Guard cell photosynthesis is critical for stomatal turgor production,yet does not directly mediate CO2‐ and ABA‐induced stomatal closing

Stomata mediate gas exchange between the inter‐cellular spaces of leaves and the atmosphere. CO₂ levels in leaves (Ci) are determined by respiration, photosynthesis, stomatal conductance and atmospheric [CO₂]. [CO₂] in leaves mediates stomatal movements. The role of guard cell photosynthesis in stomatal conductance responses is a matter of debate, and genetic approaches are needed. We have generated transgenic Arabidopsis plants that are chlorophyll‐deficient in guard cells only, expressing a constitutively active chlorophyllase in a guard cell specific enhancer trap line. Our data show that more than 90% of guard cells were chlorophyll‐deficient. Interestingly, approximately 45% of stomata had an unusual, previously not‐described, morphology of thin‐shaped chlorophyll‐less stomata. Nevertheless, stomatal size, stomatal index, plant morphology, and whole‐leaf photosynthetic parameters (PSII, qP, qN, F_V′/F_M′) were comparable with wild‐type plants. Time‐resolved intact leaf gas‐exchange analyses showed a reduction in stomatal conductance and CO₂‐assimilation rates of the transgenic plants. Normalization of CO₂ responses showed that stomata of transgenic plants respond to [CO₂] shifts. Detailed stomatal aperture measurements of normal kidney‐shaped stomata, which lack chlorophyll, showed stomatal closing responses to [CO₂] elevation and abscisic acid (ABA), while thin‐shaped stomata were continuously closed. Our present findings show that stomatal movement responses to [CO₂] and ABA are functional in guard cells that lack chlorophyll. These data suggest that guard cell CO₂ and ABA signal transduction are not directly modulated by guard cell photosynthesis/electron transport. Moreover, the finding that chlorophyll‐less stomata cause a ‘deflated’ thin‐shaped phenotype, suggests that photosynthesis in guard cells is critical for energization and guard cell turgor production.     

Photosynthetic limitations in olive cultivars with different sensitivity to salt stress

Olive (Olea europea L) is one of the most valuable and widespread fruit trees in the Mediterranean area. To breed olive for resistance to salinity, an environmental constraint typical of the Mediterranean, is an important goal. The photosynthetic limitations associated with salt stress caused by irrigation with saline (200 mm ) water were assessed with simultaneous gas‐exchange and fluorescence field measurements in six olive cultivars. Cultivars were found to possess inherently different photosynthesis when non‐stressed. When exposed to salt stress, cultivars with inherently high photosynthesis showed the highest photosynthetic reductions. There was no relationship between salt accumulation and photosynthesis reduction in either young or old leaves. Thus photosynthetic sensitivity to salt did not depend on salt exclusion or compartmentalization in the old leaves of the olive cultivars investigated. Salt reduced the photochemical efficiency, but this reduction was also not associated with photosynthesis reduction. Salt caused a reduction of stomatal and mesophyll conductance, especially in cultivars with inherently high photosynthesis. Mesophyll conductance was generally strongly associated with photosynthesis, but not in salt‐stressed leaves with a mesophyll conductance higher than 50 mmol m^?2 s^?1. The combined reduction of stomatal and mesophyll conductances in salt‐stressed leaves increased the CO₂ draw‐down between ambient air and the chloroplasts. The CO₂ draw‐down was strongly associated with photosynthesis reduction of salt‐stressed leaves but also with the variable photosynthesis of controls. The relationship between photosynthesis and CO₂ draw‐down remained unchanged in most of the cultivars, suggesting no or small changes in Rubisco activity of salt‐stressed leaves. The present results indicate that the low chloroplast CO₂ concentration set by both low stomatal and mesophyll conductances were the main limitations of photosynthesis in salt‐stressed olive as well as in cultivars with inherently low photosynthesis. It is consequently suggested that, independently of the apparent sensitivity of photosynthesis to salt, this effect may be relieved if conductances to CO₂ diffusion are restored.     

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.     

Stomatal and non-stomatal limitation to photosynthesis in two trembling aspen (Populus tremuloides Michx.) clones exposed to elevated CO2 and/or O3

Leaf gas exchange parameters and the content of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) in the leaves of two 2‐year‐old aspen (Populus tremuloides Michx.) clones (no. 216, ozone tolerant and no. 259, ozone sensitive) were determined to estimate the relative stomatal and mesophyll limitations to photosynthesis and to determine how these limitations were altered by exposure to elevated CO₂ and/or O₃. The plants were exposed either to ambient air (control), elevated CO₂ (560 p.p.m.) elevated O₃ (55 p.p.b.) or a mixture of elevated CO₂ and O₃ in a free air CO₂ enrichment (FACE) facility located near Rhinelander, Wisconsin, USA. Light‐saturated photosynthesis and stomatal conductance were measured in all leaves of the current terminal and of two lateral branches (one from the upper and one from the lower canopy) to detect possible age‐related variation in relative stomatal limitation (leaf age is described as a function of leaf plastochron index). Photosynthesis was increased by elevated CO₂ and decreased by O₃ at both control and elevated CO₂. The relative stomatal limitation to photosynthesis (l_s) was in both clones about 10% under control and elevated O₃. Exposure to elevated CO₂ + O₃ in both clones and to elevated CO₂ in clone 259, decreased l_s even further – to about 5%. The corresponding changes in Rubisco content and the stability of C_i/C_a ratio suggest that the changes in photosynthesis in response to elevated CO₂ and O₃ were primarily triggered by altered mesophyll processes in the two aspen clones of contrasting O₃ tolerance. The changes in stomatal conductance seem to be a secondary response, maintaining stable C_i under the given treatment, that indicates close coupling between stomatal and mesophyll processes.     

Weak coordination among petiole,leaf, vein,and gas‐exchange traits across Australian angiosperm species and its possible implications

Close coordination between leaf gas exchange and maximal hydraulic supply has been reported across diverse plant life forms. However, it has also been suggested that this relationship may become weak or break down completely within the angiosperms. We examined coordination between hydraulic, leaf vein, and gas‐exchange traits across a diverse group of 35 evergreen Australian angiosperms, spanning a large range in leaf structure and habitat. Leaf‐specific conductance was calculated from petiole vessel anatomy and was also measured directly using the rehydration technique. Leaf vein density (thought to be a determinant of gas exchange rate), maximal stomatal conductance, and net CO₂ assimilation rate were also measured for most species (n = 19–35). Vein density was not correlated with leaf‐specific conductance (either calculated or measured), stomatal conductance, nor maximal net CO₂ assimilation, with r² values ranging from 0.00 to 0.11, P values from 0.909 to 0.102, and n values from 19 to 35 in all cases. Leaf‐specific conductance calculated from petiole anatomy was weakly correlated with maximal stomatal conductance (r² = 0.16; P = 0.022; n = 32), whereas the direct measurement of leaf‐specific conductance was weakly correlated with net maximal CO₂ assimilation (r² = 0.21; P = 0.005; n = 35). Calculated leaf‐specific conductance, xylem ultrastructure, and leaf vein density do not appear to be reliable proxy traits for assessing differences in rates of gas exchange or growth across diverse sets of evergreen angiosperms.     

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.     

Carbon dioxide diffusion across stomata and mesophyll and photo-biochemical processes as affected by growth CO2 and phosphorus nutrition in cotton

Nutrients such as phosphorus may exert a major control over plant response to rising atmospheric carbon dioxide concentration (CO₂), which is projected to double by the end of the 21st century. Elevated CO₂ may overcome the diffusional limitations to photosynthesis posed by stomata and mesophyll and alter the photo-biochemical limitations resulting from phosphorus deficiency. To evaluate these ideas, cotton (Gossypium hirsutum) was grown in controlled environment growth chambers with three levels of phosphate (Pi) supply (0.2, 0.05 and 0.01 mM) and two levels of CO₂ concentration (ambient 400 and elevated 800 μmol mol⁻¹) under optimum temperature and irrigation. Phosphate deficiency drastically inhibited photosynthetic characteristics and decreased cotton growth for both CO₂ treatments. Under Pi stress, an apparent limitation to the photosynthetic potential was evident by CO₂ diffusion through stomata and mesophyll, impairment of photosystem functioning and inhibition of biochemical process including the carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxyganase and the rate of ribulose-1,5-bisphosphate regeneration. The diffusional limitation posed by mesophyll was up to 58% greater than the limitation due to stomatal conductance (g_s) under Pi stress. As expected, elevated CO₂ reduced these diffusional limitations to photosynthesis across Pi levels; however, it failed to reduce the photo-biochemical limitations to photosynthesis in phosphorus deficient plants. Acclimation/down regulation of photosynthetic capacity was evident under elevated CO₂ across Pi treatments. Despite a decrease in phosphorus, nitrogen and chlorophyll concentrations in leaf tissue and reduced stomatal conductance at elevated CO₂, the rate of photosynthesis per unit leaf area when measured at the growth CO₂ concentration tended to be higher for all except the lowest Pi treatment. Nevertheless, plant biomass increased at elevated CO₂ across Pi nutrition with taller plants, increased leaf number and larger leaf area.     

Effect of leaf surface wetness and wettability on photosynthesis in bean and pea

Leaf surface wetness that occurs frequently in natural environments has a significant impact on leaf photosynthesis. However, the physiological mechanisms for the photosynthetic responses to wetness are not well understood. The responses of leaf CO₂ assimilation rate (A) to 72 h of artificial mist of a wettable (bean; Phaseolus vulgaris) and a non‐wettable species (pea; Pisum sativum) were compared. Stomatal and non‐stomatal limitations to A were investigated. A 28% inhibition of A was observed in the bean leaves as a result of a 16% decrease in stomatal conductance and a 55% reduction in the amount of Rubisco. The decrease of Rubisco was mainly due to its partial degradation. In contrast to the bean leaves, a 22% stimulation of A was obtained in the 72 h mist‐treated pea leaves. Mist treatment increased stomatal conductance by 12.5% and had no effect on the amount of Rubisco. These results indicated that a positive photosynthetic response to wetness occurred only in non‐wettable species and is due to the change in stomatal regulation.     

From reproduction to production,stomata are the master regulators

The best predictor of leaf level photosynthetic rate is the porosity of the leaf surface, as determined by the number and aperture of stomata on the leaf. This remarkable correlation between stomatal porosity (or diffusive conductance to water vapour g_s) and CO₂ assimilation rate (A) applies to all major lineages of vascular plants (Figure 1) and is sufficiently predictable that it provides the basis for the model most widely used to predict water and CO₂ fluxes from leaves and canopies. Yet the Ball–Berry formulation is only a phenomenological approximation that captures the emergent character of stomatal behaviour. Progressing to a more mechanistic prediction of plant gas exchange is challenging because of the diversity of biological components regulating stomatal action. These processes are the product of more than 400 million years of co‐evolution between stomatal, vascular and photosynthetic tissues. Both molecular and structural components link the abiotic world of the whole plant with the turgor pressure of the epidermis and guard cells, which ultimately determine stomatal pore size and porosity to water and CO₂ exchange (New Phytol., 168, 2005, 275). In this review we seek to simplify stomatal behaviour by using an evolutionary perspective to understand the principal selective pressures involved in stomatal evolution, thus identifying the primary regulators of stomatal aperture. We start by considering the adaptive process that has locked together the regulation of water and carbon fluxes in vascular plants, finally examining specific evidence for evolution in the proteins responsible for regulating guard cell turgor.     

Stomatal and non-stomatal limitations to carbon assimilation: an evaluation of the path-dependent method

Abstract Environmental stresses can decrease photosynthesis by a direct effect on photosynthetic capacity of the mesophyll or by a CO₂ limitation resulting from stomatal closure. In the present study, a ‘path-dependent method’ (Jones, 1985) for the partitioning of a stress-related decline in assimilation rate between non-stomatal and stomatal factors was evaluated, using light quality as a ‘stress’. Kinetic data on assimilation rate and conductance of Phragmipedium longifolium following a change in light quality from 95 μmol m^?2s^?1 white light to 95 μmol m^?2s^?1 red light failed to generate a smooth response curve for conductance. Partitioning of limitations on assimilation by a path-dependent method that utilizes the actual trajectories of conductance and assimilation was therefore not feasible. A simplified path-dependent method (Jones, 1985) which assumes that either mesophyll cells or guard cells respond first to a stress was applied to steady-state measurements of assimilation and conductance under red and white illumination. Either 5% or 23% of the observed reduction in assimilation rate under white light was attributable to stomatal factors, depending on whether the ‘stomatal first’ or the ‘mesophyll first’ path was assumed. In the absence of additional information indicating the appropriate choice of path, arbitrary choice may therefore lead to widely divergent estimates, and potentially erroneous conclusions. An alternative approach to the evaluation of the importance to carbon assimilation of stomatal and non-stomatal factors is suggested.     

Differences in gas exchange contribute to habitat differentiation in Iberian columbines from contrasting light and water environments

During photosynthesis, respiration and transpiration, gas exchange occurs via the stomata and so plants face a trade‐off between maximising photosynthesis while minimising transpiration (expressed as water use efficiency, WUE). The ability to cope with this trade‐off and regulate photosynthetic rate and stomatal conductance may be related to niche differentiation between closely related species. The present study explored this as a possible mechanism for habitat differentiation in Iberian columbines. The roles of irradiance and water stress were assessed to determine niche differentiation among Iberian columbines via distinct gas exchange processes. Photosynthesis–irradiance curves (P–I curves) were obtained for four taxa, and common garden experiments were conducted to examine plant responses to water and irradiance stress, by measuring instantaneous gas exchange and plant performance. Gas exchange was also measured in ten individuals using two to four field populations per taxon. The taxa had different P–I curves and gas exchange in the field. At the species level, water stress and irradiance explained habitat differentiation. Within each species, a combination of irradiance and water stress explained the between‐subspecies habitat differentiation. Despite differences in stomatal conductance and CO₂ assimilation, taxa did not have different WUE under field conditions, which suggests that the environment equally modifies photosynthesis and transpiration. The P–I curves, gas exchange in the field and plant responses to experimental water and irradiance stresses support the hypothesis that habitat differentiation is associated with differences among taxa in tolerance to abiotic stress mediated by distinct gas exchange responses.     

Seasonal changes in photosynthetic characteristics of Anemone raddeana,a spring-active geophyte,in the temperate region of Japan

Summary Seasonal changes in the photosynthetic characteristics of intact involucral leaves of Anemone raddeana were investigated under laboratory conditions. Net photosynthesis and constant water vapor pressure deficit showed almost the same seasonal trend. They increased rapidly from mid-April immediately after unfolding of the leaves and reached the maximum in late-April, before the maximum expansion of the leaves. They retained the maximum values until early-May and then decreased toward late-May with a progress of leaf senescence. The calculated values of intercellular CO₂ concentration and relative stomatal limitation of photosynthesis showed no significant change throughout the season. The carboxylation efficiency as assessed by the initial slope of Ci-photosynthesis curve and the net photosynthesis under a high Ci regime varied seasonally in parallel with the change of the light-saturated photosynthesis. The results indicate that the seasonal changes in light-saturated net photosynthesis are not due to a change of stomatal conductance, but to a change in the photosynthetic capacity of mesophyll. Nevertheless, leaf conductance changed concomitantly with photosynthetic capacity, indicating that the seasonal change in stomatal conductance is modulated by the mesophyll photosynthetic capacity such that the intercellular CO₂ concentrations is maintained constant. The shape of light-photosynthesis curve was similar to that of sun-leaf type. The quantum yield also changed simultaneously with the photosynthetic capacity throughout the season.Contribution No. 2965 from the Institute of Low Temperature Science     

The influence of stomatal morphology and distribution on photosynthetic gas exchange

The intricate and interconnecting reactions of C₃ photosynthesis are often limited by one of two fundamental processes: the conversion of solar energy into chemical energy, or the diffusion of CO₂ from the atmosphere through the stomata, and ultimately into the chloroplast. In this review, we explore how the contributions of stomatal morphology and distribution can affect photosynthesis, through changes in gaseous exchange. The factors driving this relationship are considered, and recent results from studies investigating the effects of stomatal shape, size, density and patterning on photosynthesis are discussed. We suggest that the interplay between stomatal gaseous exchange and photosynthesis is complex, and that a disconnect often exists between the rates of CO₂ diffusion and photosynthetic carbon fixation. The mechanisms that allow for substantial reductions in maximum stomatal conductance without affecting photosynthesis are highly dependent on environmental factors, such as light intensity, and could be exploited to improve crop performance.     

采煤塌陷影响下土壤含水量变化对柠条气孔导度、蒸腾与光合作用速率的影响

采煤塌陷引起的土壤环境因子的变化对矿区植物生长的影响越来越受到人们的关注,气孔导度、蒸腾与光合作用作为环境变化响应的敏感因子,研究植物气孔导度、蒸腾与光合作用的变化是揭示荒漠矿区自然环境变化及其规律的重要手段之一。研究采煤塌陷条件下植物光合生理的变化是探究煤炭开采对植物叶片水分蒸腾散失和CO_2同化速率影响的关键环节,是探讨采煤塌陷影响下植物能量与水分交换动态的基础,而采煤矿区植物叶片气孔导度、蒸腾与光合作用速率对采煤塌陷影响下土壤含水量变化的响应如何尚不清楚。选取神东煤田大柳塔矿区52302工作面为实验场地,以生态修复物种柠条为研究对象,对采煤塌陷区和对照区柠条叶片气孔导度、蒸腾和光合作用速率以及土壤体积含水量进行监测,分析了采煤塌陷条件下土壤含水量的变化以及其对柠条叶片气孔导度、蒸腾与光合作用速率的影响。结果显示:(1)煤炭井工开采在地表形成大量裂缝,破坏了土体结构,潜水位埋深降低,土壤含水量均低于沉陷初期,相对于对照区,硬梁和风沙塌陷区土壤含水量分别降低了18.61%、21.12%;(2)柠条叶片气孔导度、蒸腾和光合作用速率均与土壤含水量呈正相关关系;煤炭开采沉陷增加了地表水分散失,加剧了土壤水分胁迫程度,为了减少蒸腾导致的水分散失,柠条叶片气孔阻力增加,从而气孔导度降低,阻碍了光合作用CO_2的供应,从而导致柠条叶片光合作用速率的降低,蒸腾速率也显著降低。     

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.     

Parameterization of the CO2 and H2O gas exchange of several temperate deciduous broad-leaved trees at the leaf scale considering seasonal changes

A combined model to simulate CO₂ and H₂O gas exchange at the leaf scale was parameterized using data obtained from in situ leaf‐scale observations of diurnal and seasonal changes in the CO₂ and H₂O gas exchange of four temperate deciduous broad‐leaved trees using a porometric method. The model consists of a Ball et al. type stomatal conductance submodel [Ball, Woodrow & Berry, pp. 221–224 in Progress in Photosynthesis Research (ed. I. Biggins), Martinus‐Nijhoff Publishers, Dordrecht, The Netherlands, 1987] and a Farquhar et al. type biochemical submodel of photosynthesis (Farquhar, von Caemmerer & Berry, Planta 149, 78–90, 1980). In these submodels, several parameters were optimized for each tree species as representative of the quantitative characteristics related to gas exchange. The results show that the seasonal physiological changes of V_cmax25 in the biochemical model of photosynthesis should be used to estimate the long‐term CO₂ gas exchange. For R_d25 in the biochemical model of photosynthesis and m in the Ball et al. type stomatal conductance model, the difference should be counted during the leaf expansion period.     

Simultaneous and independent effects of abscisic acid on stomata and the photosynthetic apparatus in whole leaves

(±)-Abscisic acid (ABA) at 10^-5 M was added to the transpiration stream of leaves of 16 species (C₃ and C₄, monocotyledons and dicotyledons). Stomatal responses followed one of three patterns: i) stomata that were wide and insensitive to CO₂ initially, closed partially and became sensitive to CO₂; ii) for stomata that were sensitive to CO₂ before the application of ABA, the range of highest sensitivity to CO₂ shifted from high to low intercellular partial pressures of CO₂, for instance in leaves of Zea mays from 170–350 to 70–140 bar; iii) when stomata responded strongly to ABA, their conductance was reduced to a small fraction of the initial conductance, and sensitivity to CO₂ was lost. The photosynthetic apparatus was affected by applications of ABA to various degrees, from no response at all (in agreement with several previous reports on the absence of effects of ABA on photosynthesis) through a temporary decrease of its activity to a lasting reduction. Saturation curves of photosynthesis with respect to the partial pressure of CO₂ in the intercellular spaces indicated that application of ABA could produce three phenomena: i) a reduction of the initial slope of the saturation curve (which indicates a diminished carboxylation efficiency); ii) a reduction of the level of the CO₂-saturated rate of assimilation (which indicates a reduction of the ribulose-1,5-bisphosphate regeneration capacity); and iii) an increase of the CO₂ compensation point. Photosynthesis of isolated mesophyll cells was not affected by ABA treatments. Responses of the stomatal and photosynthetic apparatus were usually synchronous and often proportional to each other, with the result that the partial pressure of CO₂ in the intercellular spaces frequently remained constant in spite of large changes in conductance and assimilation rate. Guard cells and the photosynthetic apparatus were able to recover from effects of ABA applications while the ABA supply continued. Recovery was usually partial, in the case of the photosynthetic apparatus occasionally complete. Abscisic acid did not cause stomatal closure or decreases in the rate of photosynthesis when it was applied during a phase of stomatal opening and induction of photosynthesis that followed a transition from darkness to light.Abbreviations and symbols A rate of CO₂ assimilation - ABA (±)-abscisic acid - c _a partial pressure of CO₂ in the ambient air or in the gas supplied to the leaf chambers - c _i partial pressure of CO₂ in the intercellular spaces of a leaf - e _a partial pressure of H₂O in the air - g conductance for water vapor - J quantum flux - T ₁ leaf temperature     

A coupled model of photosynthesis-transpiration based on the stomatal behavior for maize (Zea mays L.) grown in the field

The study presents a theoretical basis of a stomatal behavior-based coupled model for estimating photosynthesis, A, and transpiration, E. Outputs of the model were tested against data observed in a maize (Zea mays L.) field. The model was developed by introducing the internal conductance, g _ic, to CO₂ assimilation, and the general equation of stomatal conductance, g _sw, to H₂O diffusion, into models of CO₂ and H₂O diffusion through the stomata of plant leaves. The coupled model is easier for practical use since the model only includes environmental variables, such as ambient CO₂ concentration, leaf temperature, humidity and photosynthetic photon flux received at the leaves within the canopy. Moreover, concept of g _ic, and factors controlling A and E were discussed, and applicability of the model was examined with the data collected in the maize field.     

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.     

Diffusional conductance to CO2 is the key limitation to photosynthesis in salt‐stressed leaves of rice (Oryza sativa)

Salinity significantly limits leaf photosynthesis but the factors causing the limitation in salt‐stressed leaves remain unclear. In the present work, photosynthetic and biochemical traits were investigated in four rice genotypes under two NaCl concentration (0 and 150 mM) to assess the stomatal, mesophyll and biochemical contributions to reduced photosynthetic rate (A) in salt‐stressed leaves. Our results indicated that salinity led to a decrease in A, leaf osmotic potential, electron transport rate and CO₂ concentrations in the chloroplasts (C_c) of rice leaves. Decreased A in salt‐stressed leaves was mainly attributable to low C_c, which was determined by stomatal and mesophyll conductance. The increased stomatal limitation was mainly related to the low leaf osmotic potential caused by soil salinity. However, the increased mesophyll limitation in salt‐stressed leaves was related to both osmotic stress and ion stress. These findings highlight the importance of considering mesophyll conductance when developing salinity‐tolerant rice cultivars.