Conduit position and connectivity affect the likelihood of xylem embolism during natural drought in evergreen woodland species.

BACKGROUND AND AIMS: Hydraulic failure is considered a main cause of drought-induced forest mortality. Yet, we have a limited understanding of how the varying intensities and long time scales of natural droughts induce and propagate embolism within the xylem. METHODS: X-ray computed tomography (microCT) images were obtained from different aged branch xylem to study the number, size and spatial distribution of in situ embolized conduits among three dominant tree species growing in a woodland community. KEY RESULTS: Among the three studied tree species, those with a higher xylem vulnerability to embolism (higher water potential at 50 % loss of hydraulic conductance; P50) were more embolized than species with lower P50. Within individual stems, the probability of embolism was independent of conduit diameter but associated with conduit position. Rather than the occurrence of random or radial embolism, we observed circumferential clustering of high and low embolism density, suggesting that embolism spreads preferentially among conduits of the same age. Older xylem also appeared more likely to accumulate embolisms than young xylem, but there was no pattern suggesting that branch tips were more vulnerable to cavitation than basal regions. CONCLUSIONS: The spatial analysis of embolism occurrence in field-grown trees suggests that embolism under natural drought probably propagates by air spreading from embolized into neighbouring conduits in a circumferential pattern. This pattern offers the possibility to understand the temporal aspects of embolism occurrence by examining stem cross-sections.

Flooding constrains tree water use of a riparian forest in the lower Heihe River Basin, Northwest China.

Riparian forests in floodplains are occasionally or regularly submerged by flooding. However, controversy exists regarding the effects of flooding on water use in riparian forests, and this controversy severely restricts our ability to better utilize limited water resources to restore damaged riparian forests in arid regions.The evapotranspiration (Et) and transpiration (T) of riparian P. euphratica forests in the arid regions of northwestern China were determined using eddy covariance and sap flow technology across a 3-year period. Fortunately, the flooding introduced by ecological water diversion was occurred in 2014 and 2016 but not in 2015. Our results showed that the magnitude and seasonal pattern of Et across 3 years was comparable (approximately 900 mm), but the T was higher in 2015 (431 mm) than in the other two years (288 mm in 2014 and 290 mm in 2016). The interannual patterns in the transpiration were consistent with the net ecosystem productivity at the site. Given the similar meteorological conditions (e.g. net radiation, temperature, relative humidity, and vapor pressure deficit) among the 3 years, two aspects may contributed to the suppressed tree water use and productivity under flooding: 1) the increased soil salinity reduce the roots water uptake from soil by increasing root water potential via osmotic adjustment; and 2) the depressed tree growth (e.g. the leaf area) via suspended water upward transport along soil-plant-atmosphere continuum. Although flooding is widely known beneficial for the regeneration, we suggest that it is not appropriate for the rejuvenation of phreatophyte (e.g., Populus spp.) in arid regions. Our results were drawn from only three years of measurement and therefore longer time series are needed to confirm or refine those conclusions.

Risks to carbon dynamics in semi-arid woodlands of eastern Australia under current and future climates.

Extreme disturbance events, such as wildfire and drought, have large impacts on carbon storage and sequestration of forests and woodlands globally. Here, we present a modelling approach that assesses the relative impact of disturbances on carbon storage and sequestration, and how this will alter under climate change. Our case study is semi-arid Australia where large areas of land are managed to offset over 122 million tonnes of anthropogenic carbon emissions over a 100-year period. These carbon offsets include mature vegetation that has been protected from clearing and regenerating vegetation on degraded agricultural land. We use a Bayesian Network model to combine multiple probabilistic models of the risk posed by fire, drought, grazing and recruitment failure to carbon dynamics. The model is parameterised from a review of relevant literature and additional quantitative analyses presented here. We found that the risk of vegetation becoming a net source of carbon due to a mortality event, or failing to realise maximum sequestration potential, through recruitment failure in regenerating vegetation, was primarily a function of rainfall in this semi-arid environment. However, the relative size of an emissions event varied across vegetation communities depending on plant attributes, specifically resprouting capacity. Modelled climate change effects were variable, depending on the climate change projection used. Under 'best-case' or 'most-likely' climate scenarios for 2050, similar or increased projections of mean annual precipitation, associated with a build-up of fuel, were expected to drive an increase in fire activity (a 40-160% increase), but a decrease in drought (a 20-35% decrease). Under a 'worst-case' climate scenario, fire activity was expected to decline (a 37% decrease), but drought conditions remain similar (a 5% decrease). These projected changes to the frequency of drought and fire increase the risk that vegetation used for carbon offsetting will fail to provide anticipated amounts of carbon abatement over their lifetime.

Intraspecific variation in drought susceptibility in Eucalyptus globulus is linked to differences in leaf vulnerability.

Understanding intraspecific variation in the vulnerability of the xylem to hydraulic failure during drought is critical in predicting the response of forest tree species to climate change. However, few studies have assessed intraspecific variation in this trait, and a likely limitation is the large number of measurements required to generate the standard 'vulnerability curve' used to assess hydraulic failure. Here we explore an alternative approach that requires fewer measurements, and assess within species variation in leaf xylem vulnerability in Eucalyptus globulus Labill., an ecologically and economically important species with known genetic variation in drought tolerance. Using this approach we demonstrate significant phenotypic differences and evidence of plasticity among two provenances with contrasting drought tolerance.

Subcontinental heat wave triggers terrestrial and marine, multi-taxa responses.

Heat waves have profoundly impacted biota globally over the past decade, especially where their ecological impacts are rapid, diverse, and broad-scale. Although usually considered in isolation for either terrestrial or marine ecosystems, heat waves can straddle ecosystems of both types at subcontinental scales, potentially impacting larger areas and taxonomic breadth than previously envisioned. Using climatic and multi-species demographic data collected in Western Australia, we show that a massive heat wave event straddling terrestrial and maritime ecosystems triggered abrupt, synchronous, and multi-trophic ecological disruptions, including mortality, demographic shifts and altered species distributions. Tree die-off and coral bleaching occurred concurrently in response to the heat wave, and were accompanied by terrestrial plant mortality, seagrass and kelp loss, population crash of an endangered terrestrial bird species, plummeting breeding success in marine penguins, and outbreaks of terrestrial wood-boring insects. These multiple taxa and trophic-level impacts spanned >300,000 km2-comparable to the size of California-encompassing one terrestrial Global Biodiversity Hotspot and two marine World Heritage Areas. The subcontinental multi-taxa context documented here reveals that terrestrial and marine biotic responses to heat waves do not occur in isolation, implying that the extent of ecological vulnerability to projected increases in heat waves is underestimated.

Depressed hydraulic redistribution of roots more by stem refilling than by nocturnal transpiration for Populus euphratica Oliv. in situ measurement.

During the night, plant water loss can occur either through the roots, as hydraulic redistribution (HR), or through the leaves via the stoma, as nocturnal transpiration (En), which was methodologically difficult to separate from stem refilling (Re). While HR and En have been reported across a range of species, ecosystem, and climate zone, there is little understanding on the interactions between En and/or Re and HR. As water movement at night occurs via gradients of water potential, it is expected that during periods of high atmospheric vapor pressure deficit (VPD), water loss via En will override water loss via HR. To test this hypothesis, sap flow in stems and roots of Populus euphratica Oliv. trees, growing in a riparian zone in a hyperarid climate, was measured once in a year. Nocturnal stem sap flow was separated into En and Re using the "forecasted refilling" method. Substantial nocturnal sap flow (38% of 24-hr flux on average) was observed and positively correlated with VPD; however, the strength of the correlation was lower (R2 = .55) than diurnal sap flow (Ed) (R2 = .72), suggesting that nocturnal stem sap flow was attributed to both water loss through the canopy and replenishment of water in stem tissues. Partitioning of nocturnal sap flow shows that Re constituted approximately 80%, and En ~20%, of nocturnal sap flow. The amount of root sap flow attributed to redistribution was negatively related to Ed (R2 = .69) and the amount of acropetally sap flow in stems, Re (R2 = .41) and En (R2 = .14). It was suggested that the magnitude of HR is more strongly depressed by Re that was recharge to the water loss via Ed than by En. It was consistent with whole-tree water balance theory, that the nighttime upward sap flow to xylem, stem refilling and transpiration, may depress hydraulic redistribution of roots.

Large Partial-Thickness Tear of the Subscapularis Tendon.

A 27-year-old active-duty male sailor directly accessed physical therapy for deep left anterior shoulder pain. The patient sustained his injury the previous day, when he threw a left hook into the body of his sparring partner, creating a sudden high-energy external rotation force. Based on the patient's complaint, mechanism of injury, and internal rotation weakness, the examining physical therapist ordered magnetic resonance imaging. The images revealed a large partial tear, with retraction of the subscapularis tendon at the lesser tuberosity, a labral tear, and an intratendon biceps tendon tear with subluxation. J Orthop Sports Phys Ther 2018;48(12):983. doi:10.2519/jospt.2018.8221.

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.

Gas exchange recovery following natural drought is rapid unless limited by loss of leaf hydraulic conductance: evidence from an evergreen woodland.

Drought can cause major damage to plant communities, but species damage thresholds and postdrought recovery of forest productivity are not yet predictable. We used an El Niño drought event as a natural experiment to test whether postdrought recovery of gas exchange could be predicted by properties of the water transport system, or if metabolism, primarily high abscisic acid concentration, might delay recovery. We monitored detailed physiological responses, including shoot sapflow, leaf gas exchange, leaf water potential and foliar abscisic acid (ABA), during drought and through the subsequent rehydration period for a sample of eight canopy and understory species. Severe drought caused major declines in leaf water potential, elevated foliar ABA concentrations and reduced stomatal conductance and assimilation rates in our eight sample species. Leaf water potential surpassed levels associated with incipient loss of leaf hydraulic conductance in four species. Following heavy rainfall gas exchange in all species, except those trees predicted to have suffered hydraulic impairment, recovered to prestressed rates within 1 d. Recovery of plant gas exchange was rapid and could be predicted by the hydraulic safety margin, providing strong support for leaf vulnerability to water deficit as an index of damage under natural drought conditions.

Significant contribution from foliage-derived ABA in regulating gas exchange in Pinus radiata.

The complex regulatory system controlling stomata involves physical and chemical signals that affect guard cell turgor to bring about changes in stomatal conductance (gs). Abscisic acid (ABA) closes stomata, yet the mechanisms controlling foliar ABA status in tree species remain unclear. The importance of foliage-derived ABA in regulating gas exchange was evaluated under treatments that affected phloem export through girdling and reduced water availability in the tree species, Pinus radiata (D. Don). Branch- and whole-plant girdling increased foliar ABA levels leading to declines in gs, despite no change in plant water status. Changes in gs were largely independent of the more transient increases in foliar non-structural carbohydrates (NSC), suggesting that gradual accumulation of foliar ABA was the primary mechanism for reductions in gs and assimilation. Whole-plant girdling eventually reduced root NSC, hindering root water uptake and decreasing foliar water potential, causing a dramatic increase in ABA level in leaves and concentrations in the xylem sap of shoots (4032 ng ml-1), while root xylem sap concentrations remained low (43 ng ml-1). Contrastingly, the drought treatment caused similar increases in xylem sap ABA in both roots and shoots, suggesting that declines in water potential result in relatively consistent changes in ABA along the hydraulic pathway. ABA levels in plant canopies can be regulated independently of changes in root water status triggered by changes by both phloem export and foliar water status.

An ecoclimatic framework for evaluating the resilience of vegetation to water deficit.

The surge in global efforts to understand the causes and consequences of drought on forest ecosystems has tended to focus on specific impacts such as mortality. We propose an ecoclimatic framework that takes a broader view of the ecological relevance of water deficits, linking elements of exposure and resilience to cumulative impacts on a range of ecosystem processes. This ecoclimatic framework is underpinned by two hypotheses: (i) exposure to water deficit can be represented probabilistically and used to estimate exposure thresholds across different vegetation types or ecosystems; and (ii) the cumulative impact of a series of water deficit events is defined by attributes governing the resistance and recovery of the affected processes. We present case studies comprising Pinus edulis and Eucalyptus globulus, tree species with contrasting ecological strategies, which demonstrate how links between exposure and resilience can be examined within our proposed framework. These examples reveal how climatic thresholds can be defined along a continuum of vegetation functional responses to water deficit regimes. The strength of this framework lies in identifying climatic thresholds on vegetation function in the absence of more complete mechanistic understanding, thereby guiding the formulation, application and benchmarking of more detailed modelling.

Weak tradeoff between xylem safety and xylem-specific hydraulic efficiency across the world's woody plant species.

The evolution of lignified xylem allowed for the efficient transport of water under tension, but also exposed the vascular network to the risk of gas emboli and the spread of gas between xylem conduits, thus impeding sap transport to the leaves. A well-known hypothesis proposes that the safety of xylem (its ability to resist embolism formation and spread) should trade off against xylem efficiency (its capacity to transport water). We tested this safety-efficiency hypothesis in branch xylem across 335 angiosperm and 89 gymnosperm species. Safety was considered at three levels: the xylem water potentials where 12%, 50% and 88% of maximal conductivity are lost. Although correlations between safety and efficiency were weak (r(2)  < 0.086), no species had high efficiency and high safety, supporting the idea for a safety-efficiency tradeoff. However, many species had low efficiency and low safety. Species with low efficiency and low safety were weakly associated (r(2)  < 0.02 in most cases) with higher wood density, lower leaf- to sapwood-area and shorter stature. There appears to be no persuasive explanation for the considerable number of species with both low efficiency and low safety. These species represent a real challenge for understanding the evolution of xylem.

Exposure of trees to drought-induced die-off is defined by a common climatic threshold across different vegetation types.

Increases in drought and temperature stress in forest and woodland ecosystems are thought to be responsible for the rise in episodic mortality events observed globally. However, key climatic drivers common to mortality events and the impacts of future extreme droughts on tree survival have not been evaluated. Here, we characterize climatic drivers associated with documented tree die-off events across Australia using standardized climatic indices to represent the key dimensions of drought stress for a range of vegetation types. We identify a common probabilistic threshold associated with an increased risk of die-off across all the sites that we examined. We show that observed die-off events occur when water deficits and maximum temperatures are high and exist outside 98% of the observed range in drought intensity; this threshold was evident at all sites regardless of vegetation type and climate. The observed die-off events also coincided with at least one heat wave (three consecutive days above the 90th percentile for maximum temperature), emphasizing a pivotal role of heat stress in amplifying tree die-off and mortality processes. The joint drought intensity and maximum temperature distributions were modeled for each site to describe the co-occurrence of both hot and dry conditions and evaluate future shifts in climatic thresholds associated with the die-off events. Under a relatively dry and moderate warming scenario, the frequency of droughts capable of inducing significant tree die-off across Australia could increase from 1 in 24 years to 1 in 15 years by 2050, accompanied by a doubling in the occurrence of associated heat waves. By defining commonalities in drought conditions capable of inducing tree die-off, we show a strong interactive effect of water and high temperature stress and provide a consistent approach for assessing changes in the exposure of ecosystems to extreme drought events.

Structural adjustments in resprouting trees drive differences in post-fire transpiration.

Following disturbance many woody species are capable of resprouting new foliage, resulting in a reduced leaf-to-sapwood area ratio and altered canopy structure. We hypothesized that such changes would promote adjustments in leaf physiology, resulting in higher rates of transpiration per unit leaf area, consistent with the mechanistic framework proposed by Whitehead et al. (Whitehead D, Jarvis PG, Waring RH (1984) Stomatal conductance, transpiration and resistance to water uptake in a Pinus sylvestris spacing experiment. Can J For Res 14:692-700). We tested this in Eucalyptus obliqua L'Hér following a wildfire by comparing trees with unburnt canopies with trees that had been subject to 100% canopy scorch and were recovering their leaf area via resprouting. In resprouting trees, foliage was distributed along the trunk and on lateral branches, resulting in shorter hydraulic path lengths. We evaluated measurements of whole-tree transpiration and structural and physiological traits expected to drive any changes in transpiration. We used these structural and physiological measurements to parameterize the Whitehead et al. equation, and found that the expected ratio of transpiration per unit leaf area between resprouting and unburnt trees was 3.41. This is similar to the observed ratio of transpiration per unit leaf area, measured from sapflow observations, which was 2.89 (i.e., resprouting trees had 188% higher transpiration per unit leaf area). Foliage at low heights (<2 m) was found to be significantly different to foliage in the tree crown (14-18 m) in a number of traits, including higher specific leaf area, midday leaf water potential and higher rates of stomatal conductance and photosynthesis. We conclude that these post-fire adjustments in resprouting trees help to drive increased stomatal conductance and hydraulic efficiency, promoting the rapid return of tree-scale transpiration towards pre-disturbance levels. These transient patterns in canopy transpiration have important implications for modelling stand-level water fluxes in forests capable of resprouting, which is frequently done on the basis of the leaf area index.

Drought response strategies define the relative contributions of hydraulic dysfunction and carbohydrate depletion during tree mortality.

Plant survival during drought requires adequate hydration in living tissues and carbohydrate reserves for maintenance and recovery. We hypothesized that tree growth and hydraulic strategy determines the intensity and duration of the 'physiological drought', thereby affecting the relative contributions of loss of hydraulic function and carbohydrate depletion during mortality. We compared patterns in growth rate, water relations, gas exchange and carbohydrate dynamics in three tree species subjected to prolonged drought. Two Eucalyptus species (E. globulus, E. smithii) exhibited high growth rates and water-use resulting in rapid declines in water status and hydraulic conductance. In contrast, conservative growth and water relations in Pinus radiata resulted in longer periods of negative carbon balance and significant depletion of stored carbohydrates in all organs. The ongoing demand for carbohydrates from sustained respiration highlighted the role that duration of drought plays in facilitating carbohydrate consumption. Two drought strategies were revealed, differentiated by plant regulation of water status: plants maximized gas exchange, but were exposed to low water potentials and rapid hydraulic dysfunction; and tight regulation of gas exchange at the cost of carbohydrate depletion. These findings provide evidence for a relationship between hydraulic regulation of water status and carbohydrate depletion during terminal drought.

Global convergence in the vulnerability of forests to drought.

Shifts in rainfall patterns and increasing temperatures associated with climate change are likely to cause widespread forest decline in regions where droughts are predicted to increase in duration and severity. One primary cause of productivity loss and plant mortality during drought is hydraulic failure. Drought stress creates trapped gas emboli in the water transport system, which reduces the ability of plants to supply water to leaves for photosynthetic gas exchange and can ultimately result in desiccation and mortality. At present we lack a clear picture of how thresholds to hydraulic failure vary across a broad range of species and environments, despite many individual experiments. Here we draw together published and unpublished data on the vulnerability of the transport system to drought-induced embolism for a large number of woody species, with a view to examining the likely consequences of climate change for forest biomes. We show that 70% of 226 forest species from 81 sites worldwide operate with narrow (<1 megapascal) hydraulic safety margins against injurious levels of drought stress and therefore potentially face long-term reductions in productivity and survival if temperature and aridity increase as predicted for many regions across the globe. Safety margins are largely independent of mean annual precipitation, showing that there is global convergence in the vulnerability of forests to drought, with all forest biomes equally vulnerable to hydraulic failure regardless of their current rainfall environment. These findings provide insight into why drought-induced forest decline is occurring not only in arid regions but also in wet forests not normally considered at drought risk.

Using multiple trait associations to define hydraulic functional types in plant communities of south-western Australia.

Assessing the hydrological imbalance and associated land degradation issues facing much of southern Australia and other parts of the world requires a better understanding of the defining features of ecosystem water use and the design of sustainable agroecosystems. Thus, by grouping species with similar water-use strategies into 'hydraulic functional types' (HFTs), we investigated the characteristics of water use in species-rich plant communities of south-western Australia. HFTs were determined using multiple-trait associations between morphological and physiological traits relating to water transport, water-use efficiency and response to water deficit. Sixteen traits were assessed from a subset of 21 species from three plant communities located along a topographically determined soil- and water-availability gradient. Multivariate analyses showed that trait variation was least at sites with shallower soils and putatively lower water availability, suggesting a convergence of water-use strategies at sites where plants are exposed to large seasonal water deficits. Stem hydraulic parameters, including stem-specific hydraulic conductivity, conduit diameter and maximum percentage embolism, were positively correlated, indicating the generality that larger conduit diameter permits greater hydraulic efficiency and is associated with greater seasonal reductions in hydraulic conductivity in this ecosystem. Wood density was not correlated with these traits, but closely associated with species' ability to withstand more negative water potentials during summer. Long-term integrated water-use efficiency was lower in shallow-rooted species that exhibited more negative summer water potentials. Specific leaf area and minimum leaf water potential were correlated with a number of separate traits, and appear to represent key axes of trait variation that describe the water-use strategies of different HFTs along the topographic gradient. Five HFTs were classified using a resemblance analysis according to combinations of traits that pertain to different water-use strategies among species; year-round active tree, year-round active shrub, hemiparasite, drought-suppressed broad-leaved shrub and drought-suppressed narrow-leaved shrub.

Leaf water relations during summer water deficit: differential responses in turgor maintenance and variation in leaf structure among different plant communities in south-western Australia.

We measured leaf water relations and leaf structural traits of 20 species from three communities growing along a topographical gradient. Our aim was to assess variation in seasonal responses in leaf water status and leaf tissue physiology between sites and among species in response to summer water deficit. Species from a ridge-top heath community showed the greatest reductions in pre-dawn leaf water potentials (Psi(leaf)) and stomatal conductance during summer; species from a valley-floor woodland and a midslope mallee community showed less reductions in these parameters. Heath species also displayed greater seasonal reduction in turgor-loss point (Psi(TLP)) than species from woodland or mallee communities. In general, species that had larger reductions in Psi(leaf) during summer showed significant shifts in either their osmotic potential at full turgor (Psi(pi 100); osmotic adjustment) or in tissue elasticity (epsilon(max)). Psi(pi 100) and epsilon(max) were negatively correlated, during both spring and summer, suggesting a trade-off between these different mechanisms to cope with water stress. Specific leaf area varied greatly among species, and was significantly correlated with seasonal changes in Psi(TLP) and pre-dawn Psi(leaf). These correlations suggest that leaf structure is a prerequisite for cellular mechanisms to be effective in adjusting to water deficit.