Microbial competition for phosphorus limits the CO2 response of a mature forest.

The capacity for terrestrial ecosystems to sequester additional carbon (C) with rising CO2 concentrations depends on soil nutrient availability1,2. Previous evidence suggested that mature forests growing on phosphorus (P)-deprived soils had limited capacity to sequester extra biomass under elevated CO2 (refs. 3-6), but uncertainty about ecosystem P cycling and its CO2 response represents a crucial bottleneck for mechanistic prediction of the land C sink under climate change7. Here, by compiling the first comprehensive P budget for a P-limited mature forest exposed to elevated CO2, we show a high likelihood that P captured by soil microorganisms constrains ecosystem P recycling and availability for plant uptake. Trees used P efficiently, but microbial pre-emption of mineralized soil P seemed to limit the capacity of trees for increased P uptake and assimilation under elevated CO2 and, therefore, their capacity to sequester extra C. Plant strategies to stimulate microbial P cycling and plant P uptake, such as increasing rhizosphere C release to soil, will probably be necessary for P-limited forests to increase C capture into new biomass. Our results identify the key mechanisms by which P availability limits CO2 fertilization of tree growth and will guide the development of Earth system models to predict future long-term C storage.

The fate of carbon in a mature forest under carbon dioxide enrichment.

Atmospheric carbon dioxide enrichment (eCO2) can enhance plant carbon uptake and growth1-5, thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO2 concentration6. Although evidence gathered from young aggrading forests has generally indicated a strong CO2 fertilization effect on biomass growth3-5, it is unclear whether mature forests respond to eCO2 in a similar way. In mature trees and forest stands7-10, photosynthetic uptake has been found to increase under eCO2 without any apparent accompanying growth response, leaving the fate of additional carbon fixed under eCO2 unclear4,5,7-11. Here using data from the first ecosystem-scale Free-Air CO2 Enrichment (FACE) experiment in a mature forest, we constructed a comprehensive ecosystem carbon budget to track the fate of carbon as the forest responded to four years of eCO2 exposure. We show that, although the eCO2 treatment of +150 parts per million (+38 per cent) above ambient levels induced a 12 per cent (+247 grams of carbon per square metre per year) increase in carbon uptake through gross primary production, this additional carbon uptake did not lead to increased carbon sequestration at the ecosystem level. Instead, the majority of the extra carbon was emitted back into the atmosphere via several respiratory fluxes, with increased soil respiration alone accounting for half of the total uptake surplus. Our results call into question the predominant thinking that the capacity of forests to act as carbon sinks will be generally enhanced under eCO2, and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO2 fertilization as a driver of increased carbon sinks in global forests.

Triggers of tree mortality under drought.

Severe droughts have caused widespread tree mortality across many forest biomes with profound effects on the function of ecosystems and carbon balance. Climate change is expected to intensify regional-scale droughts, focusing attention on the physiological basis of drought-induced tree mortality. Recent work has shown that catastrophic failure of the plant hydraulic system is a principal mechanism involved in extensive crown death and tree mortality during drought, but the multi-dimensional response of trees to desiccation is complex. Here we focus on the current understanding of tree hydraulic performance under drought, the identification of physiological thresholds that precipitate mortality and the mechanisms of recovery after drought. Building on this, we discuss the potential application of hydraulic thresholds to process-based models that predict mortality.

Optimal stomatal theory predicts CO2 responses of stomatal conductance in both gymnosperm and angiosperm trees.

Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (Anet ) and minimise transpirational water loss to achieve optimal intrinsic water-use efficiency (iWUE). We tested whether this theory can predict stomatal responses to elevated atmospheric CO2 (eCO2 ), and whether it can capture differences in responsiveness among woody plant functional types (PFTs). We conducted a meta-analysis of tree studies of the effect of eCO2 on iWUE and its components Anet and stomatal conductance (gs ). We compared three PFTs, using the unified stomatal optimisation (USO) model to account for confounding effects of leaf-air vapour pressure difference (D). We expected smaller gs , but greater Anet , responses to eCO2 in gymnosperms compared with angiosperm PFTs. We found that iWUE increased in proportion to increasing eCO2 in all PFTs, and that increases in Anet had stronger effects than reductions in gs . The USO model correctly captured stomatal behaviour with eCO2 across most datasets. The chief difference among PFTs was a lower stomatal slope parameter (g1 ) for the gymnosperm, compared with angiosperm, species. Land surface models can use the USO model to describe stomatal behaviour under changing atmospheric CO2 conditions.

Decoupling between stomatal conductance and photosynthesis occurs under extreme heat in broadleaf tree species regardless of water access.

High air temperatures increase atmospheric vapor pressure deficit (VPD) and the severity of drought, threatening forests worldwide. Plants regulate stomata to maximize carbon gain and minimize water loss, resulting in a close coupling between net photosynthesis (Anet ) and stomatal conductance (gs ). However, evidence for decoupling of gs from Anet under extreme heat has been found. Such a response both enhances survival of leaves during heat events but also quickly depletes available water. To understand the prevalence and significance of this decoupling, we measured leaf gas exchange in 26 tree and shrub species growing in the glasshouse or at an urban site in Sydney, Australia on hot days (maximum Tair > 40°C). We hypothesized that on hot days plants with ample water access would exhibit reduced Anet and use transpirational cooling leading to stomatal decoupling, whereas plants with limited water access would rely on other mechanisms to avoid lethal temperatures. Instead, evidence for stomatal decoupling was found regardless of plant water access. Transpiration of well-watered plants was 23% higher than model predictions during heatwaves, which effectively cooled leaves below air temperature. For hotter, droughted plants, the increase in transpiration during heatwaves was even more pronounced-gs was 77% higher than model predictions. Stomatal decoupling was found for most broadleaf evergreen and broadleaf deciduous species at the urban site, including some wilted trees with limited water access. Decoupling may simply be a passive consequence of the physical effects of high temperature on plant leaves through increased cuticular conductance of water vapor, or stomatal decoupling may be an adaptive response that is actively regulated by stomatal opening under high temperatures. This temperature response is not yet included in any land surface model, suggesting that model predictions of evapotranspiration may be underpredicted at high temperature and high VPD.

Tropical rainforest species have larger increases in temperature optima with warming than warm-temperate rainforest trees.

While trees can acclimate to warming, there is concern that tropical rainforest species may be less able to acclimate because they have adapted to a relatively stable thermal environment. Here we tested whether the physiological adjustments to warming differed among Australian tropical, subtropical and warm-temperate rainforest trees. Photosynthesis and respiration temperature responses were quantified in six Australian rainforest seedlings of tropical, subtropical and warm-temperate climates grown across four growth temperatures in a glasshouse. Temperature-response models were fitted to identify mechanisms underpinning the response to warming. Tropical and subtropical species had higher temperature optima for photosynthesis (ToptA ) than temperate species. There was acclimation of ToptA to warmer growth temperatures. The rate of acclimation (0.35-0.78°C °C-1 ) was higher in tropical and subtropical than in warm-temperate trees and attributed to differences in underlying biochemical parameters, particularly increased temperature optima of Vcmax25 and Jmax25 . The temperature sensitivity of respiration (Q10 ) was 24% lower in tropical and subtropical compared with warm-temperate species. Overall, tropical and subtropical species had a similar capacity to acclimate to changes in growth temperature as warm-temperate species, despite being grown at higher temperatures. Quantifying the physiological acclimation in rainforests can improve accuracy of future climate predictions and assess their potential vulnerability to warming.

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.

Predicting resilience through the lens of competing adjustments to vegetation function.

There is a pressing need to better understand ecosystem resilience to droughts and heatwaves. Eco-evolutionary optimization approaches have been proposed as means to build this understanding in land surface models and improve their predictive capability, but competing approaches are yet to be tested together. Here, we coupled approaches that optimize canopy gas exchange and leaf nitrogen investment, respectively, extending both approaches to account for hydraulic impairment. We assessed model predictions using observations from a native Eucalyptus woodland that experienced repeated droughts and heatwaves between 2013 and 2020, whilst exposed to an elevated [CO2 ] treatment. Our combined approaches improved predictions of transpiration and enhanced the simulated magnitude of the CO2 fertilization effect on gross primary productivity. The competing approaches also worked consistently along axes of change in soil moisture, leaf area, and [CO2 ]. Despite predictions of a significant percentage loss of hydraulic conductivity due to embolism (PLC) in 2013, 2014, 2016, and 2017 (99th percentile PLC > 45%), simulated hydraulic legacy effects were small and short-lived (2 months). Our analysis suggests that leaf shedding and/or suppressed foliage growth formed a strategy to mitigate drought risk. Accounting for foliage responses to water availability has the potential to improve model predictions of ecosystem resilience.

Mechanisms of xylem hydraulic recovery after drought in Eucalyptus saligna.

The mechanisms by which woody plants recover xylem hydraulic capacity after drought stress are not well understood, particularly with regard to the role of embolism refilling. We evaluated the recovery of xylem hydraulic capacity in young Eucalyptus saligna plants exposed to cycles of drought stress and rewatering. Plants were exposed to moderate and severe drought stress treatments, with recovery monitored at time intervals from 24 h to 6 months after rewatering. The percentage loss of xylem vessels due to embolism (PLV) was quantified at each time point using microcomputed tomography with stem water potential (Ψx ) and canopy transpiration (Ec ) measured before scans. Plants exposed to severe drought stress suffered high levels of embolism (47.38% ± 10.97% PLV) and almost complete canopy loss. No evidence of embolism refilling was observed at 24 h, 1 week, or 3 weeks after rewatering despite rapid recovery in Ψx . Recovery of hydraulic capacity was achieved over a 6-month period by growth of new xylem tissue, with canopy leaf area and Ec recovering over the same period. These findings indicate that E. saligna recovers slowly from severe drought stress, with potential for embolism to persist in the xylem for many months after rainfall events.

High safety margins to drought-induced hydraulic failure found in five pasture grasses.

Determining the relationship between reductions in stomatal conductance (gs ) and leaf water transport during dehydration is key to understanding plant drought responses. While numerous studies have analysed the hydraulic function of woody species, minimal research has been conducted on grasses. Here, we sought to characterize hydraulic vulnerability in five widely-occurring pasture grasses (including both C3 and C4 grasses) and determine whether reductions in gs and leaf hydraulic conductance (Kleaf ) during dehydration could be attributed to xylem embolism. Using the optical vulnerability (OV) technique, we found that all species were highly resistant to xylem embolism when compared to other herbaceous angiosperms, with 50% xylem embolism (PX50 ) occurring at xylem pressures ranging from -4.4 to -6.1 MPa. We observed similar reductions in gs and Kleaf under mild water stress for all species, occurring well before PX50 . The onset of xylem embolism (PX12 ) occurred consistently after stomatal closure and 90% reduction of Kleaf . Our results suggest that factors other than xylem embolism are responsible for the majority of reductions in gs and Kleaf during drought and reductions in the productivity of pasture species under moderate drought may not be driven by embolism.

Bridge to the future: Important lessons from 20 years of ecosystem observations made by the OzFlux network.

In 2020, the Australian and New Zealand flux research and monitoring network, OzFlux, celebrated its 20th anniversary by reflecting on the lessons learned through two decades of ecosystem studies on global change biology. OzFlux is a network not only for ecosystem researchers, but also for those 'next users' of the knowledge, information and data that such networks provide. Here, we focus on eight lessons across topics of climate change and variability, disturbance and resilience, drought and heat stress and synergies with remote sensing and modelling. In distilling the key lessons learned, we also identify where further research is needed to fill knowledge gaps and improve the utility and relevance of the outputs from OzFlux. Extreme climate variability across Australia and New Zealand (droughts and flooding rains) provides a natural laboratory for a global understanding of ecosystems in this time of accelerating climate change. As evidence of worsening global fire risk emerges, the natural ability of these ecosystems to recover from disturbances, such as fire and cyclones, provides lessons on adaptation and resilience to disturbance. Drought and heatwaves are common occurrences across large parts of the region and can tip an ecosystem's carbon budget from a net CO2 sink to a net CO2 source. Despite such responses to stress, ecosystems at OzFlux sites show their resilience to climate variability by rapidly pivoting back to a strong carbon sink upon the return of favourable conditions. Located in under-represented areas, OzFlux data have the potential for reducing uncertainties in global remote sensing products, and these data provide several opportunities to develop new theories and improve our ecosystem models. The accumulated impacts of these lessons over the last 20 years highlights the value of long-term flux observations for natural and managed systems. A future vision for OzFlux includes ongoing and newly developed synergies with ecophysiologists, ecologists, geologists, remote sensors and modellers.

To what extent can rising [CO2 ] ameliorate plant drought stress?

Plant responses to elevated atmospheric carbon dioxide (eCO2 ) have been hypothesized as a key mechanism that may ameliorate the impact of future drought. Yet, despite decades of experiments, the question of whether eCO2 reduces plant water use, yielding 'water savings' that can be used to maintain plant function during periods of water stress, remains unresolved. In this Viewpoint, we identify the experimental challenges and limitations to our understanding of plant responses to drought under eCO2 . In particular, we argue that future studies need to move beyond exploring whether eCO2 played 'a role' or 'no role' in responses to drought, but instead more carefully consider the timescales and conditions that would induce an influence. We also argue that considering emergent differences in soil water content may be an insufficient means of assessing the impact of eCO2 . We identify eCO2 impact during severe drought (e.g. to the point of mortality), interactions with future changes in vapour pressure deficit and uncertainty about changes in leaf area as key gaps in our current understanding. New insights into CO2 × drought interactions are essential to better constrain model theory that governs future climate model projections of land-atmosphere interactions during periods of water stress.

Drought by CO2 interactions in trees: a test of the water savings mechanism.

Elevated atmospheric CO2 (eCa ) may benefit plants during drought by reducing stomatal conductance (gs ) but any 'water savings effect' could be neutralized by concurrent stimulation of leaf area. We investigated whether eCa enhanced water savings, thereby ameliorating the impact of drought on carbon and water relations in trees. We report leaf-level gas exchange and whole-plant and soil water relations during a short-term dry-down in two Eucalyptus species with contrasting drought tolerance. Plants had previously been established for 9 to 11 months in steady-state conditions of ambient atmospheric CO2 (aCa ) and eCa , with half of each treatment group exposed to sustained drought for 5 to 7 months. The lower stomatal conductance under eCa did not lead to soil moisture savings during the dry-down due to the counteractive effect of increased whole-plant leaf area. Nonetheless, eCa -grown plants maintained higher photosynthetic rates and leaf water potentials, making them less stressed during the dry-down, despite being larger. These effects were more pronounced in the xeric species than the mesic species, and in previously water-stressed plants. Our findings indicate that eCa may enhance plant performance during drought despite a lack of soil water savings, especially in species with more conservative growth and water-use strategies.

Hydraulic failure and tree size linked with canopy die-back in eucalypt forest during extreme drought.

Eastern Australia was subject to its hottest and driest year on record in 2019. This extreme drought resulted in massive canopy die-back in eucalypt forests. The role of hydraulic failure and tree size on canopy die-back in three eucalypt tree species during this drought was examined. We measured pre-dawn and midday leaf water potential (Ψleaf ), per cent loss of stem hydraulic conductivity and quantified hydraulic vulnerability to drought-induced xylem embolism. Tree size and tree health was also surveyed. Trees with most, or all, of their foliage dead exhibited high rates of native embolism (78-100%). This is in contrast to trees with partial canopy die-back (30-70% canopy die-back: 72-78% native embolism), or relatively healthy trees (little evidence of canopy die-back: 25-31% native embolism). Midday Ψleaf was significantly more negative in trees exhibiting partial canopy die-back (-2.7 to -6.3 MPa), compared with relatively healthy trees (-2.1 to -4.5 MPa). In two of the species the majority of individuals showing complete canopy die-back were in the small size classes. Our results indicate that hydraulic failure is strongly associated with canopy die-back during drought in eucalypt forests. Our study provides valuable field data to help constrain models predicting mortality risk.

Integrating the evidence for a terrestrial carbon sink caused by increasing atmospheric CO2.

Atmospheric carbon dioxide concentration ([CO2 ]) is increasing, which increases leaf-scale photosynthesis and intrinsic water-use efficiency. These direct responses have the potential to increase plant growth, vegetation biomass, and soil organic matter; transferring carbon from the atmosphere into terrestrial ecosystems (a carbon sink). A substantial global terrestrial carbon sink would slow the rate of [CO2 ] increase and thus climate change. However, ecosystem CO2 responses are complex or confounded by concurrent changes in multiple agents of global change and evidence for a [CO2 ]-driven terrestrial carbon sink can appear contradictory. Here we synthesize theory and broad, multidisciplinary evidence for the effects of increasing [CO2 ] (iCO2 ) on the global terrestrial carbon sink. Evidence suggests a substantial increase in global photosynthesis since pre-industrial times. Established theory, supported by experiments, indicates that iCO2 is likely responsible for about half of the increase. Global carbon budgeting, atmospheric data, and forest inventories indicate a historical carbon sink, and these apparent iCO2 responses are high in comparison to experiments and predictions from theory. Plant mortality and soil carbon iCO2 responses are highly uncertain. In conclusion, a range of evidence supports a positive terrestrial carbon sink in response to iCO2 , albeit with uncertain magnitude and strong suggestion of a role for additional agents of global change.

Patterns of post-drought recovery are strongly influenced by drought duration, frequency, post-drought wetness, and bioclimatic setting.

Understanding vegetation recovery after drought is critical for projecting vegetation dynamics in future climates. From 1997 to 2009, Australia experienced a long-lasting drought known as the Millennium Drought (MD), which led to widespread reductions in vegetation productivity. However, vegetation recovery post-drought and its determinants remain unclear. This study leverages remote sensing products from different sources-fraction of absorbed photosynthetically active radiation (FPAR), based on optical data, and canopy density, derived from microwave data-and random forest algorithms to assess drought recovery over Australian natural vegetation during a 20-year period centered on the MD. Post-drought recovery was prevalent across the continent, with 6 out of 10 drought events seeing full recovery within about 6 months. Canopy density was slower to recover than leaf area seen in FPAR. The probability of full recovery was most strongly controlled by drought return interval, post-drought hydrological condition, and drought length. Full recovery was seldom observed when drought events occurred at intervals of 3 months or less, and moderately dry (standardized water balance anomaly [SWBA] within [-1, -0.76]) post-drought conditions resulted in less complete recovery than wet (SWBA > 0.3) post-drought conditions. Press droughts, which are long term but not extreme, delayed recovery more than pulse droughts (short term but extreme) and led to a higher frequency of persistent decline. Following press droughts, the frequency of persistent decline differed little among biome types but peaked in semi-arid regions across aridity levels. Forests and savanna required the longest recovery times for press drought, while grasslands were the slowest to recover for pulse drought. This study provides quantitative thresholds that could be used to improve the modeling of ecosystem dynamics post-drought.

Increasing aridity will not offset CO2 fertilization in fast-growing eucalypts with access to deep soil water.

Rising atmospheric [CO2 ] (Ca ) generally enhances tree growth if nutrients are not limiting. However, reduced water availability and elevated evaporative demand may offset such fertilization. Trees with access to deep soil water may be able to mitigate such stresses and respond more positively to Ca . Here, we sought to evaluate how increased vapor pressure deficit and reduced precipitation are likely to modify the impact of elevated Ca (eCa ) on tree productivity in an Australian Eucalyptus saligna Sm. plantation with access to deep soil water. We parameterized a forest growth simulation model (GOTILWA+) using data from two field experiments on E. saligna: a 2-year whole-tree chamber experiment with factorial Ca (ambient =380, elevated =620 µmol mol-1 ) and watering treatments, and a 10-year stand-scale irrigation experiment. Model evaluation showed that GOTILWA+ can capture the responses of canopy C uptake to (1) rising vapor pressure deficit (D) under both Ca treatments; (2) alterations in tree water uptake from shallow and deep soil layers during soil dry-down; and (3) the impact of irrigation on tree growth. Simulations suggest that increasing Ca up to 700 µmol mol-1 alone would result in a 33% increase in annual gross primary production (GPP) and a 62% increase in biomass over 10 years. However, a combined 48% increase in D and a 20% reduction in precipitation would halve these values. Our simulations identify high D conditions as a key limiting factor for GPP. They also suggest that rising Ca will compensate for increasing aridity limitations in E. saligna trees with access to deep soil water under non-nutrient limiting conditions, thereby reducing the negative impacts of global warming upon this eucalypt species. Simulation models not accounting for water sources available to deep-rooting trees are likely to overestimate aridity impacts on forest productivity and C stocks.

Optimal stomatal drought response shaped by competition for water and hydraulic risk can explain plant trait covariation.

The classical theory of stomatal optimization stipulates that stomata should act to maximize photosynthesis while minimizing transpiration. This theory, despite its remarkable success in reproducing empirical patterns, does not account for the fact that the available water to plants is dynamically regulated by plants themselves, and that plants compete for water in most locations. Here, we develop an alternative theory in which plants maximize the expected carbon gain under stochastic rainfall in a competitive environment. We further incorporate xylem hydraulic limitation as an additional constraint to transpiration and evaluate its impacts on stomatal optimization by incorporating the direct carbon cost of xylem recovery and the opportunity cost of reduced future photosynthesis as a result of irrecoverable xylem damage. We predict stomatal behaviour to be more conservative with a higher cost induced by xylem damage. By varying the unit carbon cost and extent of xylem recovery, characterizing the direct and opportunity cost of xylem damage, respectively, our model can reproduce several key patterns of stomatal-hydraulic trait covariations. By addressing the key elements of water limitation in plant gas exchange simultaneously, including plants' self-regulation of water availability, competition for water and hydraulic risk, our study provides a comprehensive theoretical basis for understanding stomatal behaviour.

Plant profit maximization improves predictions of European forest responses to drought.

Knowledge of how water stress impacts the carbon and water cycles is a key uncertainty in terrestrial biosphere models. We tested a new profit maximization model, where photosynthetic uptake of CO2 is optimally traded against plant hydraulic function, as an alternative to the empirical functions commonly used in models to regulate gas exchange during periods of water stress. We conducted a multi-site evaluation of this model at the ecosystem scale, before and during major droughts in Europe. Additionally, we asked whether the maximum hydraulic conductance in the soil-plant continuum kmax (a key model parameter which is not commonly measured) could be predicted from long-term site climate. Compared with a control model with an empirical soil moisture function, the profit maximization model improved the simulation of evapotranspiration during the growing season, reducing the normalized mean square error by c. 63%, across mesic and xeric sites. We also showed that kmax could be estimated from long-term climate, with improvements in the simulation of evapotranspiration at eight out of the 10 forest sites during drought. Although the generalization of this approach is contingent upon determining kmax , it presents a mechanistic trait-based alternative to regulate canopy gas exchange in global models.

Visual and hydraulic techniques produce similar estimates of cavitation resistance in woody species.

Hydraulic failure of the plant vascular system is a principal cause of forest die-off under drought. Accurate quantification of this process is essential to our understanding of the physiological mechanisms underpinning plant mortality. Imaging techniques increasingly are applied to estimate xylem cavitation resistance. These techniques allow for in situ measurement of embolism formation in real time, although the benefits and trade-offs associated with different techniques have not been evaluated in detail. Here we compare two imaging methods, microcomputed tomography (microCT) and optical vulnerability (OV), to standard hydraulic methods for measurement of cavitation resistance in seven woody species representing a diversity of major phylogenetic and xylem anatomical groups. Across the seven species, there was strong agreement between cavitation resistance values (P50 ) estimated from visualization techniques (microCT and OV) and between visual techniques and hydraulic techniques. The results indicate that visual techniques provide accurate estimates of cavitation resistance and the degree to which xylem hydraulic function is impacted by embolism. Results are discussed in the context of trade-offs associated with each technique and possible causes of discrepancy between estimates of cavitation resistance provided by visual and hydraulic techniques.