No abstract

The terrestrial carbon and water cycles are intimately linked: the carbon cycle is driven by photosynthesis, while the water balance is dominated by transpiration, and both fluxes are controlled by plant stomatal conductance. The ratio between these fluxes, the plant water-use efficiency (WUE), is a useful indicator of vegetation function. WUE can be estimated using several techniques, including leaf gas exchange, stable isotope discrimination, and eddy covariance. Here we compare global compilations of data for each of these three techniques. We show that patterns of variation in WUE across plant functional types (PFTs) are not consistent among the three datasets. Key discrepancies include the following: leaf-scale data indicate differences between needleleaf and broadleaf forests, but ecosystem-scale data do not; leaf-scale data indicate differences between C and C species, whereas at ecosystem scale there is a difference between C and C crops but not grasslands; and isotope-based estimates of WUE are higher than estimates based on gas exchange for most PFTs. Our study quantifies the uncertainty associated with different methods of measuring WUE, indicates potential for bias when using WUE measures to parameterize or validate models, and indicates key research directions needed to reconcile alternative measures of WUE.

The response of photosynthesis to temperature is a central facet of plant response to climate. Such responses have been found to be highly variable among species and among studies. Understanding this variability is key when trying to predict the effects of rising global temperatures on plant productivity. There are three major factors affecting the response of leaf net photosynthesis to temperature (A(n)-T): (i) photosynthetic biochemistry, (ii) respiration and (iii) vapour pressure deficit (D) and stomatal sensitivity to vapour pressure deficit during measurements. The overall goal of our study was to quantify the relative contribution of each of these factors in determining the response of A(n) to temperature. We first conducted a sensitivity analysis with a coupled photosynthesis-stomatal (A(n)-g(s)) model, using ranges for parameters of each factor taken from the literature, and quantified how these parameters affected the A(n)-T response. Second, we applied the A(n)-g(s) model to two example sets of field data, which had different optimum temperatures (T(opt)) of A(n), to analyse which factors were most important in causing the difference. We found that each of the three factors could have an equally large effect on T(opt) of A(n). In our comparison between two field datasets, the major cause for the difference in T(opt) was not the biochemical component, but rather the differences in respiratory components and in D conditions during measurements. We concluded that shifts in A(n)-T responses are not always driven by acclimation of photosynthetic biochemistry, but can result from other factors. The D conditions during measurements and stomatal responses to D also need to be quantified if we are to better understand and predict shifts in A(n)-T with climate.

Leaf transpiration rate (E) frequently shows a peaked response to increasing vapour pressure deficit (D). The mechanisms for the decrease in E at high D, known as the 'apparent feedforward response', are strongly debated but explanations to date have exclusively focused on hydraulic processes. However, stomata also respond to signals related to photosynthesis. We investigated whether the apparent feed-forward response of E to D in the field can be explained by the response of photosynthesis to temperature (T), which normally co-varies with D in field conditions. As photosynthesis decreases with increasing T past its optimum, it may drive a decrease in g s that is additional to the response of g s to increasing D alone. If this additional decrease is sufficiently steep and coupling between A and g s occurs, it could cause an overall decrease in E with increasing D. We tested this mechanism using a gas exchange model applied to leaf-scale and whole-tree CO 2 and H 2 O fluxes measured on Eucalyptus saligna growing in whole-tree chambers. A peaked response of E to D was observed at both leaf and whole-tree scales. We found that this peaked response was matched by a gas exchange model only when T effects on photosynthesis were incorporated. Furthermore, at elevated [CO 2 ], E peaked at higher D. We hypothesize thatcould be explained by an increase in the T optimum for A, as frequently observed, however we found no support for a higher T optimum for A in elevated [CO 2 ] in this study. We conclude that field-based studies of the relationship between E and D need to consider signals related to changing photosynthesis in addition to purely hydraulic mechanisms.

Abstract. Stomatal conductance (gs) affects the fluxes of carbon, energy and water between the vegetated land surface and the atmosphere. We test an implementation of an optimal stomatal conductance model within the Community Atmosphere Biosphere Land Exchange (CABLE) land surface model (LSM). In common with many LSMs, CABLE does not differentiate between gs model parameters in relation to plant functional type (PFT), but instead only in relation to photosynthetic pathway. We therefore constrained the key model parameter "g1" which represents a plants water use strategy by PFT based on a global synthesis of stomatal behaviour. As proof of concept, we also demonstrate that the g1 parameter can be estimated using two long-term average (1960–1990) bioclimatic variables: (i) temperature and (ii) an indirect estimate of annual plant water availability. The new stomatal models in conjunction with PFT parameterisations resulted in a large reduction in annual fluxes of transpiration (~ 30% compared to the standard CABLE simulations) across evergreen needleleaf, tundra and C4 grass regions. Differences in other regions of the globe were typically small. Model performance when compared to upscaled data products was not degraded, though the new stomatal conductance scheme did not noticeably change existing model-data biases. We conclude that optimisation theory can yield a simple and tractable approach to predicting stomatal conductance in LSMs.

Plants show flexible acclimation of leaf photosynthesis to temperature that depends both on their prevailing growth environment and the climate where they originated. This acclimation has been shown to involve changes in the temperature responses of the apparent maximum rate of Rubisco carboxylation (Vcmax) and apparent maximum rate of electron transport (Jmax), as well as changes in the ratio of these parameters. We asked whether such changes in photosynthetic biochemistry attributable to climate of origin are similar in nature and magnitude to those attributable to growth environment. To address this question, we measured temperature responses of photosynthesis and chlorophyll fluorescence on six Eucalyptus species from diverse geographical and climatic regions growing in a common garden. Measurements were made in three seasons, allowing us to compare interspecific differences with seasonal changes. We found significant interspecific differences in apparent Vcmax and Jmax standardized to 25 °C, but there were no significant differences in the temperature responses of these parameters among species. Comparing data across seasons, we found significant seasonal changes in apparent Vcmax25, but not in Jmax25, causing a change in their ratio (J/V ratio). However, there were no seasonal changes in the temperature response of either parameter. We concluded that the growth environment had a much larger effect on temperature response than climate of origin among this set of species. Mean daytime temperature increased by 15 °C from winter to summer, whereas we estimated that the seasonal change in J/V ratio would cause a change in the optimum temperature (Topt) for gross photosynthesis of 3.6 °C. Use of a general relationship to describe photosynthetic temperature acclimation resulted in a strong underestimation of the Topt for photosynthesis for these species. Our results indicated that variation in photosynthetic temperature responses cannot be captured in one simple relationship with growth temperature. Further comparative research on species groups will be needed to develop a basis for modelling these interspecific differences in plant temperature acclimation.