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Rice (Oryza sativa) is a water-intensive crop, and like other plants uses stomata to balance CO2 uptake with water-loss. To identify agronomic traits related to rice stomatal complexes, an anatomical screen of 64 Thai and 100 global rice cultivars was undertaken. Epidermal outgrowths called papillae were identified on the stomatal subsidiary cells of all cultivars. These were also detected on eight other species of the Oryza genus but not on the stomata of any other plant species we surveyed. Our rice screen identified two cultivars that had “mega-papillae” that were so large or abundant that their stomatal pores were partially occluded; Kalubala Vee had extra-large papillae, and Dharia had approximately twice the normal number of papillae. These were most accentuated on the flag leaves, but mega-papillae were also detectable on earlier forming leaves. Energy dispersive X-Ray spectrometry revealed that silicon is the major component of stomatal papillae. We studied the potential function(s) of mega-papillae by assessing gas exchange and pathogen infection rates. Under saturating light conditions, mega-papillae bearing cultivars had reduced stomatal conductance and their stomata were slower to close and re-open, but photosynthetic assimilation was not significantly affected. Assessment of an F3 hybrid population treated with Xanthomonas oryzae pv. oryzicola indicated that subsidiary cell mega-papillae may aid in preventing bacterial leaf streak infection. Our results highlight stomatal mega-papillae as a novel rice trait that influences gas exchange, stomatal dynamics, and defense against stomatal pathogens which we propose could benefit the performance of future rice crops.

Land surface modellers need measurable proxies to constrain the quantity of carbon dioxide (CO2) assimilated by continental plants through photosynthesis, known as gross primary production (GPP). Carbonyl sulfide (COS), which is taken up by leaves through their stomates and then hydrolysed by photosynthetic enzymes, is a candidate GPP proxy. A former study with the ORCHIDEE land surface model used a fixed ratio of COS uptake to CO2 uptake normalised to respective ambient concentrations for each vegetation type (leaf relative uptake, LRU) to compute vegetation COS fluxes from GPP. The LRU approach is known to have limited accuracy since the LRU ratio changes with variables such as photosynthetically active radiation (PAR): while CO2 uptake slows under low light, COS uptake is not light limited. However, the LRU approach has been popular for COS–GPP proxy studies because of its ease of application and apparent low contribution to uncertainty for regional-scale applications. In this study we refined the COS–GPP relationship and implemented in ORCHIDEE a mechanistic model that describes COS uptake by continental vegetation. We compared the simulated COS fluxes against measured hourly COS fluxes at two sites and studied the model behaviour and links with environmental drivers. We performed simulations at a global scale, and we estimated the global COS uptake by vegetation to be −756 Gg S yr−1, in the middle range of former studies (−490 to −1335 Gg S yr−1). Based on monthly mean fluxes simulated by the mechanistic approach in ORCHIDEE, we derived new LRU values for the different vegetation types, ranging between 0.92 and 1.72, close to recently published averages for observed values of 1.21 for C4 and 1.68 for C3 plants. We transported the COS using the monthly vegetation COS fluxes derived from both the mechanistic and the LRU approaches, and we evaluated the simulated COS concentrations at NOAA sites. Although the mechanistic approach was more appropriate when comparing to high-temporal-resolution COS flux measurements, both approaches gave similar results when transporting with monthly COS fluxes and evaluating COS concentrations at stations. In our study, uncertainties between these two approaches are of secondary importance compared to the uncertainties in the COS global budget, which are currently a limiting factor to the potential of COS concentrations to constrain GPP simulated by land surface models on the global scale.

This study aimed to assess the toxicity of glyphosate (GLY; 0, 10, 20 and 30?mg?kg-1) in Solanum lycopersicum L., particularly focusing on the photosynthetic metabolism. By combining ecophysiological, ultrastructural, biochemical and molecular tools, the results revealed that the exposure of tomato plants to GLY led to alterations in leaf water balance regulation [increasing stomatal conductance (gs) and decreasing water use efficiency (WUEi) at higher concentrations] and induced slight alterations in the structural integrity of cells, mainly in chloroplasts, accompanied by a loss of cell viability. Moreover, the transcriptional and biochemical control of several photosynthetic-related parameters was reduced upon GLY exposure. However, in vivo chlorophyll fluorometry and IRGA gas-exchange studies revealed that the photosynthetic yield of S. lycopersicum was not repressed by GLY. Overall, GLY impacts cellular and subcellular homeostasis (by affecting chloroplast structure, reducing photosynthetic pigments and inhibiting photosynthetic-related genes transcription), and leaf structure, but is not reducing the carbon flow on a leaf area basis. Altogether, these results suggest a trade-off effect in which GLY-induced toxicity is compensated by a higher photosynthetic activity related to GLY-induced dysfunction in gs and an increase in mesophyll thickness/density, allowing the viable leaf cells to maintain their photosynthetic capacity., FCT -Fundação para a Ciência e a Tecnologia(SFRH/BD/115643/2016), info:eu-repo/semantics/publishedVersion

Abiotic stresses such as drought and salinity constantly threaten food security. Biochar as a soil amendment has the potential to ameliorate soil and alleviate drought and salinity stress. Multiple studies have been conducted to evaluate the effect of biochar in alleviating independent drought or salinity stress. However, the potential of biochar in mitigating the combined drought and salinity stress on plants has not been studied so far. Therefore, a pot experiment was conducted in the climate-controlled chamber with the objective to investigate the effect of biochar on growth, physiology, and yield of quinoa under independent and combined drought and salinity stress. Quinoa plants were subjected to three irrigation treatments i.e., full irrigation (FI), deficit irrigation (DI), and alternate root-zone drying irrigation (ARD), two saline water treatments (0 and 400 mM) and two levels of biochar (0% and 5% by weight). In the FI treatment, plants were irrigated daily to maintain pot water-holding capacity. In limited irrigation treatments, 70% water of FI was applied either to the whole pot in DI or to one side of the pot alternating in ARD, respectively. The results showed that combined drought and salinity stress drastically affected growth and performance of quinoa compared to the independent drought or salinity stress. However, soil amendment with biochar had positive effect in mitigating both independent and combined effect of drought and salinity on quinoa plants. Furthermore, biochar amendment in ARD under salinity significantly enhanced plant height, shoot biomass, and grain by 11.7%, 18.8%, and 10.2% as compared with DI under salinity, respectively. In addition, leaf photosynthetic rate (An) and stomatal conductance (gs) decreased under limited saline irrigation. Moreover, the interactive effect of biochar and ARD efficiently adjusted the balance between chemical signal (leaf ABA) and hydraulic signal (leaf water potential). Thus, intrinsic water use efficiency (WUEi) and yield in ARD were significantly enhanced compared to DI, especially under salinity stress. Overall, biochar in combination with ARD might be a wise approach for sustaining crop productivity in salt affected and drought stressed areas of the world to ensure food security.

Stomatal regulation and whole plant hydraulic signaling affect water fluxes and stress in plants. Land surface models and crop models use a coupled photosynthesis–stomatal conductance modeling approach. Those models estimate the effect of soil water stress on stomatal conductance directly from soil water content or soil hydraulic potential without explicit representation of hydraulic signals between the soil and stomata. In order to explicitly represent stomatal regulation by soil water status as a function of the hydraulic signal and its relation to the whole plant hydraulic conductance, we coupled the crop model LINTULCC2 and the root growth model SLIMROOT with Couvreur's root water uptake model (RWU) and the HILLFLOW soil water balance model. Since plant hydraulic conductance depends on the plant development, this model coupling represents a two-way coupling between growth and plant hydraulics. To evaluate the advantage of considering plant hydraulic conductance and hydraulic signaling, we compared the performance of this newly coupled model with another commonly used approach that relates root water uptake and plant stress directly to the root zone water hydraulic potential (HILLFLOW with Feddes' RWU model). Simulations were compared with gas flux measurements and crop growth data from a wheat crop grown under three water supply regimes (sheltered, rainfed, and irrigated) and two soil types (stony and silty) in western Germany in 2016. The two models showed a relatively similar performance in the simulation of dry matter, leaf area index (LAI), root growth, RWU, gross assimilation rate, and soil water content. The Feddes model predicts more stress and less growth in the silty soil than in the stony soil, which is opposite to the observed growth. The Couvreur model better represents the difference in growth between the two soils and the different treatments. The newly coupled model (HILLFLOW–Couvreur's RWU–SLIMROOT–LINTULCC2) was also able to simulate the dynamics and magnitude of whole plant hydraulic conductance over the growing season. This demonstrates the importance of two-way feedbacks between growth and root water uptake for predicting the crop response to different soil water conditions in different soils. Our results suggest that a better representation of the effects of soil characteristics on root growth is needed for reliable estimations of root hydraulic conductance and gas fluxes, particularly in heterogeneous fields. The newly coupled soil–plant model marks a promising approach but requires further testing for other scenarios regarding crops, soil, and climate.

© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust The ratio of leaf intercellular to ambient CO 2 (χ) is modulated by stomatal conductance (g s ). These quantities link carbon (C) assimilation with transpiration, and along with photosynthetic capacities (V cmax and J max ) are required to model terrestrial C uptake. We use optimization criteria based on the growth environment to generate predicted values of photosynthetic and water-use efficiency traits and test these against a unique dataset. Leaf gas-exchange parameters and carbon isotope discrimination were analysed in relation to local climate across a continental network of study sites. Sun-exposed leaves of 50 species at seven sites were measured in contrasting seasons. Values of χ predicted from growth temperature and vapour pressure deficit were closely correlated to ratios derived from C isotope (δ 13 C) measurements. Correlations were stronger in the growing season. Predicted values of photosynthetic traits, including carboxylation capacity (V cmax ), derived from δ 13 C, growth temperature and solar radiation, showed meaningful agreement with inferred values derived from gas-exchange measurements. Between-site differences in water-use efficiency were, however, only weakly linked to the plant's growth environment and did not show seasonal variation. These results support the general hypothesis that many key parameters required by Earth system models are adaptive and predictable from plants’ growth environments.

Plant functional traits have shown to be relevant predictors of forest functional responses to climate change. However, the trait-based approach to study plant performances and ecological strategies has mostly been focused on trait comparisons at the interspecific and intraspecific levels. In this study, we analyzed traits variation and association at the individual level. We measured wood and leaf traits at different height locations within the crown of five individuals of Pericopsis elata (Harms) Meeuwen (Fabaceae) from the northern tropical forest of the Democratic Republic of the Congo. All traits varied between and within individuals. The between-individual variation was more important for leaf traits (23%&ndash, 48%) than for wood traits (~10%) where the within-individual variation showed to be more important (33%&ndash, 39%). The sample location height within the crown was found to be the driving factor of this within-individual variation. In a gradient from the base to the top of the crown, theoretical specific hydraulic conductivity and specific leaf area decreased while the stomatal density increased. We found significant relationships among traits and between wood and leaf traits. However, these relationships varied with the position within the crown. The relationship between vessel size and vessel density was negative at the bottom part of the crown but positive upward. Also, the negative relationship between stomatal density and stomatal size became stronger with increasing height within the crown. Finally, the positive relationship between specific leaf area and theoretical specific hydraulic conductivity became stronger in higher parts of the crown, suggesting that P. elata constantly adapts its water use with respect to its water supply, more strongly at the top of the crown where the environment is more extreme and less buffered against environmental fluctuations.

Aim of this study was to investigate the effect of water shortage in wild jujube plants, Ziziphus lotus (L.) Lam, and how it is related to its ecological success. All leaf anatomical traits were significantly affected following soil drying. Stressed plants displayed more negative stem water potential (psi(w)) and osmotic potential values of ca. -3.5 and -4.5 MPa, respectively, after 30 d. The relative water content declined although it still maintained high values (>= 75%). The net photosynthetic rate, stomatal conductance, and transpiration rate were significantly inhibited as psi(w) decreased. However, the intrinsic water-use efficiency increased as water deficit was intensified; the difference became significant only after 30 d. As a consequence, the effective quantum yield of PSII photochemistry and the photochemical quenching coefficient significantly decreased although the maximal quantum yield of PSII photochemistry ratio remained statisticaly unchanged. Plants could maintain their water status sufficiently by increasing proline 1.7-fold and sucrose 1.8-fold in their leaves, respectively. The largest accumulation of both solutes may avoid photodamages at cellular level and play a critical role in maintaining osmotic adjustment.

Future climate change has the potential to increase drought in many regions of the globe, making it essential that land surface models (LSMs) used in coupled climate models realistically capture the drought responses of vegetation. Recent data syntheses show that drought sensitivity varies considerably among plants from different climate zones, but state-of-the-art LSMs currently assume the same drought sensitivity for all vegetation. We tested whether variable drought sensitivities are needed to explain the observed large-scale patterns of drought impact on the carbon, water and energy fluxes. We implemented data-driven drought sensitivities in the Community Atmosphere Biosphere Land Exchange (CABLE) LSM and evaluated alternative sensitivities across a latitudinal gradient in Europe during the 2003 heatwave. The model predicted an overly abrupt onset of drought unless average soil water potential was calculated with dynamic weighting across soil layers. We found that high drought sensitivity at the most mesic sites, and low drought sensitivity at the most xeric sites, was necessary to accurately model responses during drought. Our results indicate that LSMs will over-estimate drought impacts in drier climates unless different sensitivity of vegetation to drought is taken into account.

The seasonality of the NEE of the northern boreal coniferous forests was investigated by means of inversion modelling using eddy covariance data. Eddy covariance data was used to optimize the biochemical model parameters. Our study sites consisted of three Scots pine (l. Pinus sylvestris) forests and one Norway spruce (l. Picea abies) forest that were located in Finland and Sweden. We obtained temperature and seasonal dependence for the biochemical model parameters: the maximum rate of carboxylation (Vc(max)) and the maximum rate of electron transport (Jmax). Both of the parameters were optimized without assumptions about their mutual magnitude. The values obtained for the biochemical model parameters were similar at all the sites during summer time. To describe seasonality, different temperature fits were made for the spring, summer and autumn periods. During summer, average Jmax across the sites was 54.0 μmol m−2 s−1 (variance 31.2 μmol m−2 s−1) and Vc(max) was 12.0 μmol m−2 s−1 (variance 6.6 μmol m−2 s−1) at 17°C. The sensitivity of the model to LAI and atmospheric soil water stress was also studied. The impact of seasonality on annual GPP was 17% when only summertime parameterization was used throughout the year compared to seasonally changing parameterizations.

Soil is a major contributor to the biosphere– atmosphere exchange of carbonyl sulfide (COS) and carbon monoxide (CO). COS is a tracer with which to quantify terrestrial photosynthesis based on the coupled leaf uptake of COS and CO2, but such use requires separating soil COS flux, which is unrelated to photosynthesis, from ecosystem COS uptake. For CO, soil is a significant natural sink that influences the tropospheric CO budget. In the boreal forest, magnitudes and variabilities of soil COS and CO fluxes remain poorly understood. We measured hourly soil fluxes of COS, CO, and CO2 over the 2015 late growing season (July to November) in a Scots pine forest in Hyytiälä, Finland. The soil acted as a net sink of COS and CO, with average uptake rates around 3 pmol m−2 s−1 for COS and 1 nmol m−2 s−1 for CO. Soil respiration showed seasonal dynamics controlled by soil temperature, peaking at around 4 μmol m−2 s−1 in late August and September and dropping to 1–2μmolm−2 s−1 in October. In contrast, seasonal variations of COS and CO fluxes were weak and mainly driven by soil moisture changes through diffusion limitation. COS and CO fluxes did not appear to respond to temperature variation, although they both correlated well with soil respiration in specific temperature bins. However, COS : CO2 and CO : CO2 flux ratios increased with temperature, suggesting possible shifts in active COS- and CO-consuming microbial groups. Our results show that soil COS and CO fluxes do not have strong variations over the late growing season in this boreal forest and can be represented with the fluxes during the photosynthetically most active period. Well-characterized and relatively invariant soil COS fluxes strengthen the case for using COS as a photosynthetic tracer in boreal forests.

Background and Aims Experimental drought is well documented to induce a decline in photosynthetic capacity. However, if given time to acclimate to low water availability, the photosynthetic responses of plants to low soil moisture content may differ from those found in short-term experiments. This study aims to test whether plants acclimate to long-term water stress by modifying the functional relationships between photosynthetic traits and water stress, and whether species of contrasting habitat differ in their degree of acclimation. Methods Three Eucalyptus taxa from xeric and riparian habitats were compared with regard to their gas exchange responses under short- and long-term drought. Photosynthetic parameters were measured after 2 and 4 months of watering treatments, namely field capacity or partial drought. At 4 months, all plants were watered to field capacity, then watering was stopped. Further measurements were made during the subsequent ‘drying-down’, continuing until stomata were closed. Key Results Two months of partial drought consistently reduced assimilation rate, stomatal sensitivity parameters (g1), apparent maximum Rubisco activity (V′cmaxVcmax′) and maximum electron transport rate (J′maxJmax′). Eucalyptus occidentalis from the xeric habitat showed the smallest decline in V′cmaxVcmax′ and J′maxJmax′; however, after 4 months, V′cmaxVcmax′ and J′maxJmax′ had recovered. Species differed in their degree of V′cmaxVcmax′ acclimation. Eucalyptus occidentalis showed significant acclimation of the pre-dawn leaf water potential at which the V′cmaxVcmax′ and ‘true’ Vcmax (accounting for mesophyll conductance) declined most steeply during drying-down. Conclusions The findings indicate carbon loss under prolonged drought could be over-estimated without accounting for acclimation. In particular, (1) species from contrasting habitats differed in the magnitude of V′cmax reduction in short-term drought; (2) long-term drought allowed the possibility of acclimation, such that V′cmax reduction was mitigated; (3) xeric species showed a greater degree of V′cmax acclimation; and (4) photosynthetic acclimation involves hydraulic adjustments to reduce water loss while maintaining photosynthesis.

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 constrained the key model parameter "g1", which represents plant 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 model, 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 against upscaled data products was not degraded, but did not noticeably reduce existing model–data biases. We identified assumptions relating to the coupling of the vegetation to the atmosphere and the parameterisation of the minimum stomatal conductance as areas requiring further investigation in both CABLE and potentially other LSMs. We conclude that optimisation theory can yield a simple and tractable approach to predicting stomatal conductance in LSMs.

Optimization models of stomatal conductance (g(s)) attempt to explain observed stomatal behaviour in terms of cost-benefit tradeoffs. While the benefit of stomatal opening through increased CO2 uptake is clear, currently the nature of the associated cost(s) remains unclear. We explored the hypothesis that g(s) maximizes leaf photosynthesis, where the cost of stomatal opening arises from nonstomatal reductions in photosynthesis induced by leaf water stress. We analytically solved two cases, CAP and MES, in which reduced leaf water potential leads to reductions in carboxylation capacity (CAP) and mesophyll conductance (g(m)) (MES). Both CAP and MES predict the same one-parameter relationship between the intercellular:atmospheric CO2 concentration ratio (c(i)/c(a)) and vapour pressure deficit (VPD, D), viz. c(i)/c(a) approximate to xi/xi (xi+D), as that obtained from previous optimization models, with the novel feature that the parameter xi is determined unambiguously as a function of a small number of photosynthetic and hydraulic variables. These include soil-to-leaf hydraulic conductance, implying a stomatal closure response to drought. MES also predicts that g(s)/g(m) is closely related to c(i)/c(a) and is similarly conservative. These results are consistent with observations, give rise to new testable predictions, and offer new insights into the covariation of stomatal, mesophyll and hydraulic conductances.

The pull of water from the soil to the leaves causes water in the transpiration stream to be under negative pressure decreasing the water potential below zero. The osmotic concentration also contributes to the decrease in leaf water potential but withmuch lesser extent. Thus, the surface tension force is approximately balanced by a force induced by negative water potential resulting in concavely curved water-air interfaces in leaves. The lowered water potential causes a reduction in the equilibrium water vapor pressure in internal (sub-stomatal/ intercellular) cavities in relation to that over water with the potential of zero, i.e., over the flat surface. The curved surface causes a reduction also in the equilibrium vapor pressure of dissolved CO2, thus enhancing its physical solubility to water. Although the water vapor reduction is acknowledged by plant physiologists its consequences for water vapor exchange at low water potential values have received very little attention. Consequences of the enhanced CO2 solubility to a leaf water-carbon budget have not been considered at all before this study. We use theoretical calculations and modeling to show how the reduction in the vapor pressures affects transpiration and carbon assimilation rates. Our results indicate that the reduction in vapor pressures of water and CO2 could enhance plant water use efficiency up to about 10% at a leaf water potential of -2 MPa, and much more when water potential decreases further. The low water potential allows for a direct stomatal water vapor uptake from the ambient air even at sub-100% relative humidity values. This alone could explain the observed rates of foliar water uptake by e.g., the coastal redwood in the fog belt region of coastal California provided the stomata are sufficiently open. The omission of the reduction in the water vapor pressure causes a bias in the estimates of the stomatal conductance and leaf internal CO2 concentration based on leaf gas exchange measurements. Manufactures of leaf gas exchange measurement systems should incorporate leaf water potentials in measurement set-ups.

We investigated the drivers of street tree transpiration in boreal conditions, in order to better understand tree water use in the context of urban tree planning and stormwater management. Two streets built in Helsinki in 2002, hemiboreal zone that had been planted either with Tilia x vulgaris or Alnus glutinosa f. pyramidalis were used as the study sites. Tree water use was measured from sap flow over the 2008-2011 period by the heat dissipation method. Penman-Monteith based evapotranspiration models of increasing complexity were tested against the tree water use measurements to assess the role of environmental and tree related factors in tree transpiration. Alnus and Tilia respectively used 1.1 and 0.8 l of water per m(2) of leaf area per day under ample water conditions, but the annual variation was high. The Penman-Monteith evapotranspiration estimate and soil water status changes explained over 80 % of the variation in tree transpiration when the model was parameterized annually. The addition of tree crown surface area in the model improved its accuracy and diminished variation between years and sites. Using single parameterization over all four years instead of annually varying one did not produce reliable estimates of tree transpiration. Tree transpiration, scaled to different canopy cover percentages, implied that the columnar Alnus trees could transpire as much as all annual rainfall at or less than 50 % canopy cover.

To better understand the poor regulation of water loss after leaf development at high relative air humidity (RH), the relative importance of the physiological and anatomical components was analyzed focusing on cultivars with a contrasting sensitivity to elevated RH. The stomatal responsiveness to three closing stimuli (desiccation, abscisic acid feeding, light/dark transition), as well as several stomatal features (density, index, size and pore dimensions) and the cuticular transpiration rate (CTR) were determined in four rose cultivars, grown under moderate (60%) and high (95%) RH. Moreover, the effects of changes in stomatal density and pore dimensions on the stomatal conductance (gs) were quantified using a modified version of the Brown and Escombe equation. Higher water loss, as a result of plant growth at high RH, was primarily caused by an increase in residual gs, and to a lesser extent due to higher CTR. It was estimated that in leaflets subjected to desiccation the enhanced gs in high RH- as compared to moderate RH-grown plants was mostly due to poor stomatal functionality and to a lesser extent the combined result of higher stomatal density and longer pore length. It is concluded that the reduced degree and, specially, the reduced rate of stomatal closure are the primary causes of the large genotypic variation in the control of water loss in high RH-grown plants. Furthermore, it was found that although changes in stomatal length have no influence on stomatal functionality, changed anatomical features per se represent a significant and direct contribution to the increased water loss.

The river valleys in Southwest China are characterized by a dry–hot climate and relatively rich soils, and host valley-type savannas that are dominated by deciduous species. However, little is known about the ecological adaptations of Chinese savanna plants to the local environments. We hypothesize that Chinese savanna species mainly possess a drought-avoiding strategy by having a deciduous leaf habit and have productive leaf traits. To test this hypothesis, we measured 26 anatomical, morphological, physiological, and chemical traits for 33 woody species from a valley savanna in Southwest China and compared them with the literature data of other dry and wet tropical tree species and a global dataset. We found that Chinese savanna species showed drought avoidance adaptations and exhibited productive leaf traits, such as thin and dense leaves with high ratio of palisade to spongy mesophyll, leaf nutrient concentrations and photosynthetic capacity. Correlations of photosynthetic capacity with N, P, and stomatal conductance across Chinese savanna species were consistent with global patterns reported for seed plants. However, the Chinese savanna species had consistently greater carbon gain at a given specific leaf area, N, P, and stomatal conductance, suggesting higher nutrient- and intrinsic water use efficiencies. These results suggest that paradoxically, Chinese savanna species are adapted to the stressful dry–hot valley habitat by having productive leaves.

Drainage-weighing lysimeters allowed monitoring of water balance components of non-bearing olive (Olea europaea cv Barnea) trees over a 3-month period including short-term events of controlled but severe water stress. The objective of the study was to evaluate a variety of soil and plant-based water status and drought stress monitoring methods on the basis of tree-scale evapotranspiration (ET). As the trees entered into and recovered from water stress, meteorological data, actual ET (ETa), soil water content and changes in leaf turgor pressure were continuously monitored. Additionally, midday measurements of stem water potential, stomatal conductance, canopy temperature, and quantum yield of PSII photochemistry were conducted. Diurnal (dawn to dusk) measurements of all the above were made hourly on days of maximum stress. Shoot elongation rate was measured for periods of stress and recovery. Quantum yield of PSII photochemistry, stomatal conductance, and stem water potential all successfully indicated reductions in whole-tree water consumption beginning at moderate stress levels. These measured parameters fully recovered to the levels of non-stressed trees soon after water application was renewed. Shoot elongation was reduced 25–30% for the 10-day period during and following drought and recovered thereafter to levels of non-stressed trees. Whole-tree ETa was reduced by as much as 20% even following full recovery of the leaf level parameters, suggesting reduced canopy size and growth due to the stress period. Non-destructive, continuous (turgor pressure) and remotely sensed (canopy temperature) methods showed promising potential for monitoring effects of water stress, in spite of technological and data interpretation challenges requiring further attention.

The existence of an efficient but also safe hydraulic system seems to be essential for plant survival under water limiting conditions. To investigate any common pattern in this safetyefficiency trade-off, static (xylem anatomy and vulnerability to xylem cavitation) and dynamic (K(plant), soil to leaf hydraulic conductance and (leaf), leaf water potential) hydraulic properties of nine Mediterranean species belonging to four functional groups (semi-deciduous malacophyllous, sclerophylls, deciduous and herbaceous) were studied across two altitude sites. Static parameters did not show any pattern, but a strong exponential relationship between K(plant) and (leaf) was evident for all the studied species. Furthermore, each species is represented by a different part and/or range of the K(plantleaf) relationship, indicative of its adaptive mechanisms and capacity for survival under water stress conditions. The use of K(plantleaf) relationship as a tool for understanding the mechanisms of plant responses to water stress is discussed. Tree Physiology

The negative effects of drought on plant growth, development of natural plant communities and crop productivity are well established, but some of the responses remain poorly characterized, particularly the effect of long-term drought on photosynthetic capacity. We hypothesized that long-term drought results in a decline in leaf photosynthetic capacity, and not just a decrease in diffusive conductance. To test this hypothesis, we studied the effect of drought, slowly developed over 3.5 months, in leaves of eight potted mango (Mangifera indica L.) trees. We found that photosynthesis was not only limited by stomatal closure, but was also downregulated as a consequence of a strong decrease in photosynthetic capacity assessed by the measurements of maximal net photosynthesis (Amax) and the light-saturated rate of photosynthetic electron transport (Jmax). The rapid recovery of Amax and Jmax, after only 1 week of rewatering, the maintenance of a stable pool of leaf nitrogen throughout the trial, and the decrease in quantum efficiency of open centers of photosystem II, indicate that the photosynthetic machinery escaped photodamage in the drought-treated trees and was simply downregulated during drought. The hexose-to-sucrose ratio was higher in leaves from drought-treated trees than in control leaves, suggesting that photosynthetic capacity decreased as a consequence of sink limitation. (Resume d'auteur)

Summary The extent to which climate change will affect evapotranspiration and water deficits is still uncertain. Temperature increase was recently shown to lead to enhanced drought in the Netherlands. In contrast, experimental evidence shows that elevated atmospheric CO2 concentrations tend to reduce stomatal opening in plants. This leads to lower transpiration rates, although models of atmospheric and soil water feedback show that reductions may be smaller than expected from stomatal closure. We combined the various effects and feedbacks. First, we inferred partial corrections on ‘crop factors’ used in simple evaporation equations such as Makkink’s, for a range of crops and vegetation types in the Netherlands. Second, we applied these corrected factors to infer the likely effects on water deficits in the Netherlands, using a coupled set of hydrological models and national climate scenarios. The combined effects of CO2 on evapotranspiration are generally modest, between a reduction of a few percent for short crops to about 15% for tall, rough vegetation. These reductions are, however, of comparable but opposite magnitude to predicted temperature-induced increases in evapotranspiration. We show that, if combined within the coupled hydrological model, the CO2-effect would lead to a much-reduced desiccating effect of climate change. In general, it is argued that, especially for sub-regional spatial scales and seasonal time-scales, CO2 is likely to be a significant factor in the water balance even of relatively wet regions.

The responses of water relations, stomatal conductance (gs) and growth parameters of tomato (Lycopersicon esculentum Mill. cv. Royesta) plants to nitrogen fertilisation and drought were studied. The plants were subjected to a long-term, moderate and progressive water stress by adding 80 % of the water evapotranspirated by the plant the preceding day. Well-watered plants received 100 % of the water evapotranspirated. Two weeks before starting the drought period, the plants were fertilised with Hoagland’s solution with 14, 60 and 110 mM NO3 − (N14, N60 and N110, respectively). Plants of the N110 treatment had the highest leaf area. However, gs was higher for N60 plants and lower for N110 plants. At the end of the drought period, N60 plants showed the lowest values of water potential (Ψw) and osmotic potential (Ψs), and the highest values of pressure potential (Ψp). N60 plants showed the highest Ψs at maximum Ψp and the highest bulk modulus of elasticity.

Considering the combined effects of CO2 fertilization and climate change drivers on plant physiology leads to a modest increase in simulated European forest transpiration in spite of the effects of CO2-induced stomatal closure. The Earth’s carbon and hydrologic cycles are intimately coupled by gas exchange through plant stomata1,2,3. However, uncertainties in the magnitude4,5,6 and consequences7,8 of the physiological responses9,10 of plants to elevated CO2 in natural environments hinders modelling of terrestrial water cycling and carbon storage11. Here we use annually resolved long-term δ13C tree-ring measurements across a European forest network to reconstruct the physiologically driven response of intercellular CO2 (Ci) caused by atmospheric CO2 (Ca) trends. When removing meteorological signals from the δ13C measurements, we find that trees across Europe regulated gas exchange so that for one ppmv atmospheric CO2 increase, Ci increased by ∼0.76 ppmv, most consistent with moderate control towards a constant Ci/Ca ratio. This response corresponds to twentieth-century intrinsic water-use efficiency (iWUE) increases of 14 ± 10 and 22 ± 6% at broadleaf and coniferous sites, respectively. An ensemble of process-based global vegetation models shows similar CO2 effects on iWUE trends. Yet, when operating these models with climate drivers reintroduced, despite decreased stomatal opening, 5% increases in European forest transpiration are calculated over the twentieth century. This counterintuitive result arises from lengthened growing seasons, enhanced evaporative demand in a warming climate, and increased leaf area, which together oppose effects of CO2-induced stomatal closure. Our study questions changes to the hydrological cycle, such as reductions in transpiration and air humidity, hypothesized to result from plant responses to anthropogenic emissions.

In order to understand the pathway involved in the chemical enhancement of photosynthetic rate, sodium bisulfite (NaHSO3) and benzyladenine (BA), a growth regulator, were applied to strawberry plants. The influence of these compounds on gas exchange and millisecond delayed light emission (ms-DLE) was investigated using 2-month-old plants. Results showed the net photosynthetic rate (A) in leaves was promoted by both NaHSO3 and BA. Stomatal conductance (g) and transpiration rate (E) were significantly increased only by BA, while intercellular CO2 concentration (Ci) was significantly decreased by NaHSO3. The enhancement of A by NaHSO3 and BA was only a short-term effect, lasting approximately 5 days for NaHSO3 and 30 h for BA. Plants treated with NaHSO3, BA or NaHSO3 + BA, showed no significant fluctuations in carboxylation efficiency (CE), photorespiration (RP) or dark respiration (RD). These results suggest that the influences of NaHSO3 and BA on gas exchange particularly A, could be via different mechanisms: the enhancement of A by the application of low concentrations of NaHSO3 appears to be associated with increased cyclic electron flow, while BA enhancement of A is at least partially due to increased g and/or E.

In coastal environments plants have to cope with various water sources: rainwater, water table, seawater, and mixtures. These are usually characterized by different isotopic signatures ( (18)O/ (16)O and D/H ratios). Xylem water reflects the isotopic compositions of the water sources. Additionally, water-use efficiency (WUE) can be assessed with carbon isotope discrimination (Delta) analyses. Gas exchange, Delta of leaf dry matter, and isotopic composition (delta (18)O) of xylem water were measured from June to August 2001 in herbaceous perennials of mobile dunes (Ammophila littoralis, Elymus farctus) and sclerophyllous shrubs and climbers (Arbutus unedo, Pistacia lentiscus, Phillyrea angustifolia, Qercus ilex, Juniperus oxycedrus, Smilax aspera) of consolidated dunes. Assimilation rates were rather low and did not show clear seasonal patterns, possibly due to limited precipitation and generally low values of stomatal conductance. The lowest values were shown in S. aspera. Different physiological patterns were found, on the basis of delta (18)O and Delta analyses. Values of delta (18)O of xylem water of phanerophytes were remarkably constant and matched those of the water table, indicating dependence on a reliable water source; values of Delta were relatively high, indicating low intrinsic WUE, with the exception of J. oxycedrus. Surprisingly, very high delta (18)O values were found for the xylem water from S. aspera in August. This suggests retrodiffusion of leaf water to xylem sap in the stem or direct uptake of water by leaves or stems, owing to dew or fog occurrence. Low Delta values indicated high WUE in S. aspera. Contrasting strategies were shown by the species of mobile dunes: E. farctus relied on superficial water and exhibited low WUE, accordingly to its therophyte-like vegetative cycle; on the contrary, A. littoralis used deeper water sources, showing higher WUE in relation to its long-lasting vegetative habit.

Predicting the large-scale consequences of drought in contrasting environments requires that we understand how drought effects differ among species originating from those environments. A previous meta-analysis of published experiments suggested that the effects of drought on both stomatal and non-stomatal limitations to photosynthesis may vary consistently among species from different hydroclimates. Here, we explicitly tested this hypothesis with two short-term water stress experiments on congeneric mesic and xeric species. One experiment was run in Australia using Eucalyptus species and the second was run in Spain using Quercus species as well as two more mesic species. In each experiment, plants were grown under moist conditions in a glasshouse, then deprived of water, and gas exchange was monitored. The stomatal response was analysed with a recently developed stomatal model, whose single parameter g1 represents the slope of the relationship between stomatal conductance and photosynthesis. The non-stomatal response was partitioned into effects on mesophyll conductance (gm), the maximum Rubisco activity (Vcmax) and the maximum electron transport rate (Jmax). We found consistency among the drought responses of g1, gm, Vcmax and Jmax, suggesting that drought imposes limitations on Rubisco activity and RuBP regeneration capacity concurrently with declines in stomatal and mesophyll conductance. Within each experiment, the more xeric species showed relatively high g1 under moist conditions, low drought sensitivity of g1, gm, Vcmax and Jmax, and more negative values of the critical pre-dawn water potential at which Vcmax declines most steeply, compared with the more mesic species. These results indicate adaptive interspecific differences in drought responses that allow xeric tree species to continue transpiration and photosynthesis for longer during periods without rain.

How plants respond to climate change is of major concern, as plants will strongly impact future ecosystem functioning, food production and climate. Here we investigate how vegetation structure and functioning may be influenced by predicted increases in annual temperatures and atmospheric CO 2 concentration, and modeled the extent to which local plant-plant interactions may modify these effects. A canopy model was developed, which calculates photosynthesis as a function of light, nitrogen, temperature, CO 2 and water availability, and considers different degrees of light competition between neighboring plants through canopy mixing; soybean ( Glycine max ) was used as a reference system. The model predicts increased net photosynthesis and reduced stomatal conductance and transpiration under atmospheric CO 2 increase. When CO 2 elevation is combined with warming, photosynthesis is increased more, but transpiration is reduced less. Intriguingly, when competition is considered the optimal response shifts to producing larger leaf areas, but with lower stomatal conductance and associated vegetation transpiration than when competition is not considered. Furthermore, only when competition is considered are the predicted effects of elevated CO 2 on LAI well within the range of observed effects obtained by Free Air CO 2 Enrichment (FACE) experiments. Together, our results illustrate how competition between plants may modify vegetation responses to climate change.

Current understanding of physiological mechanisms governing stomatal behavior under water stress conditions is still incomplete and controversial. It has been proposed that coordination of stomatal kinetics with xylem vulnerability to cavitation [vulnerability curve (VC)] leads to different levels of isohydry/anisohydry in different plant species/cultivars. In this study, this hypothesis is tested in Vitis vinifera cultivars displaying contrasting stomatal behavior under drought stress. The cv Montepulciano (MP, near-isohydric) and Sangiovese (SG, anisohydric) were compared in terms of stomatal response to leaf and stem water potential, as possibly correlated to different petiole hydraulic conductivity (k(petiole)) and VC, as well as to leaf water relations parameters. MP leaves showed almost complete stomatal closure at higher leaf and stem water potentials than SG leaves. Moreover, MP petioles had higher maximum k(petiole) and were more vulnerable to cavitation than SG. Water potential at the turgor loss point was higher in MP than in SG. In SG, the percentage reduction of stomatal conductance (PLg(s)) under water stress was almost linearly correlated with corresponding percentage loss of k(petiole) (PLC), while in MP PLg(s) was less influenced by PLC. Our results suggest that V. vinifera near-isohydric and anisohydric genotypes differ in terms of xylem vulnerability to cavitation as well as in terms of k(petiole) and that the coordination of these traits leads to their different stomatal responses under water stress conditions.

The photosynthetic response was studied in two clones (Populus deltoides × maximowiczii Eridano and Populus × euramericana I-214), known for their differential response to ozone (O3) in terms of visible symptoms, when exposed to O3 (60 nl l−1 5 h day−1, 7 and 15 days). The photosynthetic ability was tested using gas exchange and chlorophyll fluorescence analysis. O3 caused a decrease in the CO2 assimilation rate at light saturation level in mature leaves of both clones. Alterations of Chl fluorescence parameters, in particular the Fv/Fm ratio and non-photochemical quenching were also observed. The effects were similar for both clones and it could not be concluded that differential effects on electron transport capacity were responsible for the observed reduction in photosynthesis. The reduction of photosynthetic rate in Eridano was due mainly to a reduced mesophyll activity, as evidenced by the increase in intercellular CO2 concentration and the minimal changes in stomatal conductance. In contrast, in I-214, stomatal effects were primarily responsible, although effects on the mesophyll cannot be excluded. Data obtained indicate that the effects observed at the mesophyll level may be attributed to indirect effects caused by membrane disorders.

Aims: To evaluate the impact of the amount and distribution of soil water on xylem anatomy and xylem hydraulics of current-year shoots, plant water status and stomatal conductance of mature 'Manzanilla' olive trees. Methods: Measurements of water potential, stomatal conductance, hydraulic conductivity, vulnerability to embolism, vessel diameter distribution and vessel density were made in trees under full irrigation with non-limiting soil water conditions, localized irrigation, and rain-fed conditions. Results: All trees showed lower stomatal conductance values in the afternoon than in the morning. The irrigated trees showed water potential values around -1.4 and -1.6 MPa whereas the rain-fed trees reached lower values. All trees showed similar specific hydraulic conductivity (K s) and loss of conductivity values during the morning. In the afternoon, K s of rain-fed trees tended to be lower than of irrigated trees. No differences in vulnerability to embolism, vessel-diameter distribution and vessel density were observed between treatments. Conclusions: A tight control of stomatal conductance was observed in olive which allowed irrigated trees to avoid critical water potential values and keep them in a safe range to avoid embolism. The applied water treatments did not influence the xylem anatomy and vulnerability to embolism of current-year shoots of mature olive trees., This project was funding both by the Spanish Ministry of Science and Innovation (research project AGL2006-04666/AGR) and the Austrian Science Fund (FWF) (project No. P20852-B16). J. M. Torres Ruiz held a doctoral fellowship from the Spanish Ministry of Science and Innovation (BES-2007-17149, MCINN).

The rate of photosynthesis (A) of plants exposed to water deficit is a function of stomatal (gs) and mesophyll (gm) conductance determining the availability of CO2 at the site of car- boxylation within the chloroplast. Mesophyll conductance often represents the greatest impediment to photosynthetic uptake of CO2, and a crucial determinant of the photosyn- thetic effects of drought. Abscisic acid (ABA) plays a fundamental role in signalling and co- ordination of plant responses to drought; however, the effect of ABA on gm is not well- defined. Rose, cherry, olive and poplar were exposed to exogenous ABA and their leaf gas exchange parameters recorded over a four hour period. Application with ABA induced reductions in values of A, gs and gm in all four species. Reduced gm occurred within one hour of ABA treatment in three of the four analysed species; indicating that the effect of ABA on gm occurs on a shorter timescale than previously considered. These declines in gm val- ues associated with ABA were not the result of physical changes in leaf properties due to altered turgor affecting movement of CO2, or caused by a reduction in the sub-stomatal con- centration of CO2 (Ci). Increased [ABA] likely induces biochemical changes in the properties of the interface between the sub-stomatal air-space and mesophyll layer through the actions of cooporins to regulate the transport of CO2. The results of this study provide further evi- dence that gm is highly responsive to fluctuations in the external environment, and stress signals such as ABA induce co-ordinated modifications of both gs and gm in the regulation of photosynthesis.

Estimating leaf stomatal conductance for CO 2 diffusion ( g sc ) is pivotal for further estimation of crop transpiration as well as energy and mass balances between air and plant in greenhouses. In this study, we tested two models, i.e. the Jarvis model and a new version of BWB–Leuning model (BWB–Leuning–Yin model), for estimating g sc in four greenhouse crops (cucumber, chrysanthemum, tulip and lilium), using data from extensive experiments conducted under a wide range of environmental conditions from 2003 to 2010. The models were parameterized from a subset of the experimental data. The remaining data sets for model validation were classified into two groups: group one was from experiments conducted during the similar seasons at the same sites as those for model parameterization, whereas group two was from experiments in different seasons and sites. When using data of group one, both models gave satisfactory estimations of g sc under both ample water supply and water stress conditions. When using data of group two, the BWB–Leuning–Yin model gave better estimations of g sc than the Jarvis model did. Functions or parameters of the Jarvis model, if applied to independent environmental conditions, have to be re-derived. Compared to its earlier versions, the BWB–Leuning–Yin model gave better estimated g sc under low light intensities and low CO 2 concentrations, suggesting that the new version is especially suitable for estimation of g sc for crops grown in low-investment greenhouses where levels of light and CO 2 are lower than ambient levels. In addition, analytical algorithms for the coupled FvCB model and diffusional conductance as adopted in the BWB–Leuning–Yin model make parameterization and simultaneous estimation of net photosynthetic rate and g sc an easy task, using readily obtained information.

Due to anthropogenic influences, solar UV-B irradiance at the earth’s surface is increasing. To determine the effects of enhanced UV-B radiation on photosynthetic characteristics of Prunus dulcis, two-year-old seedlings of the species were submitted to four levels of UV-B stress, namely 0 (UV-Bc), 4.42 (UV-B1), 7.32 (UV-B2) and 9.36 (UV-B3) kJ m−2 d−1. Effects of UV-B stress on a range of chlorophyll (Chl) fluorescence parameters (FPs), Chl contents and photosynthetic gas-exchange parameters were investigated. UV-B stress promoted an increase in minimal fluorescence of dark-adapted state (F0) and F0/Fm, and a decrease in variable fluorescence (Fv, Fv/Fm, Fv/F0 and F0/Fm) due to its adverse effects on photosystem II (PSII) activity. No significant change was observed for maximal fluorescence of dark-adapted state (Fm). Enhanced UV-B radiation caused a significant inhibition of net photosynthetic rate (P N) at UV-B2 and UV-B3 levels and this was accompanied by a reduction in stomatal conductance (g s) and transpiration rate (E). The contents of Chl a, b, and total Chl content (a+b) were also significantly reduced at increased UV-B stress. In general, adverse UV-B effects became significant at the highest tested radiation dose 9.36 kJ m−2 d−1. The most sensitive indicators for UV-B stress were Fv/F0, Chl a content and P N. Significant P

Hydrological models featuring root water uptake usually do not include compensation mechanisms such that reductions in uptake from dry layers are compensated by an increase in uptake from wetter layers. We developed a physically based root water uptake model with an implicit compensation mechanism.Hydrological models featuring root water uptake usually do not include compensation mechanisms such that reductions in uptake from dry layers are compensated by an increase in uptake from wetter layers. We developed a physically based root water uptake model with an implicit compensation mechanism. Based on an expression for the matric flux potential (M) as a function of the distance to the root, and assuming a depth-independent value of M at the root surface, uptake per layer is shown to be a function of layer bulk M, root surface M, and a weighting factor that depends on root length density and root radius. Actual transpiration can be calculated from the sum of layer uptake rates. The proposed reduction function (PRF) was built into the SWAP model, and predictions were compared to those made with the Feddes reduction function (FRF). Simulation results were tested against data from Canada (continuous spring wheat [(Triticum aestivum L.]) and Germany (spring wheat, winter barley [Hordeum vulgare L.], sugarbeet [Beta vulgaris L.], winter wheat rotation). For the Canadian data, the root mean square error of prediction (RMSEP) for water content in the upper soil layers was very similar for FRF and PRF; for the deeper layers, RMSEP was smaller for PRF. For the German data, RMSEP was lower for PRF in the upper layers and was similar for both models in the deeper layers. In conclusion, but dependent on the properties of the data sets available for testing,the incorporation of the new reduction function into SWAP was successful, providing new capabilities for simulating compensated root water uptake without increasing the number of input parameters or degrading model performance.

Photosynthesis rates and photosynthesis-leaf nutrient relationships were analysed in nine tropical grass and sedge species growing in three different ecosystems: a rain-fed grassland, a seasonal floodplain, and a permanent swamp, located along a hydrological gradient in the Okavango Delta, Botswana. These investigations were conducted during the rainy season, at a time of the year when differences in growth conditions between the sites were relatively uniform. At the permanent swamp, the largest variations were found for area-based leaf nitrogen contents, from 20 mmol m(-2) to 140 mmol m(-2), nitrogen use efficiencies (NUE), from 0.2 mmol (C) mol(-1) (N) s(-1) to 2.0 mmol (C) mol(-1) (N) s(-1), and specific leaf areas (SLA), from 50 cm(2) g(-1) to 400 cm(2) g(-1). For the vegetation growing at the rainfed grassland, the highest leaf gas exchange rates, high leaf nutrient levels, a low ratio of intercellular to ambient CO2 concentration, and high carboxylation efficiency were found. Taken together, these observations indicate a very efficient growth strategy that is required for survival and reproduction during the relatively brief period of water availability. The overall lowest values of light-saturated photosynthesis (A(sat)) were observed at the seasonal floodplain; around 25 mu mol m(-2) s(-1) and 30 mu mol m(-2) s(-1). To place these observations into the broader context of functional leaf trait analysis, relationships of photosynthesis rates, specific leaf area, and foliar nutrient levels were plotted, in the same way as was done for previously published 'scaling relationships' that are based largely on C-3 plants, noting the differences in the analyses between this study and the previous study. The within-and across-species variation in both A(sat) and SLA appeared better predicted by foliar phosphorus content (dry mass or area basis) rather than by foliar nitrogen concentrations, possibly because the availability of phosphorus is even more critical than the availability of nitrogen in the studied relatively oligotrophic ecosystems. [References: 49]

The global carbon and water cycles are governed by the coupling of CO2 and water vapour exchanges through the leaves of terrestrial plants, controlled by plant adaptations to balance carbon gains and hydraulic risks. We introduce a trait-based optimality theory that unifies the treatment of stomatal responses and biochemical acclimation of plants to environments changing on multiple timescales. Tested with experimental data from 18 species, our model successfully predicts the simultaneous decline in carbon assimilation rate, stomatal conductance and photosynthetic capacity during progressive soil drought. It also correctly predicts the dependencies of gas exchange on atmospheric vapour pressure deficit, temperature and CO2. Model predictions are also consistent with widely observed empirical patterns, such as the distribution of hydraulic strategies. Our unified theory opens new avenues for reliably modelling the interactive effects of drying soil and rising atmospheric CO2 on global photosynthesis and transpiration., Nature Plants, 8 (11), ISSN:2055-026X, ISSN:2055-0278

The rate of glucose equivalent production in C(4) green plants is investigated as a function of the intercellular partial pressure of CO(2), so as to find the precise physical chemistry of photosynthesis. Expressions are first formulated for the dependence of photochemical efficiency and of rubisco activation on pressure. Then a pressure-dependent rate law is derived. The latter is successfully tested for two specific C(4) plants, namely, Panicum antidotale and Panicum coloratum.