Photosynthetic responses of 13 grassland species across 11 years of free‐air CO2 enrichment is modest,consistent and independent of N supply

If long‐term responses of photosynthesis and leaf diffusive conductance to rising atmospheric carbon dioxide (CO₂) levels are similar or predictably different among species, functional types, and ecosystem types, general global models of elevated CO₂ effects can effectively be developed. To address this issue we measured gas exchange rates of 13 perennial grassland species from four functional groups across 11 years of long‐term free‐air CO₂ enrichment (eCO₂, +180 ppm above ambient CO₂) in the BioCON experiment in Minnesota, USA. Eleven years of eCO₂ produced consistent but modest increases in leaf net photosynthetic rates of 10% on average compared with plants grown at ambient CO₂ concentrations across the 13 species. This eCO₂‐induced enhancement did not depend on soil N treatment, is much less than the average across other longer‐term studies, and represents strong acclimation (i.e. downregulation) as it is also much less than the instantaneous response to eCO₂. The legume and C3 nonlegume forb species were the most responsive among the functional groups (+13% in each), the C4 grasses the least responsive (+4%), and C3 grasses intermediate in their photosynthetic response to eCO₂ across years (+9%). Leaf stomatal conductance and nitrogen content declined comparably across species in eCO₂ compared with ambient CO₂ and to degrees corresponding to results from other studies. The significant acclimation of photosynthesis is explained in part by those eCO₂‐induced decreases in leaf N content and stomatal conductance that reduce leaf photosynthetic capacity in plants grown under elevated compared with ambient CO₂ concentrations. Results of this study, probably the longest‐term with the most species, suggest that carbon cycle models that assume and thereby simulate long‐lived strong eCO₂ stimulation of photosynthesis (e.g.> 25%) for all of Earth's terrestrial ecosystems should be viewed with a great deal of caution.     

Gas exchange and photosynthetic acclimation over subambient to elevated CO2 in a C3–C4 grassland

Atmospheric CO₂ (C_a) has risen dramatically since preglacial times and is projected to double in the next century. As part of a 4‐year study, we examined leaf gas exchange and photosynthetic acclimation in C₃ and C₄ plants using unique chambers that maintained a continuous C_a gradient from 200 to 550 µmol mol^?1 in a natural grassland. Our goals were to characterize linear, nonlinear and threshold responses to increasing C_a from past to future C_a levels. Photosynthesis (A), stomatal conductance (g_s), leaf water‐use efficiency (A/g_s) and leaf N content were measured in three common species: Bothriochloa ischaemum, a C₄ perennial grass, Bromus japonicus, a C₃ annual grass, and Solanum dimidiatum, a C₃ perennial forb. Assimilation responses to internal CO₂ concentrations (A/C_i curves) and photosynthetically active radiation (A/PAR curves) were also assessed, and acclimation parameters estimated from these data. Photosynthesis increased linearly with C_a in all species (P < 0.05). S. dimidiatum and B. ischaemum had greater carboxylation rates for Rubisco and PEP carboxylase, respectively, at subambient than superambient C_a (P < 0.05). To our knowledge, this is the first published evidence of A up‐regulation at subambient C_a in the field. No species showed down‐regulation at superambient C_a. Stomatal conductance generally showed curvilinear decreases with C_a in the perennial species (P < 0.05), with steeper declines over subambient C_a than superambient, suggesting that plant water relations have already changed significantly with past C_a increases. Resource‐use efficiency (A/g_s and A/leaf N) in all species increased linearly with C_a. As both C₃ and C₄ plants had significant responses in A, g_s, A/g_s and A/leaf N to C_a enrichment, future C_a increases in this grassland may not favour C₃ species as much as originally thought. Non‐linear responses and acclimation to low C_a should be incorporated into mechanistic models to better predict the effects of past and present rising C_a on grassland ecosystems.     

Canopy radiation‐ and water‐use efficiencies as affected by elevated [CO2]

This study used an environmentally controlled plant growth facility, EcoCELLs, to measure canopy gas exchanges directly and to examine the effects of elevated [CO₂] on canopy radiation‐ and water‐use efficiencies. Sunflowers (Helianthus annus var. Mammoth) were grown at ambient (399 μmol mol^?1) and elevated [CO₂] (746 μmol mol^?1) for 53 days in EcoCELLs. Whole canopy carbon‐ and water‐fluxes were measured continuously during the period of the experiment. The results indicated that elevated [CO₂] enhanced daily total canopy carbon‐ and water‐fluxes by 53% and 11%, respectively, on a ground‐area basis, resulting in a 54% increase in radiation‐use efficiency (RUE) based on intercepted photosynthetic active radiation and a 26% increase in water‐use efficiency (WUE) by the end of the experiment. Canopy carbon‐ and water‐fluxes at both CO₂ treatments varied with canopy development. They were small at 22 days after planting (DAP) and gradually increased to the maxima at 46 DAP. When canopy carbon‐ and water‐fluxes were expressed on a leaf‐area basis, no effect of CO₂ was found for canopy water‐flux while elevated [CO₂] still enhanced canopy carbon‐flux by 29%, on average. Night‐time canopy carbon‐flux was 32% higher at elevated than at ambient [CO₂]. In addition, RUE and WUE displayed strong diurnal variations, high at noon and low in the morning or afternoon for WUE but opposite for RUE. This study provided direct evidence that plant canopy may consume more, instead of less, water but utilize both water and radiation more efficiently at elevated than at ambient [CO₂], at least during the exponential growth period as illustrated in this experiment.     

Nutrient supply enhanced the increase in intrinsic water‐use efficiency of a temperate seminatural grassland in the last century

Under the increase in atmospheric CO₂ during the last century, variable increases in the intrinsic water‐use efficiency (W_i), i.e., the ratio between carbon assimilation rate (A) and stomatal conductance (g_s), of C₃ vegetation have been observed. Here, we ask if long‐term nutrient status and especially nitrogen supply have an effect on the CO₂ response of W_i in a temperate seminatural C₃ grassland. This analysis draws on the long‐term trends (1915–2009) in W_i, derived from carbon isotope analysis, of archived hay and herbage from the Park Grass Experiment at Rothamsted (South‐East England). Plant samples came from five fertilizer treatments, each with different annual nitrogen (N; 0, 48 or 96 kg ha^?1), phosphorus (P; 0 or 35 kg ha^?1) and potassium (K; 0 or 225 kg ha^?1) applications, with lime as required to maintain soil pH near 7. Carbon isotope discrimination (¹³Δ) increased significantly (P < 0.001) on the Control (0.9‰ per 100 ppm CO₂ increase). This trend differed significantly (P < 0.01) from those observed on the fertilized treatments (PK only: 0.4‰ per 100 ppm CO₂ increase, P < 0.001; Low N only, Low N+PK, High N+PK: no significant increase). The ¹³Δ trends on fertilized treatments did not differ significantly from each other. However, N status, assessed as N fertilizer supply plus an estimate of biologically fixed N, was negatively related (r² = 0.88; P < 0.02) to the trend for ¹³Δ against CO₂. Other indices of N status exhibited similar relationships. Accordingly, the increase in W_i at High N+PK was twice that of the Control (+28% resp. +13% relative to 1915). In addition, the CO₂ responsiveness of ¹³Δ was related to the grass content of the plant community. This may have been due to the greater CO₂ responsiveness of g_s in grasses relative to forbs. Thus, the greater CO₂ response of grass‐rich fertilized swards may be related to effects of nutrient supply on botanical composition.     

Last‐century changes of alpine grassland water‐use efficiency: a reconstruction through carbon isotope analysis of a time‐series of Capra ibex horns

The ecophysiological response of an alpine grassland to recent climate change and increasing atmospheric CO₂ concentration was investigated with a new strategy to go back in time: using a time‐series of Capra ibex horns as archives of the alpine grasslands' carbon isotope discrimination (¹³Δ). From the collection of the Natural History Museum of Bern, horns of 24 males from the population of the Augstmatthorn–Brienzer Rothorn mountains, Switzerland, were sampled covering the period from 1938 to 2006. Samples were taken from the beginning of each year‐ring of the horns, representing the beginning of the horn growth period, the spring. The horns' carbon ¹³C content (Δ¹³C) declined together with that of atmospheric CO₂ over the 69‐year period, but ¹³Δ increased slightly (+0.4‰), though significantly (P<0.05), over the observation period. Estimated intercellular CO₂ concentration increased (+56 μmol mol^?1) less than the atmospheric CO₂ concentration (+81 μmol mol^?1), so that intrinsic water‐use efficiency increased by 17.8% during the 69‐year period. However, the atmospheric evaporative demand at the site increased by approximately 0.1 kPa between 1955 and 2006, thus counteracting the improvement of intrinsic water‐use efficiency. As a result, instantaneous water‐use efficiency did not change. The observed changes in intrinsic water‐use efficiency were in the same range as those of trees (as reported by others), indicating that leaf‐level control of water‐use efficiency of grassland and forests followed the same principles. This is the first reconstruction of the water‐use efficiency response of a natural grassland ecosystem to last century CO₂ and climatic changes. The results indicate that the alpine grassland community has responded to climate change by improving the physiological control of carbon gain to water loss, following the increases in atmospheric CO₂ and evaporative demand. But, effective leaf‐level water‐use efficiency has remained unchanged.     

Chloride as a macronutrient increases water‐use efficiency by anatomically driven reduced stomatal conductance and increased mesophyll diffusion to CO2

Chloride (Cl^?) has been recently described as a beneficial macronutrient, playing specific roles in promoting plant growth and water‐use efficiency (WUE). However, it is still unclear how Cl^? could be beneficial, especially in comparison with nitrate (NO₃^?), an essential source of nitrogen that shares with Cl^? similar physical and osmotic properties, as well as common transport mechanisms. In tobacco plants, macronutrient levels of Cl^? specifically reduce stomatal conductance (g_s) without a concomitant reduction in the net photosynthesis rate (A_N). As stomata‐mediated water loss through transpiration is inherent in the need of C₃ plants to capture CO₂, simultaneous increase in photosynthesis and WUE is of great relevance to achieve a sustainable increase in C₃ crop productivity. Our results showed that Cl^?‐mediated stimulation of larger leaf cells leads to a reduction in stomatal density, which in turn reduces g_s and water consumption. Conversely, Cl^? improves mesophyll diffusion conductance to CO₂ (g_m) and photosynthetic performance due to a higher surface area of chloroplasts exposed to the intercellular airspace of mesophyll cells, possibly as a consequence of the stimulation of chloroplast biogenesis. A key finding of this study is the simultaneous improvement of A_N and WUE due to macronutrient Cl^? nutrition. This work identifies relevant and specific functions in which Cl^? participates as a beneficial macronutrient for higher plants, uncovering a sustainable approach to improve crop yield.     

Maintenance of leaf N controls the photosynthetic CO2 response of grassland species exposed to 9 years of free‐air CO2 enrichment

Determining underlying physiological patterns governing plant productivity and diversity in grasslands are critical to evaluate species responses to future environmental conditions of elevated CO₂ and nitrogen (N) deposition. In a 9‐year experiment, N was added to monocultures of seven C₃ grassland species exposed to elevated atmospheric CO₂ (560 μmol CO₂ mol^?1) to evaluate how N addition affects CO₂ responsiveness in species of contrasting functional groups. Functional groups differed in their responses to elevated CO₂ and N treatments. Forb species exhibited strong down‐regulation of leaf N_mass concentrations (?26%) and photosynthetic capacity (?28%) in response to elevated CO₂, especially at high N supply, whereas C₃ grasses did not. Hence, achieved photosynthetic performance was markedly enhanced for C₃ grasses (+68%) in elevated CO₂, but not significantly for forbs. Differences in access to soil resources between forbs and grasses may distinguish their responses to elevated CO₂ and N addition. Forbs had lesser root biomass, a lower distribution of biomass to roots, and lower specific root length than grasses. Maintenance of leaf N, possibly through increased root foraging in this nutrient‐poor grassland, was necessary to sustain stimulation of photosynthesis under long‐term elevated CO₂. Dilution of leaf N and associated photosynthetic down‐regulation in forbs under elevated [CO₂], relative to the C₃ grasses, illustrates the potential for shifts in species composition and diversity in grassland ecosystems that have significant forb and grass components.     

Interactive effects of elevated CO2, warming,reduced rainfall,and nitrogen on leaf gas exchange in five perennial grassland species

Global changes can interact to affect photosynthesis and thus ecosystem carbon capture, yet few multi-factor field studies exist to examine such interactions. Here, we evaluate leaf gas exchange responses of five perennial grassland species from four functional groups to individual and interactive global changes in an open-air experiment in Minnesota, USA, including elevated CO₂ (eCO₂), warming, reduced rainfall and increased soil nitrogen supply. All four factors influenced leaf net photosynthesis and/or stomatal conductance, but almost all effects were context-dependent, i.e. they differed among species, varied with levels of other treatments and/or depended on environmental conditions. Firstly, the response of photosynthesis to eCO₂ depended on species and nitrogen, became more positive as vapour pressure deficit increased and, for a C₄ grass and a legume, was more positive under reduced rainfall. Secondly, reduced rainfall increased photosynthesis in three functionally distinct species, potentially via acclimation to low soil moisture. Thirdly, warming had positive, neutral or negative effects on photosynthesis depending on species and rainfall. Overall, our results show that interactions among global changes and environmental conditions may complicate predictions based on simple theoretical expectations of main effects, and that the factors and interactions influencing photosynthesis vary among herbaceous species.     

Long‐term elevated air [CO2] strengthens photosynthetic functioning and mitigates the impact of supra‐optimal temperatures in tropical Coffea arabica and C. canephora species

The tropical coffee crop has been predicted to be threatened by future climate changes and global warming. However, the real biological effects of such changes remain unknown. Therefore, this work aims to link the physiological and biochemical responses of photosynthesis to elevated air [CO₂] and temperature in cultivated genotypes of Coffea arabica L. (cv. Icatu and IPR108) and Coffea canephora cv. Conilon CL153. Plants were grown for ca. 10 months at 25/20 °C (day/night) and 380 or 700 μl CO₂ l^?1 and then subjected to temperature increase (0.5 °C day^?1) to 42/34 °C. Leaf impacts related to stomatal traits, gas exchanges, C isotope composition, fluorescence parameters, thylakoid electron transport and enzyme activities were assessed at 25/20, 31/25, 37/30 and 42/34 °C. The results showed that (1) both species were remarkably heat tolerant up to 37/30 °C, but at 42/34 °C a threshold for irreversible nonstomatal deleterious effects was reached. Impairments were greater in C. arabica (especially in Icatu) and under normal [CO₂]. Photosystems and thylakoid electron transport were shown to be quite heat tolerant, contrasting to the enzymes related to energy metabolism, including RuBisCO, which were the most sensitive components. (2) Significant stomatal trait modifications were promoted almost exclusively by temperature and were species dependent. Elevated [CO₂], (3) strongly mitigated the impact of temperature on both species, particularly at 42/34 °C, modifying the response to supra‐optimal temperatures, (4) promoted higher water‐use efficiency under moderately higher temperature (31/25 °C) and (5) did not provoke photosynthetic downregulation. Instead, enhancements in [CO₂] strengthened photosynthetic photochemical efficiency, energy use and biochemical functioning at all temperatures. Our novel findings demonstrate a relevant heat resilience of coffee species and that elevated [CO₂] remarkably mitigated the impact of heat on coffee physiology, therefore playing a key role in this crop sustainability under future climate change scenarios.     

Change in terrestrial ecosystem water‐use efficiency over the last three decades

Defined as the ratio between gross primary productivity (GPP) and evapotranspiration (ET), ecosystem‐scale water‐use efficiency (EWUE) is an indicator of the adjustment of vegetation photosynthesis to water loss. The processes controlling EWUE are complex and reflect both a slow evolution of plants and plant communities as well as fast adjustments of ecosystem functioning to changes of limiting resources. In this study, we investigated EWUE trends from 1982 to 2008 using data‐driven models derived from satellite observations and process‐oriented carbon cycle models. Our findings suggest positive EWUE trends of 0.0056, 0.0007 and 0.0001 g C m^?2 mm^?1 yr^?1 under the single effect of rising CO₂ (‘CO₂’), climate change (‘CLIM’) and nitrogen deposition (‘NDEP’), respectively. Global patterns of EWUE trends under different scenarios suggest that (i) EWUE‐CO₂ shows global increases, (ii) EWUE‐CLIM increases in mainly high latitudes and decreases at middle and low latitudes, (iii) EWUE‐NDEP displays slight increasing trends except in west Siberia, eastern Europe, parts of North America and central Amazonia. The data‐driven MTE model, however, shows a slight decline of EWUE during the same period (?0.0005 g C m^?2 mm^?1 yr^?1), which differs from process‐model (0.0064 g C m^?2 mm^?1 yr^?1) simulations with all drivers taken into account. We attribute this discrepancy to the fact that the nonmodeled physiological effects of elevated CO₂ reducing stomatal conductance and transpiration (TR) in the MTE model. Partial correlation analysis between EWUE and climate drivers shows similar responses to climatic variables with the data‐driven model and the process‐oriented models across different ecosystems. Change in water‐use efficiency defined from transpiration‐based WUE_t (GPP/TR) and inherent water‐use efficiency (IWUE_t, GPP×VPD/TR) in response to rising CO₂, climate change, and nitrogen deposition are also discussed. Our analyses will facilitate mechanistic understanding of the carbon–water interactions over terrestrial ecosystems under global change.     

A dynamic leaf gas‐exchange strategy is conserved in woody plants under changing ambient CO2: evidence from carbon isotope discrimination in paleo and CO2 enrichment studies

Rising atmospheric [CO₂], c_a, is expected to affect stomatal regulation of leaf gas‐exchange of woody plants, thus influencing energy fluxes as well as carbon (C), water, and nutrient cycling of forests. Researchers have proposed various strategies for stomatal regulation of leaf gas‐exchange that include maintaining a constant leaf internal [CO₂], c_i, a constant drawdown in CO₂ (c_a ? c_i), and a constant c_i/c_a. These strategies can result in drastically different consequences for leaf gas‐exchange. The accuracy of Earth systems models depends in part on assumptions about generalizable patterns in leaf gas‐exchange responses to varying c_a. The concept of optimal stomatal behavior, exemplified by woody plants shifting along a continuum of these strategies, provides a unifying framework for understanding leaf gas‐exchange responses to c_a. To assess leaf gas‐exchange regulation strategies, we analyzed patterns in c_i inferred from studies reporting C stable isotope ratios (δ¹³C) or photosynthetic discrimination (?) in woody angiosperms and gymnosperms that grew across a range of c_a spanning at least 100 ppm. Our results suggest that much of the c_a‐induced changes in c_i/c_a occurred across c_a spanning 200 to 400 ppm. These patterns imply that c_a ? c_i will eventually approach a constant level at high c_a because assimilation rates will reach a maximum and stomatal conductance of each species should be constrained to some minimum level. These analyses are not consistent with canalization toward any single strategy, particularly maintaining a constant c_i. Rather, the results are consistent with the existence of a broadly conserved pattern of stomatal optimization in woody angiosperms and gymnosperms. This results in trees being profligate water users at low c_a, when additional water loss is small for each unit of C gain, and increasingly water‐conservative at high c_a, when photosystems are saturated and water loss is large for each unit C gain.     

Seasonal responses of terrestrial ecosystem water‐use efficiency to climate change

Ecosystem water‐use efficiency (EWUE) is an indicator of carbon–water interactions and is defined as the ratio of carbon assimilation (GPP) to evapotranspiration (ET). Previous research suggests an increasing long‐term trend in annual EWUE over many regions and is largely attributed to the physiological effects of rising CO₂. The seasonal trends in EWUE, however, have not yet been analyzed. In this study, we investigate seasonal EWUE trends and responses to various drivers during 1982–2008. The seasonal cycle for two variants of EWUE, water‐use efficiency (WUE, GPP/ET), and transpiration‐based WUE (WUE_t, the ratio of GPP and transpiration), is analyzed from 0.5° gridded fields from four process‐based models and satellite‐based products, as well as a network of 63 local flux tower observations. WUE derived from flux tower observations shows moderate seasonal variation for most latitude bands, which is in agreement with satellite‐based products. In contrast, the seasonal EWUE trends are not well captured by the same satellite‐based products. Trend analysis, based on process‐model factorial simulations separating effects of climate, CO₂, and nitrogen deposition (NDEP), further suggests that the seasonal EWUE trends are mainly associated with seasonal trends of climate, whereas CO₂ and NDEP do not show obvious seasonal difference in EWUE trends. About 66% grid cells show positive annual WUE trends, mainly over mid‐ and high northern latitudes. In these regions, spring climate change has amplified the effect of CO₂ in increasing WUE by more than 0.005 gC m⁻² mm⁻¹ yr⁻¹ for 41% pixels. Multiple regression analysis further shows that the increase in springtime WUE in the northern hemisphere is the result of GPP increasing faster than ET because of the higher temperature sensitivity of GPP relative to ET. The partitioning of annual EWUE to seasonal components provides new insight into the relative sensitivities of GPP and ET to climate, CO_2, and NDEP.     

Intrinsic water‐use efficiency of temperate seminatural grassland has increased since 1857: an analysis of carbon isotope discrimination of herbage from the Park Grass Experiment

A 150‐year‐long record of intrinsic water‐use efficiency (W_i) was derived from community‐level carbon isotope discrimination (¹³Δ) in the herbage of the unfertilized, unlimed control treatment (plot 3) of the Park Grass Experiment at Rothamsted (England) between 1857 and 2007. ¹³Δ during spring growth (first cut harvested in June) averaged 21.0‰ (±0.5‰ SD) and has not shown a long‐term trend (P=0.5) since 1857. ¹³Δ of summer/autumn growth (second cut harvested between September and November) increased from 21.3‰ to 22.0‰ (P < 0.001) between 1875 and 2007. W_i during spring growth has therefore increased by 33% since the beginning of the experiment, and W_i of summer/autumn growth has increased by 18%. The variation in ¹³Δ was mainly related to weather conditions. Plant available soil water explained 51% and 40% of the variation in spring growth ¹³Δ and summer/autumn growth ¹³Δ, respectively. In the 1857–2007 period yields have not increased, suggesting that community‐level photosynthesis has not increased either. Therefore, the increased W_i probably resulted from a decreased stomatal conductance. Vapour pressure deficit (VPD) during spring growth (March–June) has not changed since 1915, meaning that instantaneous water‐use efficiency (W_t) in spring time has increased and transpiration has probably decreased, provided that leaf temperature followed air temperature. Conversely, VPD in the months between the first and second cut has increased by 0.07 kPa since 1915, offsetting the effect of increased W_i on W_t during summer and early autumn. Our results suggest that vegetation has adjusted physiologically to elevated CO₂ by decreasing stomatal conductance in this nutrient‐limited grassland.     

Forest water use and water use efficiency at elevated CO2: a model‐data intercomparison at two contrasting temperate forest FACE sites

Predicted responses of transpiration to elevated atmospheric CO₂ concentration (eCO₂) are highly variable amongst process‐based models. To better understand and constrain this variability amongst models, we conducted an intercomparison of 11 ecosystem models applied to data from two forest free‐air CO₂ enrichment (FACE) experiments at Duke University and Oak Ridge National Laboratory. We analysed model structures to identify the key underlying assumptions causing differences in model predictions of transpiration and canopy water use efficiency. We then compared the models against data to identify model assumptions that are incorrect or are large sources of uncertainty. We found that model‐to‐model and model‐to‐observations differences resulted from four key sets of assumptions, namely (i) the nature of the stomatal response to elevated CO₂ (coupling between photosynthesis and stomata was supported by the data); (ii) the roles of the leaf and atmospheric boundary layer (models which assumed multiple conductance terms in series predicted more decoupled fluxes than observed at the broadleaf site); (iii) the treatment of canopy interception (large intermodel variability, 2–15%); and (iv) the impact of soil moisture stress (process uncertainty in how models limit carbon and water fluxes during moisture stress). Overall, model predictions of the CO₂ effect on WUE were reasonable (intermodel μ = approximately 28% ± 10%) compared to the observations (μ = approximately 30% ± 13%) at the well‐coupled coniferous site (Duke), but poor (intermodel μ = approximately 24% ± 6%; observations μ = approximately 38% ± 7%) at the broadleaf site (Oak Ridge). The study yields a framework for analysing and interpreting model predictions of transpiration responses to eCO₂, and highlights key improvements to these types of models.     

Evidence of changing intrinsic water‐use efficiency under rising atmospheric CO2 concentrations in Boreal Fennoscandia from subfossil leaves and tree ring δ13C ratios

Investigating the many internal feedbacks within the climate system is a vital component of the effort to quantify the full effects of future anthropogenic climate change. The stomatal apertures of plants tend to close and decrease in number under elevated CO₂ concentrations, increasing water‐use efficiency (WUE) and reducing canopy evapotranspiration. Experimental and modelling studies reveal huge variations in these changes such that the warming associated with reduced evapotranspiration (known as physiological forcing) is neither well understood or constrained. Palaeo‐observations of changes in stomatal response and plant WUE under rising CO₂ might be used to better understand the processes underlying the physiological forcing feedback and to link measured changes in plant WUE to a specific physiological change in stomata. Here we use time series of tree ring (Pinus sylvestris L.) δ¹³C and subfossil leaf (Betula nana L.) measurements of stomatal density and geometry to derive records of changes in intrinsic water‐use efficiency (iWUE) and maximum stomatal conductance in the Boreal zone of northern Finland and Sweden. We investigate the rate of change in both proxies, over the recent past. The independent lines of evidence from these two different Boreal species indicate increased iWUE and reduced maximum stomatal conductance of similar magnitude from preindustrial times (ca. ad 1850) to around ad 1970. After this maximum stomatal conductance continues to decrease to ad 2000 in B. nana but iWUE in P. sylvestris reaches a plateau. We suggest that northern boreal P. sylvestris might have reached a threshold in its ability to increase WUE as CO₂ rises.     

Environmental and stomatal control of photosynthetic enhancement in the canopy of a sweetgum (Liquidambar styraciflua L.) plantation during 3 years of CO2 enrichment

Light‐saturated photosynthetic and stomatal responses to elevated CO₂ were measured in upper and mid‐canopy foliage of a sweetgum (Liquidambar styraciflua L) plantation exposed to free‐air CO₂ enrichment (FACE) for 3 years, to characterize environmental interactions with the sustained CO₂ effects in an intact deciduous forest stand. Responses were evaluated in relation to one another, and to seasonal patterns and natural environmental stresses, including high  temperatures, vapour pressure deficits (VPD), and drought. Photosynthetic CO₂ assimilation (A) averaged 46% higher in the +200 µmol mol^?1 CO₂ treatment, in mid‐ and upper canopy foliage. Stomatal conductance (g_s) averaged 14% (mid‐canopy) and 24% (upper canopy) lower under CO₂ enrichment. Variations in the relative responses of A and g_s were linked, such that greater relative stimulation of A was observed on dates when relative reductions in g_s were slight. Dry soils and high VPD reduced g_s and A in both treatments, and tended to diminish treatment differences. The absolute effects of CO₂ on A and g_s were minimized whenever g_s was low (<0·15 mol m^?2 s^?1), but relative effects, as the ratio of elevated to ambient rates, varied greatly under those conditions. Both stomatal and non‐stomatal limitations of A were involved during late season droughts. Leaf temperature had a limited influence on A and g_s, and there was no detectable relationship between prevailing temperature and CO₂ effects on A or g_s. The responsiveness of A and g_s to elevated CO₂, both absolute and relative, was maintained through time and within the canopy of this forest stand, subject to seasonal constraints and variability associated with limiting air and soil moisture.     

Acclimation response of spring wheat in a free-air CO2 enrichment (FACE) atmosphere with variable soil nitrogen regimes. 2. Net assimilation and stomatal conductance of leaves

Atmospheric CO₂ concentration continues to rise. It is important, therefore, to determine what acclimatory changes will occur within the photosynthetic apparatus of wheat (Triticum aestivum L. cv. Yecora Rojo) grown in a future high-CO₂ world at ample and limited soil N contents. Wheat was grown in an open field exposed to the CO₂ concentration of ambient air [370 μmol (CO₂) mol⁻¹; Control] and air enriched to ∼200 μmol (CO₂) mol⁻¹ above ambient using a Free-Air CO₂ Enrichment (FACE) apparatus (main plot). A High (35 g m⁻²) or Low (7 and 1.5 g m⁻² for 1996 and 1997, respectfully) level of N was applied to each half of the main CO₂ treatment plots (split-plot). Under High-N, FACE reduced stomatal conductance (g _s) by 30% at mid-morning (2 h prior to solar noon), 36% at midday (solar noon) and 27% at mid-afternoon (2.5 h after solar noon), whereas under Low-N, g _s was reduced by as much as 31% at mid-morning, 44% at midday and 28% at mid-afternoon compared with Control. But, no significant CO₂ × N interaction effects occurred. Across seasons and growth stages, daily accumulation of carbon (A′) was 27% greater in FACE than Control. High-N increased A′ by 18% compared with Low-N. In contrast to results for g _s, however, significant CO₂ × N interaction effects occurred because FACE increased A′ by 30% at High-N, but by only 23% at Low-N. FACE enhanced the seasonal accumulation of carbon (A′′) by 29% during 1996 (moderate N-stress), but by only 21% during 1997 (severe N-stress). These results support the premise that in a future high-CO₂ world an acclimatory (down-regulation) response in the photosynthetic apparatus of field-grown wheat is anticipated. They also demonstrate, however, that the stimulatory effect of a rise in atmospheric CO₂ on carbon gain in wheat can be maintained if nutrients such as nitrogen are in ample supply. This revised version was published online in June 2006 with corrections to the Cover Date.     

Response of wheat growth,grain yield and water use to elevated CO2 under a Free‐Air CO2 Enrichment (FACE) experiment and modelling in a semi‐arid environment

The response of wheat crops to elevated CO₂ (eCO₂) was measured and modelled with the Australian Grains Free‐Air CO₂ Enrichment experiment, located at Horsham, Australia. Treatments included CO₂ by water, N and temperature. The location represents a semi‐arid environment with a seasonal VPD of around 0.5 kPa. Over 3 years, the observed mean biomass at anthesis and grain yield ranged from 4200 to 10 200 kg ha^?1 and 1600 to 3900 kg ha^?1, respectively, over various sowing times and irrigation regimes. The mean observed response to daytime eCO₂ (from 365 to 550 μmol mol^?1 CO₂) was relatively consistent for biomass at stem elongation and at anthesis and LAI at anthesis and grain yield with 21%, 23%, 21% and 26%, respectively. Seasonal water use was decreased from 320 to 301 mm (P = 0.10) by eCO₂, increasing water use efficiency for biomass and yield, 36% and 31%, respectively. The performance of six models (APSIM‐Wheat, APSIM‐Nwheat, CAT‐Wheat, CROPSYST, OLEARY‐CONNOR and SALUS) in simulating crop responses to eCO₂ was similar and within or close to the experimental error for accumulated biomass, yield and water use response, despite some variations in early growth and LAI. The primary mechanism of biomass accumulation via radiation use efficiency (RUE) or transpiration efficiency (TE) was not critical to define the overall response to eCO₂. However, under irrigation, the effect of late sowing on response to eCO₂ to biomass accumulation at DC65 was substantial in the observed data (~40%), but the simulated response was smaller, ranging from 17% to 28%. Simulated response from all six models under no water or nitrogen stress showed similar response to eCO₂ under irrigation, but the differences compared to the dryland treatment were small. Further experimental work on the interactive effects of eCO₂, water and temperature is required to resolve these model discrepancies.     

Water availability affects seasonal CO2‐induced photosynthetic enhancement in herbaceous species in a periodically dry woodland

Elevated atmospheric CO₂ (eCO₂) is expected to reduce the impacts of drought and increase photosynthetic rates via two key mechanisms: first, through decreased stomatal conductance (g_s) and increased soil water content (V_SWC) and second, through increased leaf internal CO₂ (C_i) and decreased stomatal limitations (S_lim). It is unclear if such findings from temperate grassland studies similarly pertain to warmer ecosystems with periodic water deficits. We tested these mechanisms in three important C₃ herbaceous species in a periodically dry Eucalyptus woodland and investigated how eCO₂‐induced photosynthetic enhancement varied with seasonal water availability, over a 3 year period. Leaf photosynthesis increased by 10%–50% with a 150 μmol mol^?1 increase in atmospheric CO₂ across seasons. This eCO₂‐induced increase in photosynthesis was a function of seasonal water availability, given by recent precipitation and mean daily V_SWC. The highest photosynthetic enhancement by eCO₂ (>30%) was observed during the most water‐limited period, for example, with V_SWC <0.07 in this sandy surface soil. Under eCO₂ there was neither a significant decrease in g_s in the three herbaceous species, nor increases in V_SWC, indicating no “water‐savings effect” of eCO₂. Periods of low V_SWC showed lower g_s (less than ≈ 0.12 mol m^?2 s^?1), higher relative S_lim (>30%) and decreased C_i under the ambient CO₂ concentration (aCO₂), with leaf photosynthesis strongly carboxylation‐limited. The alleviation of S_lim by eCO₂ was facilitated by increasing C_i, thus yielding a larger photosynthetic enhancement during dry periods. We demonstrated that water availability, but not eCO₂, controls g_s and hence the magnitude of photosynthetic enhancement in the understory herbaceous plants. Thus, eCO₂ has the potential to alter vegetation functioning in a periodically dry woodland understory through changes in stomatal limitation to photosynthesis, not by the “water‐savings effect” usually invoked in grasslands.