductivity were ~0.4 MPa more negative in TE than in CO branches. Foliage hydraulic vulnerability (expressed as water potential at 50% loss of leaf/end-twig hydraulic conductance) and water potential at turgor loss point were also, respectively, 0.4 and 0.5 MPa lower in TE trees. Minor differences were observed in conduit mean hydraulic diameter and cell wall reinforcement. Our findings indicate significant and fast hydraulic acclimation under relatively mild drought in both tree species. Acclimation was well coordinated between branches and foliage, which might be essential for survival and productivity of mature trees under future drought periods.

Xylem Sap Surface Tension May Be Crucial for Hydraulic Safety.

The surface tension (γ) of xylem sap plays a key role in stabilizing air-water interfaces at the pits between water- and gas-filled conduits to avoid air seeding at low water potentials. We studied seasonal changes in xylem sap γ in Picea abies and Pinus mugo growing at the alpine timberline. We analyzed their vulnerability to drought-induced embolism using solutions of different γ and estimated the potential effect of seasonal changes in γ on hydraulic vulnerability. In both species, xylem sap γ showed distinct seasonal courses between about 50 and 68 mn m-1 Solutions with low γ caused higher vulnerability to drought-induced xylem embolism. The water potential at 50% loss of hydraulic conductivity in P. abies and P. mugo was -3.35 and -3.86 MPa at γ of 74 mn m-1 but -2.11 and -2.09 MPa at 45 mn m-1 This indicates up to about 1 MPa seasonal variation in 50% loss of hydraulic conductivity. The results revealed pronounced effects of changes in xylem sap γ on the hydraulic safety of trees in situ. These effects also are relevant in vulnerability analyses, where the use of standard solutions with high γ overestimates hydraulic safety. Thus, γ should be considered carefully in hydraulic studies.

A multi-species synthesis of physiological mechanisms in drought-induced tree mortality.

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere-atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.

Hydraulics link leaf shape and environmental niche in terrestrial bromeliads

AbstractTerrestrial species of the megadiverse bromeliad family display a wide variety of leaf shapes, many of which have evolved convergently in different lineages. Here, I examine the links between leaf shape, venation architecture, hydraulic function, and bioclimatic relations in two bromeliad groups displaying diverse leaf shapes, the genus Pitcairnia (Pitcairnioideae) and the terrestrial grade of the Bromelioideae subfamily. Leaf shapes with broader leaf blades, notably petiolate and lanceolate morphologies, tend to show wider vein spacing, which is associated with reduced hydraulic capacity and higher hydraulic vulnerability. In turn, these leaf shapes tend to occur in species restricted to moist, aseasonal environments, suggesting that hydraulic function is an important mediator of the relationship between leaf shape and bromeliad environmental niches. This network of trait–trait and trait–environment relationships may have been of profound important in the ecological and evolutionary diversification of the bromeliads. Similar structure–function principles are likely to apply in other tropical herbaceous monocots, which are of great ecological importance but generally neglected in plant hydraulic research.

Analysis of bridge and stream conditions of over 300 Vermont bridges damaged in Tropical Storm Irene

The 2011 Tropical Storm Irene deposited 100–200 mm of rain in Vermont with a rainfall recurrence interval for a 12-hour storm exceeding 500 years in some areas. This single hurricane-related event damaged over 300 bridges. The wide range of damage prompted a networkwide analysis of flood, scour, stream and structural conditions. A first step was the assembly of a unique data set containing information on 326 damaged bridges, 1936 undamaged bridges and the surrounding stream conditions. Descriptions of the damage appear as case studies that include pre-storm bridge and stream geomorphology conditions. The assembled and georeferenced data include rainfall, damage type and extent, estimated and actual repair costs, bridge characteristics, bridge ratings and stream geomorphic assessments from a number of sources. The analyses identified significant features of bridge vulnerability under extreme floods. The bridge age and rating assessment characteristics, such as substructure, channel and structural adequacy ratings, followed by scour, waterway adequacy and sufficiency ratings, correlated strongly to damage. The stream geomorphic features have promise to supplement future bridge rating systems and in identifying hydraulic vulnerability of bridges. Empirical fragility curves relating probability of meeting or exceeding different bridge damage levels based on channel and waterway adequacy ratings are also presented.

Visualization of xylem embolism by X‐ray microtomography: a direct test against hydraulic measurements

X-ray microtomography (microCT) is becoming a valuable noninvasive tool for advancing our understanding of plant-water relations. Laboratory-based microCT systems are becoming more affordable and provide better access than synchrotron facilities. However, some systems come at the cost of comparably lower signal quality and spatial resolution than synchrotron facilities. In this study, we evaluated laboratory-based X-ray microCT imaging as a tool to nondestructively analyse hydraulic vulnerability to drought-induced embolism in a woody plant species. We analysed the vulnerability to drought-induced embolism of benchtop-dehydrated Eucalyptus camaldulensis plants using microCT and hydraulic flow measurements on the same sample material, allowing us to directly compare the two methods. Additionally, we developed a quantitative procedure to improve microCT image analysis at limited resolution and accurately measure vessel lumens. Hydraulic measurements matched closely with microCT imaging of the current-year growth ring, with similar hydraulic conductivity and loss of conductivity due to xylem embolism. Optimized thresholding of vessel lumens during image analysis, based on a physiologically meaningful parameter (theoretical conductivity), allowed us to overcome common potential constraints of some lab-based systems. Our results indicate that estimates of vulnerability to embolism provided by microCT visualization agree well with those obtained from hydraulic measurements on the same sample material.

Genetic adaptation and phenotypic plasticity contribute to greater leaf hydraulic tolerance in response to drought in warmer climates.

The ability of plants to maintain an intact water transport system in leaves under drought conditions is intimately linked to survival and can been be seen as adaptive in shaping species climatic limits. Large differences in leaf hydraulic vulnerability to drought are known among species from contrasting climates, yet whether this trait varies among populations within a single species and, furthermore, whether it is altered by changes in growth conditions, remain unclear. We examined intraspecific variation in both leaf water transport capacity (Kleaf) and leaf hydraulic vulnerability to drought (P50leaf) among eight populations of Corymbia calophylla (R. Br.) K.D. Hill & L.A.S. Johnson (Myrtaceae) from both cool and warm climatic regions grown reciprocally under two temperature treatments representing the cool and warm edge of the species distribution. Kleaf did not vary between cool and warm-climate populations, nor was it affected by variable growth temperature. In contrast, population origin and growth temperature independently altered P50leaf. Using data pooled across growth temperatures, cool-climate populations showed significantly higher leaf hydraulic vulnerability (P50leaf = -3.55 ± 0.18 MPa) than warm-climate populations (P50leaf = -3.78 ± 0.08 MPa). Across populations, P50leaf decreased as population home-climate temperature increased, but was unrelated to rainfall and aridity. For populations from both cool and warm climatic regions, P50leaf was lower under the warmer growth conditions. These results provide evidence of trait plasticity in leaf hydraulic vulnerability to drought in response to variable growth temperature. Furthermore, they suggest that climate, and in particular temperature, may be a strong selective force in shaping intraspecific variation in leaf hydraulic vulnerability to drought.

Fitplc - an R package to fit hydraulic vulnerability curves

We describe a toolkit to fit hydraulic vulnerability curves, such as the percent loss of xylem hydraulic conductivity ('PLC curves') as a function of the water potential. The toolkit is implemented as an R package, and is thus free to use and open source. The package fits the Weibull or sigmoidal function to measurements of PLC, conductance or conductivity, at corresponding leaf or stem water potentials. From the fitted curve, estimates of Px (the water potential at which x% conductivity is lost, e.g. the P50), and slope parameter (Sx) are provided together with confidence intervals (CI) around the fitted line. The CIs are estimated with the bootstrap. We also demonstrate the advantages of using mixed-effects models in situations where multiple individuals are measured on a species, as compared to the more traditional approach of fitting curves separately and averaging the parameters. We demonstrate the use of the new package with example data on seven species measured with two different techniques.

Outside-Xylem Vulnerability, Not Xylem Embolism, Controls Leaf Hydraulic Decline during Dehydration.

Leaf hydraulic supply is crucial to maintaining open stomata for CO2 capture and plant growth. During drought-induced dehydration, the leaf hydraulic conductance (Kleaf) declines, which contributes to stomatal closure and, eventually, to leaf death. Previous studies have tended to attribute the decline of Kleaf to embolism in the leaf vein xylem. We visualized at high resolution and quantified experimentally the hydraulic vulnerability of xylem and outside-xylem pathways and modeled their respective influences on plant water transport. Evidence from all approaches indicated that the decline of Kleaf during dehydration arose first and foremost due to the vulnerability of outside-xylem tissues. In vivo x-ray microcomputed tomography of dehydrating leaves of four diverse angiosperm species showed that, at the turgor loss point, only small fractions of leaf vein xylem conduits were embolized, and substantial xylem embolism arose only under severe dehydration. Experiments on an expanded set of eight angiosperm species showed that outside-xylem hydraulic vulnerability explained 75% to 100% of Kleaf decline across the range of dehydration from mild water stress to beyond turgor loss point. Spatially explicit modeling of leaf water transport pointed to a role for reduced membrane conductivity consistent with published data for cells and tissues. Plant-scale modeling suggested that outside-xylem hydraulic vulnerability can protect the xylem from tensions that would induce embolism and disruption of water transport under mild to moderate soil and atmospheric droughts. These findings pinpoint outside-xylem tissues as a central locus for the control of leaf and plant water transport during progressive drought.