Xylem cavitation resistance has profound implications for plant physiology and ecology. This process is characterized by a ‘vulnerability curve’ (VC) showing the variation of the percentage of ...cavitation as a function of xylem pressure potential. The shape of this VC varies from ‘sigmoidal’ to ‘exponential’. This review provides a panorama of the techniques that have been used to generate such a curve. The techniques differ by (i) the way cavitation is induced (e.g. bench dehydration, centrifugation, or air injection), and (ii) the way cavitation is measured (e.g. percentage loss of conductivity (PLC) or acoustic emission), and a nomenclature is proposed based on these two methods. A survey of the literature of more than 1200 VCs was used to draw statistics on the usage of these methods and on their reliability and validity. Four methods accounted for more than 96% of all curves produced so far: bench dehydration–PLC, centrifugation–PLC, pressure sleeve-PLC, and Cavitron. How the shape of VCs varies across techniques and species xylem anatomy was also analysed. Strikingly, it was found that the vast majority of curves obtained with the reference bench dehydration-PLC method are ‘sigmoidal’. ‘Exponential’ curves were more typical of the three other methods and were remarkably frequent for species having large xylem conduits (ring-porous), leading to a substantial overestimation of the vulnerability of cavitation for this functional group. We suspect that ‘exponential’ curves may reflect an open-vessel artefact and call for more precautions with the usage of the pressure sleeve and centrifugation techniques.
Climate change is expected to exacerbate drought for many plants, making drought tolerance a key driver of species and ecosystem responses. Plant drought tolerance is determined by multiple traits, ...but the relationships among traits, either within individual plants or across species, have not been evaluated for general patterns across plant diversity. We synthesized the published data for stomatal closure, wilting, declines in hydraulic conductivity in the leaves, stems, and roots, and plant mortality for 262 woody angiosperm and 48 gymnosperm species. We evaluated the correlations among the drought tolerance traits across species, and the general sequence of water potential thresholds for these traits within individual plants. The trait correlations across species provide a framework for predicting plant responses to a wide range of water stress from one or two sampled traits, increasing the ability to rapidly characterize drought tolerance across diverse species. Analyzing these correlations also identified correlations among the leaf and stem hydraulic traits and the wilting point, or turgor loss point, beyond those expected from shared ancestry or independent associations with water stress alone. Further, on average, the angiosperm species generally exhibited a sequence of drought tolerance traits that is expected to limit severe tissue damage during drought, such as wilting and substantial stem embolism. This synthesis of the relationships among the drought tolerance traits provides crucial, empirically supported insight into representing variation in multiple traits in models of plant and ecosystem responses to drought.
Summary
Although transpiration‐driven transport of xylem sap is well known to operate under absolute negative pressure, many terrestrial, vascular plants show positive xylem pressure above ...atmospheric pressure on a seasonal or daily basis, or during early developmental stages. The actual location and mechanisms behind positive xylem pressure remain largely unknown, both in plants that show seasonal xylem pressure before leaf flushing, and those that show a diurnal periodicity of bleeding and guttation. Available evidence shows that positive xylem pressure can be driven based on purely physical forces, osmotic exudation into xylem conduits, or hydraulic pressure in parenchyma cells associated with conduits. The latter two mechanisms may not be mutually exclusive and can be understood based on a similar modelling scenario. Given the renewed interest in positive xylem pressure, this review aims to provide a constructive way forward by discussing similarities and differences of mechanistic models, evaluating available evidence for hydraulic functions, such as rehydration of tissues, refilling of water stores, and embolism repair under positive pressure, and providing recommendations for future research, including methods that avoid or minimise cutting artefacts.
Drought-induced tree mortality has been observed globally and is expected to increase under climate change scenarios, with large potential consequences for the terrestrial carbon sink. Predicting ...mortality across species is crucial for assessing the effects of climate extremes on forest community biodiversity, composition, and carbon sequestration. However, the physiological traits associated with elevated risk of mortality in diverse ecosystems remain unknown, although these traits could greatly improve understanding and prediction of tree mortality in forests. We performed a meta-analysis on species’ mortality rates across 475 species from 33 studies around the globe to assess which traits determine a species’ mortality risk. We found that species-specific mortality anomalies from community mortality rate in a given drought were associated with plant hydraulic traits. Across all species, mortality was best predicted by a low hydraulic safety margin—the difference between typical minimum xylem water potential and that causing xylem dysfunction—and xylem vulnerability to embolism. Angiosperms and gymnosperms experienced roughly equal mortality risks. Our results provide broad support for the hypothesis that hydraulic traits capture key mechanisms determining tree death and highlight that physiological traits can improve vegetation model prediction of tree mortality during climate extremes.
Pit membranes between xylem vessels have been suggested to have functional adaptive traits because of their influence on hydraulic resistance and vulnerability to embolism in plants. Observations of ...intervessel pit membranes in 26 hardwood species using electron microscopy showed significant variation in their structure, with a more than 25-fold difference in thickness (70-1892 nm) and observed maximum pore diameter (10-225 nm). In some SEM images, pit membrane porosity was affected by sample preparation, although pores were resolvable in intact pit membranes of many species. A significant relationship (r² = 0.7, P = 0.002) was found between pit membrane thickness and maximum pore diameter, indicating that the thinner membranes are usually more porous. In a subset of nine species, maximum pore diameter determined from SEM was correlated with pore diameter calculated from air-seeding thresholds (r² = 0.8, P < 0.001). Our data suggest that SEM images of intact pit membranes underestimate the porosity of pit membranes in situ. Pit membrane porosity based on SEM offers a relative estimate of air-seeding thresholds, but absolute pore diameters must be treated with caution. The implications of variation in pit membrane thickness and porosity to plant function are discussed.
Wood performs several essential functions in plants, including mechanically supporting aboveground tissue, storing water and other resources, and transporting sap. Woody tissues are likely to face ...physiological, structural and defensive trade-offs. How a plant optimizes among these competing functions can have major ecological implications, which have been under-appreciated by ecologists compared to the focus they have given to leaf function. To draw together our current understanding of wood function, we identify and collate data on the major wood functional traits, including the largest wood density database to date (8412 taxa), mechanical strength measures and anatomical features, as well as clade-specific features such as secondary chemistry. We then show how wood traits are related to one another, highlighting functional trade-offs, and to ecological and demographic plant features (growth form, growth rate, latitude, ecological setting). We suggest that, similar to the manifold that tree species leaf traits cluster around the 'leaf economics spectrum', a similar 'wood economics spectrum' may be defined. We then discuss the biogeography, evolution and biogeochemistry of the spectrum, and conclude by pointing out the major gaps in our current knowledge of wood functional traits.
Bordered pits are cavities in the lignified cell walls of xylem conduits (vessels and tracheids) that are essential components in the water-transport system of higher plants. The pit membrane, which ...lies in the center of each pit, allows water to pass between xylem conduits but limits the spread of embolism and vascular pathogens in the xylem. Averaged across a wide range of species, pits account for > 50% of total xylem hydraulic resistance, indicating that they are an important factor in the overall hydraulic efficiency of plants. The structure of pits varies dramatically across species, with large differences evident in the porosity and thickness of pit membranes. Because greater porosity reduces hydraulic resistance but increases vulnerability to embolism, differences in pit structure are expected to correlate with trade-offs between efficiency and safety of water transport. However, trade-offs in hydraulic function are influenced both by pit-level differences in structure (e.g. average porosity of pit membranes) and by tissue-level changes in conduit allometry (average length, diameter) and the total surface area of pit membranes that connects vessels. In this review we address the impact of variation in pit structure on water transport in plants from the level of individual pits to the whole plant.
PREMISE OF THE STUDY: Concentrations of nonstructural carbohydrates (NSCs) are used as proxies for the net carbon balance of trees and as indicators of carbon starvation resulting from environmental ...stress. Woody organs are the largest NSC-storing compartments in forest ecosystems; therefore, it is essential to understandthe factors that affect the size of this important storage pool. In wood, NSC are predominantly deposited in ray and axial parenchyma (RAP); however, direct links between nutrient storage and RAP anatomy have not yet been established. Here, we tested whether the NSC storage capacity of wood is influenced by the amount of RAP. METHODS: We measured NSC concentrations and RAP fractions in root and stem sapwood of 12 temperate species sampled at the onset of winter dormancy and in stem sapwood of four tropical trees growing in an evergreen lowland rainforest. The patterns of starch distribution were visualized by staining with Lugol's solution. KEY RESULTS: The concentration of NSCs in sapwood of temperate trees scales tightly with the amount of RAP and living fibers (LFs), with almost all RAP and LFs being densely packed with starch grains. In contrast, the tropical species had lower NSC concentrations despite their higher RAP and LFs fraction and had considerable interspecific differences in starch distribution. CONCLUSIONS: The differences in RAP and LFs abundance affect the ability of sapwood to store NSC in temperate trees, whereas a more diverse set of functions of RAP might be pronounced in species growing in a tropical environment with little seasonally.
Hydraulic failure induced by xylem embolism is one of the primary mechanisms of plant dieback during drought. However, many of the methods used to evaluate the vulnerability of different species to ...drought-induced embolism are indirect and invasive, increasing the possibility that measurement artifacts may occur. Here, we utilize x-ray computed microtomography (microCT) to directly visualize embolism formation in the xylem of living, intact plants with contrasting wood anatomy (Quercus robur, Populus tremula × Populus alba, and Pinus pinaster). These observations were compared with widely used centrifuge techniques that require destructive sampling. MicroCT imaging provided detailed spatial information regarding the dimensions and functional status of xylem conduits during dehydration. Vulnerability curves based on microCT observations of intact plants closely matched curves based on the centrifuge technique for species with short vessels (P. tremula × P. alba) or tracheids (P. pinaster). For ring porous Q. robur, the centrifuge technique significantly overestimated vulnerability to embolism, indicating that caution should be used when applying this technique to species with long vessels. These findings confirm that microCT can be used to assess the vulnerability to embolism on intact plants by direct visualization.
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