Variations in nitrogen use efficiency reflect the biochemical subtype while variations in water use efficiency reflect the evolutionary lineage of C4 grasses at inter‐glacial CO2

C₄ photosynthesis evolved multiple times in diverse lineages. Most physiological studies comparing C₄ plants were not conducted at the low atmospheric CO₂ prevailing during their evolution. Here, 24 C₄ grasses belonging to three biochemical subtypes [nicotinamide adenine dinucleotide malic enzyme (NAD‐ME), phosphoenolpyruvate carboxykinase (PCK) and nicotinamide adenine dinucleotide phosphate malic enzyme (NADP‐ME)] and six major evolutionary lineages were grown under ambient (400 μL L^?1) and inter‐glacial (280 μL L^?1) CO₂. We hypothesized that nitrogen‐related and water‐related physiological traits are associated with subtypes and lineages, respectively. Photosynthetic rate and stomatal conductance were constrained by the shared lineage, while variation in leaf mass per area (LMA), leaf N per area, plant dry mass and plant water use efficiency were influenced by the subtype. Subtype and lineage were equally important for explaining variations in photosynthetic nitrogen use efficiency (PNUE) and photosynthetic water use efficiency (PWUE). CO₂ treatment impacted most parameters. Overall, higher LMA and leaf N distinguished the Chloridoideae/NAD‐ME group, while NADP‐ME and PCK grasses were distinguished by higher PNUE regardless of lineage. Plants were characterized by high photosynthesis and PWUE when grown at ambient CO₂ and by high conductance at inter‐glacial CO₂. In conclusion, the evolutionary and biochemical diversity among C₄ grasses was aligned with discernible leaf physiology, but it remains unknown whether these traits represent ecophysiological adaptation.     

Strong photosynthetic acclimation and enhanced water‐use efficiency in grassland functional groups persist over 21 years of CO2 enrichment,independent of nitrogen supply

Uncertainty about long‐term leaf‐level responses to atmospheric CO₂ rise is a major knowledge gap that exists because of limited empirical data. Thus, it remains unclear how responses of leaf gas exchange to elevated CO₂ (eCO₂) vary among plant species and functional groups, or across different levels of nutrient supply, and whether they persist over time for long‐lived perennials. Here, we report the effects of eCO₂ on rates of net photosynthesis and stomatal conductance in 14 perennial grassland species from four functional groups over two decades in a Minnesota Free‐Air CO₂ Enrichment experiment, BioCON. Monocultures of species belonging to C₃ grasses, C₄ grasses, forbs, and legumes were exposed to two levels of CO₂ and nitrogen supply in factorial combinations over 21 years. eCO₂ increased photosynthesis by 12.9% on average in C₃ species, substantially less than model predictions of instantaneous responses based on physiological theory and results of other studies, even those spanning multiple years. Acclimation of photosynthesis to eCO₂ was observed beginning in the first year and did not strengthen through time. Yet, contrary to expectations, the response of photosynthesis to eCO₂ was not enhanced by increased nitrogen supply. Differences in responses among herbaceous plant functional groups were modest, with legumes responding the most and C₄ grasses the least as expected, but did not further diverge over time. Leaf‐level water‐use efficiency increased by 50% under eCO₂ primarily because of reduced stomatal conductance. Our results imply that enhanced nitrogen supply will not necessarily diminish photosynthetic acclimation to eCO₂ in nitrogen‐limited systems, and that significant and consistent declines in stomatal conductance and increases in water‐use efficiency under eCO₂ may allow plants to better withstand drought.     

Physiological acclimation dampens initial effects of elevated temperature and atmospheric CO2 concentration in mature boreal Norway spruce

Physiological processes of terrestrial plants regulate the land–atmosphere exchange of carbon, water, and energy, yet few studies have explored the acclimation responses of mature boreal conifer trees to climate change. Here we explored the acclimation responses of photosynthesis, respiration, and stomatal conductance to elevated temperature and/or CO₂ concentration ([CO₂]) in a 3‐year field experiment with mature boreal Norway spruce. We found that elevated [CO₂] decreased photosynthetic carboxylation capacity (?23% at 25 °C) and increased shoot respiration (+64% at 15 °C), while warming had no significant effects. Shoot respiration, but not photosynthetic capacity, exhibited seasonal acclimation. Stomatal conductance at light saturation and a vapour pressure deficit of 1 kPa was unaffected by elevated [CO₂] but significantly decreased (?27%) by warming, and the ratio of intercellular to ambient [CO₂] was enhanced (+17%) by elevated [CO₂] and decreased (?12%) by warming. Many of these responses differ from those typically observed in temperate tree species. Our results show that long‐term physiological acclimation dampens the initial stimulation of plant net carbon assimilation to elevated [CO₂], and of plant water use to warming. Models that do not account for these responses may thus overestimate the impacts of climate change on future boreal vegetation–atmosphere interactions.     

The acclimation of leaf photosynthesis of wheat and rice to seasonal temperature changes in T‐FACE environments

Crops show considerable capacity to adjust their photosynthetic characteristics to seasonal changes in temperature. However, how photosynthesis acclimates to changes in seasonal temperature under future climate conditions has not been revealed. We measured leaf photosynthesis (A_n) of wheat (Triticum aestivum L.) and rice (Oryza sativa L.) grown under four combinations of two levels of CO₂ (ambient and enriched up to 500 µmol/mol) and two levels of canopy temperature (ambient and increased by 1.5–2.0°C) in temperature by free‐air CO₂ enrichment (T‐FACE) systems. Parameters of a biochemical C₃‐photosynthesis model and of a stomatal conductance (g_s) model were estimated for the four conditions and for several crop stages. Some biochemical parameters related to electron transport and most g_s parameters showed acclimation to seasonal growth temperature in both crops. The acclimation response did not differ much between wheat and rice, nor among the four treatments of the T‐FACE systems, when the difference in the seasonal growth temperature was accounted for. The relationships between biochemical parameters and leaf nitrogen content were consistent across leaf ranks, developmental stages, and treatment conditions. The acclimation had a strong impact on g_s model parameters: when parameter values of a particular stage were used, the model failed to correctly estimate g_s values of other stages. Further analysis using the coupled g_s–biochemical photosynthesis model showed that ignoring the acclimation effect did not result in critical errors in estimating leaf photosynthesis under future climate, as long as parameter values were measured or derived from data obtained before flowering.     

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.     

Asymmetric responses of adaxial and abaxial stomata to elevated CO2: impacts on the control of gas exchange by leaves

The response of adaxial and abaxial stomatal conductance in Rumex obtusifolius to growth at elevated atmospheric concentrations of CO₂ (250 μmol mol^?1 above ambient) was investigated over two growing seasons. The conductance of both the adaxial and abaxial leaf surfaces was found to be reduced by elevated concentrations of CO₂. Elevated CO₂ caused a much greater reduction in conductance for the adaxial surface than for the abaxial surface. The absence of effects upon stomatal density indicated that the reductions were probably the result of changes in stomatal aperture. Partitioning of gas exchange between the leaf surfaces revealed that increased concentrations of CO₂ caused increased rates of photosynthesis only via the abaxial surface. Additionally, leaf thickness was found to increase during growth at elevated concentrations of CO₂. The tendency for these amphistomatous leaves to develop a distribution of conductance approaching that of hypostomatous leaves clearly reduced their maximum photosynthetic potential. This conclusion was supported by measurements of stomatal limitation, which showed greater values for the adaxial surfaces, and greater values at elevated CO₂. This reduction in photosynthesis may in part be caused by higher diffusive limitations imposed because of increased leaf thickness. In an uncoupled canopy, asymmetrical stomatal responses of the kind identified here may appreciably reduce transpiration. Species which show symmetrical responses are less likely to show reduced transpirational rates, and a redistribution of water loss between species may occur. The implications of asymmetrical stomatal responses for photosynthesis and canopy transpiration are discussed.     

Photosynthetic acclimation to elevated CO2 in Phaseolus vulgaris L. is altered by growth response to nitrogen supply

Abstract Long‐term exposure of plants to elevated CO₂ often leads to downward photosynthetic acclimation. Nitrogen (N) deficiency could potentially exacerbate this response by reducing growth rate and the sink for photosynthates, but this has not always been observed. Experimentally, the interpretation of N effects on CO₂ responses can be confounded by increasing severity of tissue N deficiency over time when N supply is not adjusted as demand increases. In this study, N supply ranged from sub‐ to supra‐optimal (20–540 kgN ha^–l equivalent), and relatively stable levels of tissue N concentration were obtained in all treatments by varying twice‐weekly application rates in proportion to plant growth. The effects of N on photosynthesis and growth of beans (Phaseolus vulgaris L.) raised at ambient (35 Pa) and three elevated (70, 105, 140 Pa) CO₂ partial pressures (pCO₂) were evaluated. Averaging across N treatments, leaf total non‐structural carbohydrates (TNC) were 2.5‐ to 3‐fold higher and leaf N concentrations were 31–35% lower at elevated compared to ambient pCO₂. Light‐saturated net CO₂ assimilation rates measured at growth pCO₂ (A_satg) were significantly higher (26–40% depending on N supply) in plants grown at elevated compared to ambient pCO₂. When measured at a common pCO₂ of 35 Pa, the A_sat of plants grown at elevated CO₂ was 15–29% less than that of plants grown at 35 Pa, indicative of downward photosynthetic acclimation. The magnitude of downward photosynthetic acclimation to elevated CO₂ was greater in plants grown at high (180 and 540 kgN ha^–l) compared to low (20 and 60 kgN ha^–l) N supply, and this was associated with a higher A_sat at growth pCO₂, higher leaf area ratio (leaf area/total biomass), and higher TNC in leaves of high‐N plants. Our results indicate that the effect of N on acclimation to CO₂ will depend on the balance between supply and demand for N during the growing period, and the effect this has on biomass allocation and source‐sink C balance at the whole‐plant level.     

The apparent feed-forward response to vapour pressure deficit of stomata in droughted, field-grown Eucalyptus globulus Labill

This study tested a multiplicative model of stomatal response to environment for drought‐affected trees of Eucalyptus globulus Labill. growing in southern Australia. The model incorporates a feed‐forward response to vapour pressure deficit of ambient air (δe_a) and performed well if evaluated using reduced major axis regression and log‐transformed data. There was strong evidence from gas‐exchange data, leaf water potentials and sapflow measurements of the feed‐forward response by stomata to leaf‐to‐air vapour pressure deficit (δe_l). The response of stomata to δe_l was irreversible. Stomatal conductance and the rate of net photosynthesis were highly correlated and declined, together with the rate of transpiration, throughout the afternoon as δe_a increased despite increasing leaf water potentials. The concentration of CO₂ inside leaves (c_i) increased as stomatal conductance declined indicating increasing non‐stomatal limitations to photosynthesis. The stomatal response to δe_l of E. globulus in the field is best described as an ‘apparent feed‐forward response’ that probably results from both slowly reversible depression of net photosynthesis and abscisic acid accumulation in guard cells. We suggest that the stomatal response to c_i may strengthen the link between photosynthetic capacity and stomatal conductance during leaf drying as a result of either drought or large δ e_l.     

Regulation and acclimation of leaf gas exchange in a piñon–juniper woodland exposed to three different precipitation regimes

Leaf gas‐exchange regulation plays a central role in the ability of trees to survive drought, but forecasting the future response of gas exchange to prolonged drought is hampered by our lack of knowledge regarding potential acclimation. To investigate whether leaf gas‐exchange rates and sensitivity to drought acclimate to precipitation regimes, we measured the seasonal variations of leaf gas exchange in a mature piñon–juniper Pinus edulis–Juniperus monosperma woodland after 3 years of precipitation manipulation. We compared trees receiving ambient precipitation with those in an irrigated treatment (+30% of ambient precipitation) and a partial rainfall exclusion (?45%). Treatments significantly affected leaf water potential, stomatal conductance and photosynthesis for both isohydric piñon and anisohydric juniper. Leaf gas exchange acclimated to the precipitation regimes in both species. Maximum gas‐exchange rates under well‐watered conditions, leaf‐specific hydraulic conductance and leaf water potential at zero photosynthetic assimilation all decreased with decreasing precipitation. Despite their distinct drought resistance and stomatal regulation strategies, both species experienced hydraulic limitation on leaf gas exchange when precipitation decreased, leading to an intraspecific trade‐off between maximum photosynthetic assimilation and resistance of photosynthesis to drought. This response will be most detrimental to the carbon balance of piñon under predicted increases in aridity in the southwestern USA.     

The effects on Arbutus unedo L. of long-term exposure to elevated CO2

Arbutus unedo is a sclerophyllous evergreen, characteristic of Mediterranean coastal scrub vegetation. In Italy, trees of A. unedo have been found close to natural CO₂ vents where the mean atmospheric carbon dioxide concentration is about 2200 μmol mol^?1. Comparisons were made between trees growing in elevated and ambient CO₂ concentrations to test for evidence of adaptation to long-term exposure to elevated CO₂. Leaves formed at elevated CO₂ have a lower stomatal density and stomatal index and higher specific leaf area than those formed at ambient CO₂, but there was no change in carbon to nitrogen ratios of the leaf tissue. Stomatal conductance was lower at elevated CO₂ during rapid growth in the spring. In mid-summer, under drought stress, stomatal closure of all leaves occurred and in the autumn, when stress was relieved, the conductance of leaves at both elevated and ambient CO₂ increased. In the spring, the stomatal conductance of the new flush of leaves at ambient CO₂ was higher than the leaves at elevated CO₂, increasing instantaneous water use efficiency at elevated CO₂. Chlorophyll fluorescence measurements suggested that elevated CO₂ provided some protection against photoinhibition in mid-summer. Analysis of A/C_i curves showed that there was no evidence of either upward or downward regulation of photosynthesis at elevated CO₂. It is therefore anticipated that A. unedo will have higher growth rates as the ambient CO₂ concentrations increase.     

Hydrologic balance in an intact temperate forest ecosystem under ambient and elevated atmospheric CO2 concentration

Abstract Increasing atmospheric CO₂ concentration decreases stomatal conductance in many species, but the savings of water from reduced transpiration may permit the forest to retain greater leaf area index (L). Therefore, the net effect on water use in forest ecosystems under a higher CO₂ atmosphere is difficult to predict. The free air CO₂ enrichment (FACE) facility (n = 3) in a 14‐m tall (in 1996) Pinus taeda L. stand was designed to reduce uncertainties in predicting such responses. Continuous measurements of precipitation, throughfall precipitation, sap flux, and soil moisture were made over 3.5 years under ambient (CO₂^a) and elevated (CO₂^e) ambient + 200 µmol mol^?1). Annual stand transpiration under ambient CO₂ conditions accounted for 84–96% of latent heat flux measured with the eddy‐covariance technique above the canopy. Under CO₂^e, P. taeda transpired less per unit of leaf area only when soil drought was severe. Liquidambar styraciflua, the other major species in the forest, used progressively less water, settling at 25% reduction in sap flux density after 3.5 years under CO₂^e. Because P. taeda dominated the stand, and severe drought periods were of relatively short duration, the direct impact of CO₂^e on water savings in the stand was undetectable. Moreover, the forest used progressively more water under CO₂^e, probably because soil moisture availability progressively increased, probably owing to a reduction in soil evaporation caused by more litter buildup in the CO₂^e plots. The results suggest that, in this forest, the effect of CO₂^e on transpiration was greater indirectly through enhanced litter production than directly through reduced stomatal conductance. In forests composed of species more similar to L. styraciflua, water savings from stomatal closure may dominate the response to CO₂^e.     

Photosynthetic response of Tempranillo grapevine to climate change scenarios

Photosynthesis in C₃ plants is CO₂ limited and therefore any increase in Rubisco carboxylation substrate may increase net CO₂ fixation, unless plants experience acclimation or other limitations. These aspects are largely unexplored in grapevine. Photosynthesis analysis was used to assess the stomatal, mesophyll, photochemical and biochemical contributions to the decreasing photosynthesis observed in Tempranillo grapevines (Vitis vinifera) from veraison to ripeness, modulated by CO₂, temperature and water availability. Photosynthesis and photosystem II photochemistry decreased from veraison to ripeness. The elevated CO₂ and temperature increased photosynthesis, but transiently, in both well irrigated (WI) and water‐stressed plants. Photosynthetic rates were maxima 1 week after the start of elevated CO₂ and temperature treatments, but differences with treatments of ambient conditions disappeared with time. There were not marked changes in leaf water status, leaf chlorophyll or leaf protein that could limit photosynthesis at ripeness. Leaf total soluble sugars remained at ripeness as high as 2 weeks after the start of treatments. On the other hand, and as expected, CO₂ diffusional limitations impaired photosynthesis in grapevine plants grown under water scarcity, stomatal and mesophyll conductances to CO₂ decreased and in turn low chloroplastic CO₂ concentrations limited photosynthetic CO₂ fixation. In summary, photochemistry and photosynthesis from veraison to ripeness in Tempranillo grapevine were dominated by a developmental‐related decreasing trend that was only transiently influenced by elevated CO₂ concentrations.     

Seasonal patterns of photosynthetic response and acclimation to elevated carbon dioxide in field-grown strawberry

Strawberry (Fragaria × ananassa) plants were grown in field plots at the current ambient [CO₂], and at ambient + 300 and ambient + 600 μmol mol⁻¹ [CO₂]. Approximately weekly measurements were made of single leaf gas exchange of upper canopy leaves from early spring through fall of two years, in order to determine the temperature dependence of the stimulation of photosynthesis by elevated [CO₂], whether growth at elevated [CO₂] resulted in acclimation of photosynthesis, and whether any photosynthetic acclimation was reduced when fruiting created additional demand for the products of photosynthesis. Stimulation of photosynthetic CO₂ assimilation by short-term increases in [CO₂] increased strongly with measurement temperature. The stimulation exceeded that predicted from the kinetic characteristics of ribulose-1,5-bisphosphate carboxylase at all temperatures. Acclimation of photosynthesis to growth at elevated [CO₂] was evident from early spring through summer, including the fruiting period in early summer, with lower rates under standard measurement conditions in plants grown at elevated [CO₂]. The degree of acclimation increased with growth [CO₂]. However, there were no significant differences between [CO₂] treatments in total nitrogen per leaf area, and photosynthetic acclimation was reversed one day after switching the [CO₂] treatments. Tests showed that acclimation did not result from a limitation of photosynthesis by triose phosphate utilization rate at elevated [CO₂]. Photosynthetic acclimation was not evident during dry periods in midsummer, when the elevated [CO₂] treatments conserved soil water and photosynthesis declined more at ambient than at elevated [CO₂]. Acclimation was also not evident during the fall, when plants were vegetative, despite wet conditions and continued higher leaf starch content at elevated [CO₂]. Stomatal conductance responded little to short-term changes in [CO₂] except during drought, and changed in parallel with photosynthetic acclimation through the seasons in response to the long-term [CO₂] treatments. The data do not support the hypothesis that source-sink balance controls the seasonal occurrence of photosynthetic acclimation to elevated [CO₂] in this species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Mesophyll conductance in land surface models: effects on photosynthesis and transpiration

The CO₂ transfer conductance within plant leaves (mesophyll conductance, g_m) is currently not considered explicitly in most land surface models (LSMs), but instead treated implicitly as an intrinsic property of the photosynthetic machinery. Here, we review approaches to overcome this model deficiency by explicitly accounting for g_m, which comprises the re‐adjustment of photosynthetic parameters and a model describing the variation of g_m in dependence of environmental conditions. An explicit representation of g_m causes changes in the response of photosynthesis to environmental factors, foremost leaf temperature, and ambient CO₂ concentration, which are most pronounced when g_m is small. These changes in leaf‐level photosynthesis translate into a stronger climate and CO₂ response of gross primary productivity (GPP) and transpiration at the global scale. The results from two independent studies show consistent latitudinal patterns of these effects with biggest differences in GPP in the boreal zone (up to ~15%). Transpiration and evapotranspiration show spatially similar, but attenuated, changes compared with GPP. These changes are indirect effects of g_m caused by the assumed strong coupling between stomatal conductance and photosynthesis in current LSMs. Key uncertainties in these simulations are the variation of g_m with light and the robustness of its temperature response across plant types and growth conditions. Future research activities focusing on the response of g_m to environmental factors and its relation to other plant traits have the potential to improve the representation of photosynthesis in LSMs and to better understand its present and future role in the Earth system.     

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.     

Do all leaf photosynthesis parameters of rice acclimate to elevated CO2, elevated temperature,and their combination,in FACE environments?

Leaf photosynthesis of crops acclimates to elevated CO₂ and temperature, but studies quantifying responses of leaf photosynthetic parameters to combined CO₂ and temperature increases under field conditions are scarce. We measured leaf photosynthesis of rice cultivars Changyou 5 and Nanjing 9108 grown in two free‐air CO₂ enrichment (FACE) systems, respectively, installed in paddy fields. Each FACE system had four combinations of two levels of CO₂ (ambient and enriched) and two levels of canopy temperature (no warming and warmed by 1.0–2.0°C). Parameters of the C₃ photosynthesis model of Farquhar, von Caemmerer and Berry (the FvCB model), and of a stomatal conductance (g_s) model were estimated for the four conditions. Most photosynthetic parameters acclimated to elevated CO₂, elevated temperature, and their combination. The combination of elevated CO₂ and temperature changed the functional relationships between biochemical parameters and leaf nitrogen content for Changyou 5. The g_s model significantly underestimated g_s under the combination of elevated CO₂ and temperature by 19% for Changyou 5 and by 10% for Nanjing 9108 if no acclimation was assumed. However, our further analysis applying the coupled g_s–FvCB model to an independent, previously published FACE experiment showed that including such an acclimation response of g_s hardly improved prediction of leaf photosynthesis under the four combinations of CO₂ and temperature. Therefore, the typical procedure that crop models using the FvCB and g_s models are parameterized from plants grown under current ambient conditions may not result in critical errors in projecting productivity of paddy rice under future global change.     

Reversibility of photosynthetic acclimation of swiss chard and sugarbeet grown at elevated concentrations of CO2

Although leaf photosynthesis and plant growth are initially stimulated by elevated CO₂ concentrations, increasing insensitivity to CO₂ (acclimation) is a frequent occurrence. In order to examine the acclimation process, we studied photosynthesis and whole plant development in swiss chard (Beta vulgaris L. Koch ssp. ciela) and sugarbeet (Beta vulgaris L. ssp. vulgaris) grown at either ambient or twice ambient concentrations of CO₂. In an initial controlled environment study, photosynthetic acclimation to elevated CO₂ levels was observed in both subspecies 24 days after sowing (DAS) but was not observed at 42 and 49 DAS for sugarbeet or at 49 DAS for swiss chard. Although sugarbeet and swiss chard differed in root size and morphology, this was not a factor in the onset of photosynthetic acclimation. The reversal of photosynthetic acclimation that was observed in older plants grown at elevated CO₂, concentrations was associated with a rapid increase in root development (i.e. increased root: shoot [R/S] ratio), increased sucrose levels in sinks (roots) and no differences in total soluble leaf protein of either subspecies relative to the ambient CO₂ condition. In a second set of experiments, swiss chard and sugarbeet were grown in outdoor Plexiglass chambers at different times of the year (i.e. summer and early fall). Average 24-h temperature was 30.7 and 19.4°C for the summer and fall plantings, respectively. In agreement with the controlled environment study, lack of photosynthetic acclimation, determined from the response of photosynthesic rate to internal CO₂ concentration, was correlated with increased root biomass and sucrose concentration relative to the ambient condition. However, photo-synthetic acclimation was observed depending on the season, i.e. summer (swiss chard) or fall (sugarbeet), suggesting that acclimation was affected by environmental factors, such as temperature. Data from both experiments suggest that continued long-term photosynthetic stimulation may be dependent upon the ability of increased CO₂ to stimulate new sink development which would allow full utilization of the additional carbon made available in a high CO₂ environment.     

Elevated CO2 enhances water relations and productivity and affects gas exchange in C3 and C4 grasses of the Colorado shortgrass steppe.

Six open‐top chambers were installed on the shortgrass steppe in north‐eastern Colorado, USA from late March until mid‐October in 1997 and 1998 to evaluate how this grassland will be affected by rising atmospheric CO₂. Three chambers were maintained at current CO₂ concentration (ambient treatment), three at twice ambient CO₂, or approximately 720 μmol mol^?1 (elevated treatment), and three nonchambered plots served as controls. Above‐ground phytomass was measured in summer and autumn during each growing season, soil water was monitored weekly, and leaf photosynthesis, conductance and water potential were measured periodically on important C₃ and C₄ grasses. Mid‐season and seasonal above‐ground productivity were enhanced from 26 to 47% at elevated CO₂, with no differences in the relative responses of C₃/C₄ grasses or forbs. Annual above‐ground phytomass accrual was greater on plots which were defoliated once in mid‐summer compared to plots which were not defoliated during the growing season, but there was no interactive effect of defoliation and CO₂ on growth. Leaf photosynthesis was often greater in Pascopyrum smithii (C₃) and Bouteloua gracilis (C₄) plants in the elevated chambers, due in large part to higher soil water contents and leaf water potentials. Persistent downward photosynthetic acclimation in P. smithii leaves prevented large photosynthetic enhancement for elevated CO₂‐grown plants. Shoot N concentrations tended to be lower in grasses under elevated CO₂, but only Stipa comata (C₃) plants exhibited significant reductions in N under elevated compared to ambient CO₂ chambers. Despite chamber warming of 2.6 °C and apparent drier chamber conditions compared to unchambered controls, above‐ground production in all chambers was always greater than in unchambered plots. Collectively, these results suggest increased productivity of the shortgrass steppe in future warmer, CO₂ enriched environments.     

Increased stomatal conductance induces rapid changes to photosynthetic rate in response to naturally fluctuating light conditions in rice

A close correlation between stomatal conductance and the steady-state photosynthetic rate has been observed for diverse plant species under various environmental conditions. However, it remains unclear whether stomatal conductance is a major limiting factor for the photosynthetic rate under naturally fluctuating light conditions. We analysed a SLAC1 knockout rice line to examine the role of stomatal conductance in photosynthetic responses to fluctuating light. SLAC1 encodes a stomatal anion channel that regulates stomatal closure. Long exposures to weak light before treatments with strong light increased the photosynthetic induction time required for plants to reach a steady-state photosynthetic rate and also induced stomatal limitation of photosynthesis by restricting the diffusion of CO₂ into leaves. The slac1 mutant exhibited a significantly higher rate of stomatal opening after an increase in irradiance than wild-type plants, leading to a higher rate of photosynthetic induction. Under natural conditions, in which irradiance levels are highly variable, the stomata of the slac1 mutant remained open to ensure efficient photosynthetic reaction. These observations reveal that stomatal conductance is important for regulating photosynthesis in rice plants in the natural environment with fluctuating light.     

Do photosynthetic limitations of evergreen Quercus ilex leaves change with long‐term increased drought severity?

Seasonal drought can severely impact leaf photosynthetic capacity. This is particularly important for Mediterranean forests, where precipitation is expected to decrease as a consequence of climate change. Impacts of increased drought on the photosynthetic capacity of the evergreen Quercus ilex were studied for two years in a mature forest submitted to long‐term throughfall exclusion. Gas exchange and chlorophyll fluorescence were measured on two successive leaf cohorts in a control and a dry plot. Exclusion significantly reduced leaf water potential in the dry treatment. In both treatments, light‐saturated net assimilation rate (A_max), stomatal conductance (g_s), maximum carboxylation rate (V_cmax), maximum rate of electron transport (J_max), mesophyll conductance to CO₂ (g_m) and nitrogen investment in photosynthesis decreased markedly with soil water limitation during summer. The relationships between leaf photosynthetic parameters and leaf water potential remained identical in the two treatments. Leaf and canopy acclimation to progressive, long‐term drought occurred through changes in leaf area index, leaf mass per area and leaf chemical composition, but not through modifications of physiological parameters.