The peaked response of transpiration rate to vapour pressure deficit in field conditions can be explained by the temperature optimum of photosynthesis

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 feed-forward 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 stomatal conductance (gs) that is additional to the response of gs to increasing D alone. If this additional decrease is sufficiently steep and coupling between A and gs 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 CO2 and H2O 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. We conclude that field-based studies of the relationship between E and D need to consider signals related to changing photosynthetic rates in addition to purely hydraulic mechanisms. © 2014 Elsevier B.V

Rooting depth explains [CO <inf>2</inf>]× drought interaction in Eucalyptus saligna

Elevated atmospheric [CO 2] (eCa) often decreases stomatal conductance, which may delay the start of drought, as well as alleviate the effect of dry soil on plant water use and carbon uptake. We studied the interaction between drought and eCa in a whole-tree chamber experiment with Eucalyptus saligna. Trees were grown for 18 months in their Ca treatments before a 4-month dry-down. Trees grown in eCa were smaller than those grown in ambient Ca (aCa) due to an early growth setback that was maintained throughout the duration of the experiment. Pre-dawn leaf water potentials were not different between Ca treatments, but were lower in the drought treatment than the irrigated control. Counter to expectations, the drought treatment caused a larger reduction in canopy-average transpiration rates for trees in the eCa treatment compared with aCa. Total tree transpiration over the dry-down was positively correlated with the decrease in soil water storage, measured in the top 1.5 m, over the drying cycle; however, we could not close the water budget especially for the larger trees, suggesting soil water uptake below 1.5 m depth. Using neutron probe soil water measurements, we estimated fractional water uptake to a depth of 4.5 m and found that larger trees were able to extract more water from deep soil layers. These results highlight the interaction between rooting depth and response of tree water use to drought. The responses of tree water use to eCa involve interactions between tree size, root distribution and soil moisture availability that may override the expected direct effects of eCa. It is essential that these interactions be considered when interpreting experimental results. © 2011 The Author. Published by Oxford University Press. A ll rights reserved

Reconciling the optimal and empirical approaches to modelling stomatal conductance

Models of vegetation function are widely used to predict the effects of climate change on carbon, water and nutrient cycles of terrestrial ecosystems, and their feedbacks to climate. Stomatal conductance, the process that governs plant water use and carbon uptake, is fundamental to such models. In this paper, we reconcile two long-standing theories of stomatal conductance. The empirical approach, which is most commonly used in vegetation models, is phenomenological, based on experimental observations of stomatal behaviour in response to environmental conditions. The optimal approach is based on the theoretical argument that stomata should act to minimize the amount of water used per unit carbon gained. We reconcile these two approaches by showing that the theory of optimal stomatal conductance can be used to derive a model of stomatal conductance that is closely analogous to the empirical models. Consequently, we obtain a unified stomatal model which has a similar form to existing empirical models, but which now provides a theoretical interpretation for model parameter values. The key model parameter, g1, is predicted to increase with growth temperature and with the marginal water cost of carbon gain. The new model is fitted to a range of datasets ranging from tropical to boreal trees. The parameter g1 is shown to vary with growth temperature, as predicted, and also with plant functional type. The model is shown to correctly capture responses of stomatal conductance to changing atmospheric CO2, and thus can be used to test for stomatal acclimation to elevated CO2. The reconciliation of the optimal and empirical approaches to modelling stomatal conductance is important for global change biology because it provides a simple theoretical framework for analyzing, and simulating, the coupling between carbon and water cycles under environmental change. © 2011 Blackwell Publishing Ltd

Interactive effects of elevated CO <inf>2</inf> and drought on nocturnal water fluxes in Eucalyptus saligna

Nocturnal water flux has been observed in trees under a variety of environmental conditions and can be a significant contributor to diel canopy water flux. Elevated atmospheric CO 2 (elevated [CO 2]) can have an important effect on day-time plant water fluxes, but it is not known whether it also affects nocturnal water fluxes. We examined the effects of elevated [CO 2] on nocturnal water flux of field-grown Eucalyptus saligna trees using sap flux through the tree stem expressed on a sapwood area (J s) and leaf area (E t) basis. After 19 months growth under well-watered conditions, drought was imposed by withholding water for 5 months in the summer, ending with a rain event that restored soil moisture. Reductions in J s and E t were observed during the severe drought period in the dry treatment under elevated [CO 2], but not during moderate- and post-drought periods. Elevated [CO 2] affected night-time sap flux density which included the stem recharge period, called 'total night flux' (19:00 to 05:00, J s,r), but not during the post-recharge period, which primarily consisted of canopy transpiration (23:00 to 05:00, J s,c). Elevated [CO 2] wet (EW) trees exhibited higher J s,r than ambient [CO 2] wet trees (AW) indicating greater water flux in elevated [CO 2] under well-watered conditions. However, under drought conditions, elevated [CO 2] dry (ED) trees exhibited significantly lower J s,r than ambient [CO 2] dry trees (AD), indicating less water flux during stem recharge under elevated [CO 2]. J s,c did not differ between ambient and elevated [CO 2]. Vapour pressure deficit (D) was clearly the major influence on night-time sap flux. D was positively correlated with J s,r and had its greatest impact on J s,r at high D in ambient [CO 2]. Our results suggest that elevated [CO 2] may reduce night-time water flux in E. saligna when soil water content is low and D is high. While elevated [CO 2] affected J s,r, it did not affect day-time water flux in wet soil, suggesting that the responses of J s,r to environmental factors cannot be directly inferred from day-time patterns. Changes in J s,r are likely to influence pre-dawn leaf water potential, and plant responses to water stress. Nocturnal fluxes are clearly important for predicting effects of climate change on forest physiology and hydrology. © 2011 The Author. Published by Oxford University Press. A ll rights reserved

Whole-tree chambers for elevated atmospheric CO<inf>2</inf> experimentation and tree scale flux measurements in south-eastern Australia: The Hawkesbury Forest Experiment

Resolving ecophysiological processes in elevated atmospheric CO2 (Ca) at scales larger than single leaves poses significant challenges. Here, we describe a field-based experimental system designed to grow trees up to 9m tall in elevated Ca with the capacity to control air temperature and simultaneously measure whole-tree gas exchange. In western Sydney, Australia, we established the Hawkesbury Forest Experiment (HFE) where we built whole-tree chambers (WTC) to measure whole-tree CO2 and water fluxes of an evergreen broadleaf tree, Eucalyptus saligna. A single E. saligna tree was grown from seedling to small tree within each of 12 WTCs; six WTCs were maintained at ambient Ca and six WTCs were maintained at elevated Ca, targeted at ambient Ca +240μmolmol-1. All 12 WTCs were controlled to track ambient outside air temperature (Tair) and air water vapour deficit (Dair). During the experimental period, Tair, Dair and Ca in the WTCs were within 0.5°C, 0.3kPa, and 15μmolmol-1 of the set-points for 90% of the time, respectively. Diurnal responses of whole-tree CO2 and water vapour fluxes are analysed, demonstrating the ability of the tree chamber system to measure rapid environmental responses of these fluxes of entire trees. The light response of CO2 uptake for entire trees showed a clear diurnal hysteresis, attributed to stomatal closure at high Dair. Tree scale CO2 fluxes confirm the hypothesised deleterious effect of chilling night-time temperatures on whole-tree carbon gain in this subtropical Eucalyptus. The whole-tree chamber flux data add an invaluable scale to measurements in both ambient and elevated Ca and allow us to elucidate the mechanisms driving tree productivity responses to elevated Ca in interaction with water availability and temperature. © 2010 Elsevier B.V

Optimal stomatal behaviour around the world

© 2015 Macmillan Publishers Limited. All rights reserved. Stomatal conductance (g s) is a key land-surface attribute as it links transpiration, the dominant component of global land evapotranspiration, and photosynthesis, the driving force of the global carbon cycle. Despite the pivotal role of g s in predictions of global water and carbon cycle changes, a global-scale database and an associated globally applicable model of g s that allow predictions of stomatal behaviour are lacking. Here, we present a database of globally distributed g s obtained in the field for a wide range of plant functional types (PFTs) and biomes. We find that stomatal behaviour differs among PFTs according to their marginal carbon cost of water use, as predicted by the theory underpinning the optimal stomatal model and the leaf and wood economics spectrum. We also demonstrate a global relationship with climate. These findings provide a robust theoretical framework for understanding and predicting the behaviour of g s across biomes and across PFTs that can be applied to regional, continental and global-scale modelling of ecosystem productivity, energy balance and ecohydrological processes in a future changing climate

Optimal stomatal behaviour around the world

This is the author accepted manuscript. The final version is available from Springer Nature via the DOI in this recordStomatal conductance (g s) is a key land-surface attribute as it links transpiration, the dominant component of global land evapotranspiration, and photosynthesis, the driving force of the global carbon cycle. Despite the pivotal role of g s in predictions of global water and carbon cycle changes, a global-scale database and an associated globally applicable model of g s that allow predictions of stomatal behaviour are lacking. Here, we present a database of globally distributed g s obtained in the field for a wide range of plant functional types (PFTs) and biomes. We find that stomatal behaviour differs among PFTs according to their marginal carbon cost of water use, as predicted by the theory underpinning the optimal stomatal model and the leaf and wood economics spectrum. We also demonstrate a global relationship with climate. These findings provide a robust theoretical framework for understanding and predicting the behaviour of g s across biomes and across PFTs that can be applied to regional, continental and global-scale modelling of ecosystem productivity, energy balance and ecohydrological processes in a future changing climate.This research was supported by the Australian Research Council (ARC MIA Discovery Project 1433500-2012-14). A.R. was financially supported in part by The Next-Generation Ecosystem Experiments (NGEE-Arctic) project, which is supported by the Office of Biological and Environmental Research in the Department of Energy, Office of Science, and through the United States Department of Energy contract No. DE-AC02-98CH10886 to Brookhaven National Laboratory. M.O.d.B. acknowledges that the Brassica data were obtained within a research project financed by the Belgian Science Policy (OFFQ, contract number SD/AF/02) and coordinated by K. Vandermeiren at the Open-Top Chamber research facilities of CODA-CERVA (Tervuren, Belgium)