Increasing leaf hydraulic conductance with transpiration rate minimizes the water potential drawdown from stem to leaf.

Leaf hydraulic conductance (k leaf) is a central element in the regulation of leaf water balance but the properties of k leaf remain uncertain. Here, the evidence for the following two models for k leaf in well-hydrated plants is evaluated: (i) k leaf is constant or (ii) k leaf increases as transpiration rate (E) increases. The difference between stem and leaf water potential (ΔΨstem-leaf), stomatal conductance (g s), k leaf, and E over a diurnal cycle for three angiosperm and gymnosperm tree species growing in a common garden, and for Helianthus annuus plants grown under sub-ambient, ambient, and elevated atmospheric CO₂ concentration were evaluated. Results show that for well-watered plants k leaf is positively dependent on E. Here, this property is termed the dynamic conductance, k leaf(E), which incorporates the inherent k leaf at zero E, which is distinguished as the static conductance, k leaf(0). Growth under different CO₂ concentrations maintained the same relationship between k leaf and E, resulting in similar k leaf(0), while operating along different regions of the curve owing to the influence of CO₂ on g s. The positive relationship between k leaf and E minimized variation in ΔΨstem-leaf. This enables leaves to minimize variation in Ψleaf and maximize g s and CO₂ assimilation rate over the diurnal course of evaporative demand

Variation in xylem hydraulic structure and function of two mangrove species across a latitudinal gradient in eastern Australia

Mangroves growing in tropical and subtropical intertidal zones face challenges from warming and altered rainfall patterns associated with global climate change. Intraspecific variation in hydraulic traits may allow a mangrove species to acclimate to novel climatic conditions, yet little is known regarding the potential for adaptive plasticity in these traits. In this study, we aimed to quantify the variation in plant hydraulic traits of two widespread mangrove species growing across a latitudinal gradient. We investigated the xylem hydraulic structure and function of Avicennia marina and Aegiceras corniculatum, across three sites spanning a latitudinal gradient of 17.45◦ in eastern Australia. We found that both species were highly resistant to xylem embolism and that there was significant intraspecific variation in hydraulic traits between sites. The highest embolism resistance and sapwood-specific hydraulic conductivity (KS) were found at the lowest latitude site that had the highest mean annual temperature and precipitation. A. marina showed no differences in vessel size and density among sites. It has other special features such as successive cambia enhancing its ability to adapt to a large environmental gradient. In contrast, A. corniculatum showed higher vessel densities at lower latitudes. There was a significant and positive correlation (R 2 = 0.72, p < 0.05) between KS and embolism resistance across species and sites, suggesting the absence of a tradeoff between hydraulic efficiency and safety. Both embolism resistance and KS were negatively correlated with wood density but positively with vessel wall reinforcement. This study reveals that these two widespread mangrove species were adapted to warmer climates by enhancing both hydraulic efficiency and safety

Unlocking drought-induced tree mortality : physiological mechanisms to modeling

Drought-related tree mortality has become a major concern worldwide due to its pronounced negative impacts on the functioning and sustainability of forest ecosystems. However, our ability to identify the species that are most vulnerable to drought, and to pinpoint the spatial and temporal patterns of mortality events, is still limited. Model is useful tools to capture the dynamics of vegetation at spatiotemporal scales, yet contemporary land surface models (LSMs) are often incapable of predicting the response of vegetation to environmental perturbations with sufficient accuracy, especially under stressful conditions such as drought. Significant progress has been made regarding the physiological mechanisms underpinning plant drought response in the past decade, and plant hydraulic dysfunction has emerged as a key determinant for tree death due to water shortage. The identification of pivotal physiological events and relevant plant traits may facilitate forecasting tree mortality through a mechanistic approach, with improved precision. In this review, we (1) summarize current understanding of physiological mechanisms leading to tree death, (2) describe the functionality of key hydraulic traits that are involved in the process of hydraulic dysfunction, and (3) outline their roles in improving the representation of hydraulic function in LSMs. We urge potential future research on detailed hydraulic processes under drought, pinpointing corresponding functional traits, as well as understanding traits variation across and within species, for a better representation of drought-induced tree mortality in models

Non-invasive imaging reveals convergence in root and stem vulnerability to cavitation across five tree species

Root vulnerability to cavitation is challenging to measure and under-represented in current datasets. This gap limits the precision of models used to predict plant responses to drought because roots comprise the critical interface between plant and soil. In this study we measured vulnerability to drought induced cavitation in woody roots and stems of five tree species (Acacia aneura, Cedrus deodara, Eucalyptus crebra, Eucalytus saligna, and Quercus palustris) with a wide range of xylem anatomies. X-ray microtomography was used to visualize the accumulation of xylem embolism in stems and roots of intact plants that were naturally dehydrated to varying levels of water stress. Vulnerability to cavitation, defined as the water potential causing a 50% loss of hydraulic function (P50), varied broadly among the species (-4.51 to -11.93 MPa in stems and -3.13 to -9.64 MPa in roots). The P50 of roots and stems was significantly related across species, with species that had more vulnerable stems also having more vulnerable roots. While there was strong convergence in root and stem vulnerability to cavitation, the P50 of roots was significantly higher than the P50 of stems in three species. However, the difference in root and stem vulnerability for these species was small; between 1% to 31% of stem P50. Thus, while some differences existed between organs, roots were not dramatically more vulnerable to embolism than stems and the differences observed were less than those reported in previous studies. Further study is required to evaluate the vulnerability across root orders and to extend these conclusions to a greater number of species and xylem functional types

An empirical method that separates irreversible stem radial growth from bark water content changes in trees : theory and case studies

Substantial uncertainty surrounds our knowledge of tree stem growth, with some of the most basic questions, such as when stem radial growth occurs through the daily cycle, still unanswered. We employed high-resolution point dendrometers, sap flow sensors, and developed theory and statistical approaches, to devise a novel method separating irreversible radial growth from elastic tension-driven and elastic osmotically driven changes in bark water content. We tested this method using data from five case study species. Experimental manipulations, namely a field irrigation experiment on Scots pine and a stem girdling experiment on red forest gum trees, were used to validate the theory. Time courses of stem radial growth following irrigation and stem girdling were consistent with a-priori predictions. Patterns of stem radial growth varied across case studies, with growth occurring during the day and/or night, consistent with the available literature. Importantly, our approach provides a valuable alternative to existing methods, as it can be approximated by a simple empirical interpolation routine that derives irreversible radial growth using standard regression techniques. Our novel method provides an improved understanding of the relative source-sink carbon dynamics of tree stems at a sub-daily time scale.Peer reviewe

Using dense Sentinel-2 time series to explore combined fire and drought impacts in eucalypt forests

Following one of the driest years on record, millions of hectares of forests in southeast Australia were burned in the 2019-20200 "Black Summer" wildfires. In addition to the areas burned, drought related canopy collapse, dieback and tree mortality was widely observed. In this paper, we present a method to map canopy damage due to drought and fire across a large area. Sentinel-2 satellite imagery was used in a monthly time series to highlight areas of forest where the Normalized Burn Ratio index was significantly below a pre-disturbance "stable" period. The stable period was defined as the 3 years prior to 2019 and the disturbance thresholds are based on bioregion specific standard deviations below pre-disturbance means. The novel methods enabled drought impacted forests to be identified, including those which were subsequently burned by wildfire. Across the 20 Mha of forests studied, 9.9 Mha (49%) fell below the disturbance threshold. Of that, 5.8 Mha was disturbed by fire and a further 4.1 Mha by drought outside of the fire extent. Within the fire extent, almost 0.9 Mha was identified as being significantly drought affected prior to being burned. An analysis of spectral recovery following substantial rainfall from February 2020 onward indicates that most of the areas impacted by both drought and fire have similar rates of recovery to those impacted only by fire. There are some areas, however, where the combined effects of the "double disturbance "appears to be hindering recovery. The methods presented here are easily transferrable and demonstrate an approach for monitoring forest disturbance at higher temporal and spatial scales than those typically used

The role of hydraulic failure in a massive mangrove die-off event

Between late 2015 and early 2016, more than 7,000 ha of mangrove forest died along the coastline of the Gulf of Carpentaria, in northern Australia. This massive die-off was preceded by a strong 2015/2016 El Niño event, resulting in lower precipitation, a drop in sea level and higher than average temperatures in northern Australia. In this study, we investigated the role of hydraulic failure in the mortality and recovery of the dominant species, Avicennia marina, 2 years after the mortality event. We measured predawn water potential (Ψpd) and percent loss of stem hydraulic conductivity (PLC) in surviving individuals across a gradient of impact. We also assessed the vulnerability to drought-induced embolism (Ψ50) for the species. Areas with severe canopy dieback had higher native PLC (39%) than minimally impacted areas (6%), suggesting that hydraulic recovery was ongoing. The high resistance of A. marina to water-stress-induced embolism (Ψ50 = −9.6 MPa), indicates that severe water stress (Ψpd < −10 MPa) would have been required to cause mortality in this species. Our data indicate that the natural gradient of water-stress enhanced the impact of El Niño, leading to hydraulic failure and mortality in A. marina growing on severely impacted (SI) zones. It is likely that lowered sea levels and less frequent inundation by seawater, combined with lower inputs of fresh water, high evaporative demand and high temperatures, led to the development of hyper-salinity and extreme water stress during the 2015/16 summer

Towards species-level forecasts of drought-induced tree mortality risk

Predicting species-level responses to drought at the landscape scale is critical to reducing uncertainty in future terrestrial carbon and water cycle projections. We embedded a stomatal optimisation model in the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model and parameterised the model for 15 canopy dominant eucalypt tree species across South-Eastern Australia (mean annual precipitation range: 344–1424 mm yr−1). We conducted three experiments: applying CABLE to the 2017–2019 drought; a 20% drier drought; and a 20% drier drought with a doubling of atmospheric carbon dioxide (CO2). The severity of the drought was highlighted as for at least 25% of their distribution ranges, 60% of species experienced leaf water potentials beyond the water potential at which 50% of hydraulic conductivity is lost due to embolism. We identified areas of severe hydraulic stress within-species’ ranges, but we also pinpointed resilience in species found in predominantly semiarid areas. The importance of the role of CO2 in ameliorating drought stress was consistent across species. Our results represent an important advance in our capacity to forecast the resilience of individual tree species, providing an evidence base for decision-making around the resilience of restoration plantings or net-zero emission strategies

Towards species‐level forecasts of drought‐induced tree mortality risk

Predicting species-level responses to drought at the landscape scale is critical to reducing uncertainty in future terrestrial carbon and water cycle projections. We embedded a stomatal optimisation model in the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model and parameterised the model for 15 canopy dominant eucalypt tree species across South-Eastern Australia (mean annual precipitation range: 344–1424 mm yr−1). We conducted three experiments: applying CABLE to the 2017–2019 drought; a 20% drier drought; and a 20% drier drought with a doubling of atmospheric carbon dioxide (CO2). The severity of the drought was highlighted as for at least 25% of their distribution ranges, 60% of species experienced leaf water potentials beyond the water potential at which 50% of hydraulic conductivity is lost due to embolism. We identified areas of severe hydraulic stress within-species’ ranges, but we also pinpointed resilience in species found in predominantly semiarid areas. The importance of the role of CO2 in ameliorating drought stress was consistent across species. Our results represent an important advance in our capacity to forecast the resilience of individual tree species, providing an evidence base for decision-making around the resilience of restoration plantings or net-zero emission strategies

One Stomatal Model to Rule Them All?:Toward Improved Representation of Carbon and Water Exchange in Global Models

Stomatal conductance schemes that optimize with respect to photosynthetic and hydraulic functions have been proposed to address biases in land-surface model (LSM) simulations during drought. However, systematic evaluations of both optimality-based and alternative empirical formulations for coupling carbon and water fluxes are lacking. Here, we embed 12 empirical and optimization approaches within a LSM framework. We use theoretical model experiments to explore parameter identifiability and understand how model behaviors differ in response to abiotic changes. We also evaluate the models against leaf-level observations of gas-exchange and hydraulic variables, from xeric to wet forest/woody species spanning a mean annual precipitation range of 361–3,286 mm yr−1. We find that models differ in how easily parameterized they are, due to: (a) poorly constrained optimality criteria (i.e., resulting in multiple solutions), (b) low influence parameters, (c) sensitivities to environmental drivers. In both the idealized experiments and compared to observations, sensitivities to variability in environmental drivers do not agree among models. Marked differences arise in sensitivities to soil moisture (soil water potential) and vapor pressure deficit. For example, stomatal closure rates at high vapor pressure deficit range between −45% and +70% of those observed. Although over half the new generation of stomatal schemes perform to a similar standard compared to observations of leaf-gas exchange, two models do so through large biases in simulated leaf water potential (up to 11 MPa). Our results provide guidance for LSM development, by highlighting key areas in need for additional experimentation and theory, and by constraining currently viable stomatal hypotheses