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Abstract Background Stomata respond to vapour pressure deficit (D) – when D increases, stomata begin to close. Closure is the result of a decline in guard cell turgor, but the link between D and turgor is poorly understood. We describe a model for stomatal responses to increasing D based upon cellular water relations. The model also incorporates impacts of increasing levels of water stress upon stomatal responses to increasing D. Results The model successfully mimics the three phases of stomatal responses to D and also reproduces the impact of increasing plant water deficit upon stomatal responses to increasing D. As water stress developed, stomata regulated transpiration at ever decreasing values of D. Thus, stomatal sensitivity to D increased with increasing water stress. Predictions from the model concerning the impact of changes in cuticular transpiration upon stomatal responses to increasing D are shown to conform to experimental data. Sensitivity analyses of stomatal responses to various parameters of the model show that leaf thickness, the fraction of leaf volume that is air-space, and the fraction of mesophyll cell wall in contact with air have little impact upon behaviour of the model. In contrast, changes in cuticular conductance and membrane hydraulic conductivity have significant impacts upon model behaviour. Conclusion Cuticular transpiration is an important feature of stomatal responses to D and is the cause of the 3 phase response to D. Feed-forward behaviour of stomata does not explain stomatal responses to D as feedback, involving water loss from guard cells, can explain these responses.

Phytotoxicity due to excessive boron (B) uptake by plants impedes routine agronomic utilisation of coal fly ash. We assessed 11 fly ashes (pH 3.14-10.77) having total B content ([B.sub.t]) of 12-136 mg/kg, of which 20-30% was hot water soluble ([B.sub.s]) in the acidic ashes (pH We conducted two pot trials in which canola was grown in soils amended with fly ash. In the first trial, an alkaline fly ash ([B.sub.t] 66 mg/kg) was incorporated at 5 rates of up to 625 Mg/ha into the top 50 mm of 2 acidic soils in 0.30-m-long intact cores, and sown with canola. Boron concentration in leaves at flowering reached the phytotoxic threshold, and both plant growth and seed yield were reduced, only at 625 Mg/ha. In the second trial, 4 fly ashes (pH 3.29-10.77, [B.sub.t] 12127 mg/kg) were incorporated at 4 rates of up to 108 Mg/ha into the top 0.10 m of 2 acidic soils in 1.0-m-long intact cores and then sown with canola. Ashes with highest [B.sub.t], when applied at 108 Mg/ha, increased B concentration in the topsoil only. Of the 2 ashes with the highest [B.sub.t], only that which produced low soil pH and applied at 108 Mg/ha increased B concentration in the shoot, but was still below phytotoxic threshold. The results suggest that B derived from these ashes may not cause phytotoxicity and excessive soil B accumulation if the ashes are applied at modest rates ( Additional keywords: trace elements, boron toxicity, soil boron, soil pH., Introduction Coal fly ash is a useful soil amendment for supplying plant and animal nutrients and improving soil physical and chemical properties. However, certain trace elements in fly ash can [...]

Vapour pressure deficit (D) is projected to increase in the future as temperature rises. In response to increased D , stomatal conductance (gs) and photosynthesis (A) are reduced, which may result in significant reductions in terrestrial carbon, water and energy fluxes. It is thus important for gas exchange models to capture the observed responses of gs and A with increasing D. We tested a series of coupled A – gs models against leaf gas exchange measurements from the Cumberland Plain Woodland (Australia), where D regularly exceeds 2 kPa and can reach 8 kPa in summer. Two commonly used A – gs models were not able to capture the observed decrease in A and gs with increasing D at the leaf scale. To explain this decrease in A and gs , two alternative hypotheses were tested: hydraulic limitation (i.e., plants reduce gs and/or A due to insufficient water supply) and non-stomatal limitation (i.e., downregulation of photosynthetic capacity). We found that the model that incorporated a non-stomatal limitation captured the observations with high fidelity and required the fewest number of parameters. Whilst the model incorporating hydraulic limitation captured the observed A and gs , it did so via a physical mechanism that is incorrect. We then incorporated a non-stomatal limitation into the stand model, MAESPA, to examine its impact on canopy transpiration and gross primary production. Accounting for a non-stomatal limitation reduced the predicted transpiration by ~19%, improving the correspondence with sap flow measurements, and gross primary production by ~14%. Given the projected global increases in D associated with future warming, these findings suggest that models may need to incorporate non-stomatal limitation to accurately simulate A and gs in the future with high D. Further data on non-stomatal limitation at high D should be a priority, in order to determine the generality of our results and develop a widely applicable model. [ABSTRACT FROM AUTHOR]

A direct relationship between gross ecosystem productivity (GEP) measured by the eddy covariance (EC) method and Moderate Resolution Imaging Spectroradiometer (MODIS) vegetation indices (VIs) has been observed in many temperate and tropical ecosystems. However, in Australian evergreen forests, and particularly sclerophyll woodlands, MODIS VIs do not capture seasonality of GEP. In this study, we re-evaluate the connection between satellite and flux tower data at four contrasting Australian ecosystems, through comparisons of ecosystem photosynthetic activity (GEP) and potential (e.g. ecosystem light use efficiency and quantum yield) with MODIS vegetation satellite products, including VIs, gross primary productivity (GPPMOD), leaf area index (LAIMOD), and fraction of photosynthetic active radiation (fPARMOD). We found that satellite derived greenness products constitute a measurement of ecosystem structure (e.g. leaf area index - quantity of leaves) and function (e.g. leaf level photosynthetic assimilation capacity - quality of leaves), rather than productivity. Our results show that in primarily meteorological-driven (e.g. photosynthetic active radiation, air temperature and/or precipitation) and relatively aseasonal vegetation photosynthetic potential ecosystems (e.g. evergreen wet sclerophyll forests), there were no statistically significant relationships between GEP and satellite derived measures of greenness. In contrast, for phenology-driven ecosystems (e.g. tropical savannas), changes in the vegetation status drove GEP, and tower-based measurements of photosynthetic activity were best represented by VIs. We observed the highest correlations between MODIS products and GEP in locations where key meteorological variables and vegetation phenology were synchronous (e.g. semi-arid Acacia woodlands) and low correlation at locations where they were asynchronous (e.g. Mediterranean ecosystems). Eddy covariance data offer much more than validation and/or calibration of satellite data and models. Knowledge of the conditions in which flux tower measurements and VIs or other remote sensing products converge greatly advances our understanding of the mechanisms driving the carbon cycle (phenology and climate drivers) and provides an ecological basis for interpretation of satellite derived measures of greenness. [ABSTRACT FROM AUTHOR]

ABSTRACTEucalypt saplings in north Australian savannas commonly die back, sometimes to ground level, during the 5 months of the long dry season. Water potentials are lower in saplings than large trees during the dry season, and we hypothesized that low water potentials may lead to high levels of xylem embolism and consequent death of branches and whole shoots. As the dry season progressed, hydraulic conductance of terminal branches decreased by 50% in Eucalyptus tetrodonta but not in Eucalyptus miniata saplings. Hydraulic conductance per leaf area decreased seasonally by 34% in E. tetrodonta branches. These decreases may be associated with the loss of leaves recorded from E. tetrodonta but not E. miniata branches. We modelled the effect of sequential loss of parallel resistors, representing petioles on a branch. This showed there is a non-linear decrease in flow as basal resistors are lost, which can lead to a decrease in mean flow per resistor due to increased mean path-length. Thus the observed loss of basal leaves, together with xylem embolism, probably contributed to the seasonal loss of hydraulic conductance in E. tetrodonta saplings. Loss of hydraulic conductance due to xylem embolism was generally low (< 15%) in both species, and the seasonal increase in embolism could not fully account for the decline in hydraulic conductance of E. tetrodonta branches. There was little evidence that branch and shoot death was caused by these levels of embolism. Developing an embolism vulnerability curve for species with long vessels is problematic and this issue is discussed. [ABSTRACT FROM AUTHOR]

Groundwater-dependent ecosystems (GDEs) are at risk globally due to unsustainable levels of groundwater extraction, especially in arid and semi-arid regions. In this review, we examine recent developments in the ecohydrology of GDEs with a focus on three knowledge gaps: (1) how do we locate GDEs, (2) how much water is transpired from shallow aquifers by GDEs and (3) what are the responses of GDEs to excessive groundwater extraction? The answers to these questions will determine water allocations that are required to sustain functioning of GDEs and to guide regulations on groundwater extraction to avoid negative impacts on GDEs. We discuss three methods for identifying GDEs: (1) techniques relying on remotely sensed information; (2) fluctuations in depth-to-groundwater that are associated with diurnal variations in transpiration; and (3) stable isotope analysis of water sources in the transpiration stream. We then discuss several methods for estimating rates of GW use, including direct measurement using sapflux or eddy covariance technologies, estimation of a climate wetness index within a Budyko framework, spatial distribution of evapotranspiration (ET) using remote sensing, groundwater modelling and stable isotopes. Remote sensing methods often rely on direct measurements to calibrate the relationship between vegetation indices and ET. ET from GDEs is also determined using hydrologic models of varying complexity, from the White method to fully coupled, variable saturation models. Combinations of methods are typically employed to obtain clearer insight into the components of groundwater discharge in GDEs, such as the proportional importance of transpiration versus evaporation (e.g. using stable isotopes) or from groundwater versus rainwater sources. Groundwater extraction can have severe consequences for the structure and function of GDEs. In the most extreme cases, phreatophytes experience crown dieback and death following groundwater drawdown. We provide a brief review of two case studies of the impacts of GW extraction and then provide an ecosystem-scale, multiple trait, integrated metric of the impact of differences in groundwater depth on the structure and function of eucalypt forests growing along a natural gradient in depth-to-groundwater. We conclude with a discussion of a depth-to-groundwater threshold in this mesic GDE. Beyond this threshold, significant changes occur in ecosystem structure and function. [ABSTRACT FROM AUTHOR]

Groundwater-dependent ecosystems (GDEs) are at risk globally due to unsustainable levels of groundwater extraction, especially in arid and semi-arid regions. In this review, we examine recent developments in the ecohydrology of GDEs with a focus on three knowledge gaps: (1) how do we locate GDEs, (2) how much water is transpired from shallow aquifers by GDEs; and (3) what are the responses of GDEs to excessive groundwater extraction? The answers to these questions will determine water allocations that are required to sustain functioning of GDEs and to guide regulations on groundwater extraction to avoid negative impacts on GDEs. We discuss three methods for identifying GDEs: (1) fluctuations in depth-to-groundwater that are associated with diurnal variations in transpiration, (2) stable isotope analysis of water sources in the transpiration stream; and (3) remote sensing methods. We then discuss several methods for estimating rates of GW use, including direct measurement using sapflux or eddy covariance technologies, estimation of a climate wetness index within a Budyko framework, spatial distribution of ET using remote sensing, groundwater modelling and stable isotopes. Remote sensing methods often rely on direct measurements to calibrate the relationship between vegetation indices and ET. ET from GDEs is also determined using hydrologic models of varying complexity, from the "White method" to fully coupled, variable saturation models. Combinations of methods are typically employed to obtain clearer insight into the components of groundwater discharge in GDEs, such as the proportional importance of transpiration vs. evaporation (e.g., using stable isotopes) or from groundwater vs. rainwater sources. Groundwater extraction can have severe consequences on structure and function of GDEs. In the most extreme cases, phreatophytes experience crown dieback and death following groundwater drawdown. We provide a brief review of two case studies of the impacts of GW extraction and discuss the use of C isotope ratios in xylem to reveal past influences of GW extraction. We conclude with a discussion of a depth-to-groundwater threshold in mesic and semi-arid GDEs. Across this threshold, significant changes occur in ecosystem structure and function. [ABSTRACT FROM AUTHOR]

Long-term declines in rainfall in south-western Australia have resulted in increased interest in the hydraulic characteristics of jarrah (Eucalyptus marginata Donn ex Smith) forest established in the region's drinking water catchments on rehabilitated bauxite mining sites. We hypothesized that in jarrah forest established on rehabilitated mine sites: (1) leaf area index (L) is independent of initial tree spacing; and (2) more densely planted trees have less leaf area for the same leaf mass, or the same sapwood area, and have denser sapwood. Initial stand densities ranged from about 600 to 9000 stems ha-1, and trees were 18 years old at the time of sampling. Leaf area index was unaffected by initial stand density, except in the most sparsely stocked stands where L was 1.2 compared with 2.0-2.5 in stands at other spacings. The ratio of leaf area to sapwood area (Al:As) was unaffected by tree spacing or tree size and was 0.2 at 1.3 m height and 0.25 at the crown base. There were small increases in sapwood density and decreases in leaf specific area with increased spacing. Tree diameter or basal area was a better predictor of leaf area than sapwood area. At the stand scale, basal area was a good predictor of L (r² = 0.98, n = 15) except in the densest stands. We conclude that the hydraulic attributes of this forest type are largely independent of initial tree spacing, thus simplifying parameterization of stand and catchment water balance models. [ABSTRACT FROM AUTHOR]