Effects of elevated atmospheric [CO2] on instantaneous transpiration efficiency at leaf and canopy scales in Eucalyptus saligna

Rising atmospheric concentrations of CO₂ (C_a) can reduce stomatal conductance and transpiration rate in trees, but the magnitude of this effect varies considerably among experiments. The theory of optimal stomatal behaviour predicts that the ratio of photosynthesis to transpiration (instantaneous transpiration efficiency, ITE) should increase in proportion to C_a. We hypothesized that plants regulate stomatal conductance optimally in response to rising C_a. We tested this hypothesis with data from young Eucalyptus saligna Sm. trees grown in 12 climate‐controlled whole‐tree chambers for 2 years at ambient and elevated C_a. Elevated C_a was ambient + 240 ppm, 60% higher than ambient C_a. Leaf‐scale gas exchange was measured throughout the second year of the study and leaf‐scale ITE increased by 60% under elevated C_a, as predicted. Values of leaf‐scale ITE depended strongly on vapour pressure deficit (D) in both CO₂ treatments. Whole‐canopy CO₂ and H₂O fluxes were also monitored continuously for each chamber throughout the second year. There were small differences in D between C_a treatments, which had important effects on values of canopy‐scale ITE. However, when C_a treatments were compared at the same D, canopy‐scale ITE was consistently increased by 60%, again as predicted. Importantly, leaf and canopy‐scale ITE were not significantly different, indicating that ITE was not scale‐dependent. Observed changes in transpiration rate could be explained on the basis that ITE increased in proportion to C_a. The effect of elevated C_a on photosynthesis increased with rising D. At high D, C_a had a large effect on photosynthesis and a small effect on transpiration rate. At low D, in contrast, there was a small effect of C_a on photosynthesis, but a much larger effect on transpiration rate. If shown to be a general response, the proportionality of ITE with C_a will allow us to predict the effects of C_a on transpiration rate.     

Root biomass distribution and soil properties of an open woodland on a duplex soil

Data on the distribution of root biomass are critical to understanding the ecophysiology of vegetation communities. This is particularly true when models are applied to describe ecohydrology and vegetation function. However, there is a paucity of such information across continental Australia. We quantified vertical and horizontal root biomass distribution in a woodland dominated by Angophora bakeri and Eucalyptus sclerophylla on the Cumberland Plains near Richmond, New South Wales. The site was characterised by a duplex (texture contrast) soil with the A horizon (to 70 cm) consisting of loamy sand and the B horizon (to > 10 m) consisting of sandy clay. The topsoil had a smaller bulk density, a smaller water holding capacity but a larger organic component and a larger hydraulic conductivity in comparison to the subsoil. Root biomass was sampled to 1.5 m depth and declined through the soil profile. Whilst total biomass in the B horizon was relatively small, its contribution to the function of the trees was highly significant. Coarse roots accounted for approximately 82% of the root mass recovered. Lateral distribution of fine roots was generally even but coarse roots were more likely to occur closer to tree stems. Variation in tree diameter explained 75% of the variation in total below-ground biomass. The trench method suggested the belowground biomass was 6.03?±?1.21 kg m^?2 but this method created bias towards sampling close to tree stems. We found that approximately 68% of root material was within a 2 m radius of tree stems and this made up 54% of the total number of samples but in reality, only approximately 5 to 10% of the site is within a 2 m radius of tree stems. Based on these proportions, our recalculated belowground biomass was 2.93?±?0.59 kg m^?2. These measurements provide valuable data for modeling of ecosystem water use and productivity.     

Comparing model predictions and experimental data for the response of stomatal conductance and guard cell turgor to manipulations of cuticular conductance, leaf-to-air vapour pressure difference and temperature: feedback mechanisms are able to account for all observations

Stomata respond to increasing leaf-to-air vapour pressure difference (LAVPD) ( D ) by closing. The mechanism by which this occurs is debated. A role for feedback and peristomatal transpiration has been proposed. In this paper, we apply a recent mechanistic model of stomatal behaviour, and compare model and experimental data for the influence of increasing D on stomatal conductance.
We manipulated cuticular conductance ( g _c) by three independent methods. First, we increased g _c by using a solvent mixture applied to both leaf surfaces prior to determining stomatal responses to D ; second, we increased g _c by increasing leaf temperature at constant D ; and third, we coated a small area of leaf with a light oil to decrease g _c. In all three experiments, experimental data and model outputs showed very close agreement.
We conclude, from the close agreement between model and experimental data and the fact that manipulations of g _c, and hence cuticular transpiration, influenced g _s in ways consistent with a feedback mechanism, that feedback is central in determining stomatal responses to D .     

Carbon balance of a tropical savanna of northern Australia

Through estimations of above- and below-ground standing biomass, annual biomass increment, fine root production and turnover, litterfall, canopy respiration and total soil CO₂ efflux, a carbon balance on seasonal and yearly time-scales is developed for a Eucalypt open-forest savanna in northern Australia. This carbon balance is compared to estimates of carbon fluxes derived from eddy covariance measurements conducted at the same site. The total carbon (C) stock of the savanna was 204±53 ton C ha^–1, with approximately 84% below-ground and 16% above-ground. Soil organic carbon content (0–1 m) was 151±33 ton C ha^–1, accounting for about 74% of the total carbon content in the ecosystem. Vegetation biomass was 53±20 ton C ha^–1, 39% of which was found in the root component and 61% in above-ground components (trees, shrubs, grasses). Annual gross primary production was 20.8 ton C ha^–1, of which 27% occurred in above-ground components and 73% below-ground components. Net primary production was 11 ton C ha^–1 year^–1, of which 8.0 ton C ha^–1 (73%) was contributed by below-ground net primary production and 3.0 ton C ha^–1 (27%) by above-ground net primary production. Annual soil carbon efflux was 14.3 ton C ha^–1 year^–1. Approximately three-quarters of the carbon flux (above-ground, below-ground and total ecosystem) occur during the 5–6 months of the wet season. This savanna site is a carbon sink during the wet season, but becomes a weak source during the dry season. Annual net ecosystem production was 3.8 ton C ha^–1 year^–1.     

Modelling Seasonal and Inter-annual Variations in Carbon and Water Fluxes in an Arid-Zone <Emphasis Type="Italic">Acacia</Emphasis> Savanna Woodland, 1981–2012

Changes in climatic characteristics such as seasonal and inter-annual variability may affect ecosystem structure and function, hence alter carbon and water budgets of ecosystems. Studies of modelling combined with field experiments can provide essential information to investigate interactions between carbon and water cycles and climate. Here we present a first attempt to investigate the long-term climate controls on seasonal patterns and inter-annual variations in water and carbon exchanges in an arid-zone savanna-woodland ecosystem using a detailed mechanistic soil–plant–atmosphere model (SPA), driven by leaf area index (LAI) simulated by an ecohydrological model (WAVES) and observed climate data during 1981–2012. The SPA was tested against almost 3 years of eddy covariance flux measurements in terms of gross primary productivity (GPP) and evapotranspiration (ET). The model was able to explain 80 and 71% of the variability of observed daily GPP and ET, respectively. Long-term simulations showed that carbon accumulation rates and ET ranged from 20.6 g C m^?2 mon^?1 in the late dry season to 45.8 g C m^?2 mon^?1 in the late wet season, respectively, primarily driven by seasonal variations in LAI and soil moisture. Large climate variations resulted in large seasonal variation in ecosystem water-use efficiency (eWUE). Simulated annual GPP varied between 146.4 and 604.7 g C m^?2 y^?1. Variations in annual ET coincided with that of GPP, ranging from 110.2 to 625.8 mm y^?1. Annual variations in GPP and ET were driven by the annual variations in precipitation and vapour pressure deficit (VPD) but not temperature. The linear coupling of simulated annual GPP and ET resulted in eWUE having relatively small year-to-year variation.     

Differences in osmotic adjustment,foliar abscisic acid dynamics,and stomatal regulation between an isohydric and anisohydric woody angiosperm during drought

Species are often classified along a continuum from isohydric to anisohydric, with isohydric species exhibiting tighter regulation of leaf water potential through stomatal closure in response to drought. We investigated plasticity in stomatal regulation in an isohydric (Eucalyptus camaldulensis) and an anisohydric (Acacia aptaneura) angiosperm species subject to repeated drying cycles. We also assessed foliar abscisic acid (ABA) content dynamics, aboveground/belowground biomass allocation and nonstructural carbohydrates. The anisohydric species exhibited large plasticity in the turgor loss point (Ψ_TLP), with plants subject to repeated drying exhibiting lower Ψ_TLP and correspondingly larger stomatal conductance at low water potential, compared to plants not previously exposed to drought. The anisohydric species exhibited a switch from ABA to water potential‐driven stomatal closure during drought, a response previously only reported for anisohydric gymnosperms. The isohydric species showed little osmotic adjustment, with no evidence of switching to water potential‐driven stomatal closure, but did exhibit increased root:shoot ratios. There were no differences in carbohydrate depletion between species. We conclude that a large range in Ψ_TLP and biphasic ABA dynamics are indicative of anisohydric species, and these traits are associated with exposure to low minimum foliar water potential, dense sapwood and large resistance to xylem embolism.     

Embolism recovery strategies and nocturnal water loss across species influenced by biogeographic origin

Drought‐induced tree mortality is expected to increase in future climates with the potential for significant consequences to global carbon, water, and energy cycles. Xylem embolism can accumulate to lethal levels during drought, but species that can refill embolized xylem and recover hydraulic function may be able to avoid mortality. Yet the potential controls of embolism recovery, including cross‐biome patterns and plant traits such as nonstructural carbohydrates (NSCs), hydraulic traits, and nocturnal stomatal conductance, are unknown. We exposed eight plant species, originating from mesic (tropical and temperate) and semi‐arid environments, to drought under ambient and elevated CO₂ levels, and assessed recovery from embolism following rewatering. We found a positive association between xylem recovery and NSCs, and, surprisingly, a positive relationship between xylem recovery and nocturnal stomatal conductance. Arid‐zone species exhibited greater embolism recovery than mesic zone species. Our results indicate that nighttime stomatal conductance often assumed to be a wasteful use of water, may in fact be a key part of plant drought responses, and contribute to drought survival. Findings suggested distinct biome‐specific responses that partially depended on species climate‐of‐origin precipitation or aridity index, which allowed some species to recover from xylem embolism. These findings provide improved understanding required to predict the response of diverse plant communities to drought. Our results provide a framework for predicting future vegetation shifts in response to climate change.     

Convergence of tree water use within an arid-zone woodland

We examined spatial and temporal patterns of tree water use and aspects of hydraulic architecture in four common tree species of central Australia—Corymbia opaca, Eucalyptus victrix, E. camaldulensis and Acacia aneura—to better understand processes that constrain water use in these environments. These four widely distributed species occupy contrasting niches within arid environments including woodlands, floodplains and riparian environments. Measurements of tree water use and leaf water potential were made at two sites with contrasting water table depths during a period of high soil water availability following summer rainfall and during a period of low soil water availability following 7 months of very little rainfall during 2007. There were significant differences in specific leaf area (SLA), sapwood area to leaf area ratios and sapwood density between species. Sapwood to leaf area ratio increased in all species from April to November indicating a decline in leaf area per unit sapwood area. Despite very little rainfall in the intervening period three species, C. opaca, E. victrix and E. camaldulensis maintained high leaf water potentials and tree water use during both periods. In contrast, leaf water potential and water use in the A. aneura were significantly reduced in November compared to April. Despite contrasting morphology and water use strategies, we observed considerable convergence in water use among the four species. Wood density in particular was strongly related to SLA, sapwood area to leaf area ratios and soil to leaf conductance, with all four species converging on a common relationship. Identifying convergence in hydraulic traits can potentially provide powerful tools for scaling physiological processes in natural ecosystems.