A stomatal optimization theory to describe the effects of atmospheric CO2 on leaf photosynthesis and transpiration

Background and Aims

Global climate models predict decreases in leaf stomatal conductance and transpiration due to increases in atmospheric CO₂. The consequences of these reductions are increases in soil moisture availability and continental scale run-off at decadal time-scales. Thus, a theory explaining the differential sensitivity of stomata to changing atmospheric CO₂ and other environmental conditions must be identified. Here, these responses are investigated using optimality theory applied to stomatal conductance.

Methods

An analytical model for stomatal conductance is proposed based on: (a) Fickian mass transfer of CO₂ and H₂O through stomata; (b) a biochemical photosynthesis model that relates intercellular CO₂ to net photosynthesis; and (c) a stomatal model based on optimization for maximizing carbon gains when water losses represent a cost. Comparisons between the optimization-based model and empirical relationships widely used in climate models were made using an extensive gas exchange dataset collected in a maturing pine (Pinus taeda) forest under ambient and enriched atmospheric CO₂.

Key Results and Conclusion

In this interpretation, it is proposed that an individual leaf optimally and autonomously regulates stomatal opening on short-term (approx. 10-min time-scale) rather than on daily or longer time-scales. The derived equations are analytical with explicit expressions for conductance, photosynthesis and intercellular CO₂, thereby making the approach useful for climate models. Using a gas exchange dataset collected in a pine forest, it is shown that (a) the cost of unit water loss λ (a measure of marginal water-use efficiency) increases with atmospheric CO₂; (b) the new formulation correctly predicts the condition under which CO₂-enriched atmosphere will cause increasing assimilation and decreasing stomatal conductance.     

Changes in gas exchange characteristics and water use efficiency of mangroves in response to salinity and vapour pressure deficit

Summary Measurements were made of the photosynthetic gas exchange properties and water use efficiency of 19 species of mangrove in 9 estuaries with different salinity and climatic regimes in north eastern Australia and Papua New Guinea. Stomatal conductance and CO₂ assimilation rates differed significantly between species at the same locality, with the salt-secreting species, Avicennia marina, consistently having the highest CO₂ assimilation rates and stomatal conductances. Proportional changes in stomatal conductance and CO₂ assimilation rate resulted in constant and similar intercellular CO₂ concentrations for leaves exposed to photon flux densities above 800 mol·m^-2·s^-1 in all species at a particular locality. In consequence, all species at the same locality had similar water use efficiencies. There were, however, significant differences in gas exchange properties between different localities. Stomatal conductance and CO₂ assimilation rate both decreased with increasing salinity and with increasing leaf to air vapour pressure deficit (VPD). Furthermore, the slope of the relationship between assimilation rate and stomatal conductance increased, while intercellular CO₂ concentration decreased, with increasing salinity and with decreasing ambient relative humidity. It is concluded from these results that the water use efficiency of mangroves increases with increasing environmental stress, in this case aridity, thereby maximising photosynthetic carbon fixation while minimising water loss.Contribution No. 459 from the Australian Institute of Marine Science     

Photosynthesis of cotton plants exposed to elevated levels of carbon dioxide in the field

The cotton (Gossypium hirsutum L.) plant responds to a doubling of atmospheric CO₂ with almost doubled yield. Gas exchange of leaves was monitored to discover the photosynthetic basis of this large response. Plants were grown in the field in open-top chambers with ambient (nominally 350 l/l) or enriched (nominally either 500 or 650 l/l) concentrations of atmospheric CO₂. During most of the season, in fully-irrigated plants the relationship between assimilation (A) and intercellular CO₂ concentration (c_i) was almost linear over an extremely wide range of c_i. CO₂ enrichment did not alter this relationship or diminish photosynthetic capacity (despite accumulation of starch to very high levels) until very late in the season, when temperature was somewhat lower than at midseason. Stomatal conductance at midseason was very high and insensitive to CO₂, leading to estimates of c_i above 85% of atmospheric CO₂ concentration in both ambient and enriched chambers. Water stress caused A to show a saturation response with respect to c_i, and it increased stomatal closure in response to CO₂ enrichment. In fully-irrigated plants CO₂ enrichment to 650 l/l increased A more than 70%, but in water-stressed plants enrichment increased A only about 52%. The non-saturating response of A to c_i, the failure of CO₂ enrichment to decrease photosynthetic capacity for most of the season, and the ability of the leaves to maintain very high c_i, form in part the basis for the very large response to CO₂ enrichment.Abbreviations c_a- atmospheric CO₂ concentration - c_i- intercellular CO₂ concentration - A- rate of assimilation of CO₂ - g_s- stomatal conductance to water vapor - g_b- boundary layer conductance to water vapor - g_m- mesophyll conductance to CO₂ - VPD- vapor pressure deficit - _w leaf water potential - L- stomatal limitation to CO₂ uptake     

Control of photosynthesis by the carbohydrate level in leaves of the C4 plant Amaranthus edulis L.

Photosynthesis was studied in relation to the carbohydrate status in intact leaves of the C₄ plant Amaranthus edulis. The rate of leaf net CO₂ assimilation, stomatal conductance and intercellular partial pressure of CO₂ remained constant or showed little decline towards the end of an 8-h period of illumination in ambient air (340 bar CO₂, 21% O₂). When sucrose export from the leaf was inhibited by applying a 4-h cold-block treatment (1°C) to the petiole, the rate of photosynthesis rapidly decreased with time. After the removal of the cold block from the petiole, further reduction in photosynthetic rate occurred, and there was no recovery in the subsequent light period. Although stomatal conductance declined with time, intercellular CO₂ partial pressure remained relatively constant, indicating that the inhibition of photosynthesis was not primarily caused by changes in stomatal aperture. Analysis of the leaf carbohydrate status showed a five- to sixfold increase in the soluble sugar fraction (mainly sucrose) in comparison with the untreated controls, whereas the starch content was the same. Leaf osmotic potential increased significantly with the accumulation of soluble sugars upon petiole chilling, and leaf water potential became slightly more negative. After 14 h recovery in the dark, photosynthesis returned to its initial maximum value within 1 h of illumination, and this was associated with a decline in leaf carbohydrate levels overnight. These data show that, in Amaranthus edulis, depression in photosynthesis when translocation is impaired is closely related to the accumulation of soluble sugars (sucrose) in source leaves, indicating feedback control of C₄ photosynthesis. Possible mechanisms by which sucrose accumulation in the leaf may affect the rate of photosynthesis are discussed with regard to the leaf anatomy of C₄ plants.Abbreviations and symbols A net CO₂ assimilation rate - Ci intercellular CO₂ partial pressure - PEP phosphoenolpyruvate - RuBP ribulose-1,5-bisphosphate - water potential - osmotic pressure     

The drelationship between CO2-assimilation rate,Rubisco carbamylation and Rubisco activase content in activase-deficient transgenic tobacco suggests a simple model of activase action

Transgenic tobacco (Nicotiana tabacum L. cv. W38) plants with an antisense gene directed against the mRNA of ribulose-1,5-bisphosphate carboxylase/ oxygenase (Rubisco) activase were used to examine the relationship between CO₂-assimilation rate, Rubisco carbamylation and activase content. Plants used were those members of the r₁ progeny of a primary transformant with two independent T-DNA inserts that could be grown without CO₂ supplementation. These plants had from < 1% to 20% of the activase content of control plants. Severe suppression of activase to amounts below 5% of those present in the controls was required before reductions in CO₂-assimilation rate and Rubisco carbamylation were observed, indicating that one activase tetramer is able to service as many as 200 Rubisco hexadecamers and maintain wild-type carbamylation levels in vivo. The reduction in CO₂-assimilation rate was correlated with the reduction in Rubisco carbamylation. The anti-activase plants had similar ribulose-1,5-bisphosphate pool sizes but reduced 3-phosphoglycerate pool sizes compared to those of control plants. Stomatal conductance was not affected by reduced activase content or CO₂-assimilation rate. A mathematical model of activase action is used to explain the observed hyperbolic dependence of Rubisco carbamylation on activase content.Abbreviations CA1P 2-carboxyarabinitol-1-phosphate - P_ip_a intercellular, ambient partial pressure of CO₂ - PGA 3-phospho-glycerate - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase - RuBP ribulose-1,5-bisphosphate - SSU small subunit of Rubisco     

Plant water relations and the effects of elevated CO2: a review and suggestions for future research

Increased ambient carbon dioxide (CO₂) has been found to ameliorate water stress in the majority of species studied. The results of many studies indicate that lower evaporative flux density is associated with high CO₂-induced stomatal closure. As a result of decreases in evaporative flux density and increases in net photosynthesis, also found to occur in high CO₂ environments, plants have often been shown to maintain higher water use efficiencies when grown at high CO₂ than when grown in normal, ambient air. Plants grown at high CO₂ have also been found to maintain higher total water potentials, to increase biomass production, have larger root-to-shoot ratios, and to be generally more drought resistant (through avoidance mechanisms) than those grown at ambient CO₂ levels. High CO₂-induced changes in plant structure (i.e., vessel or tracheid anatomy, leaf specific conductivity) may be associated with changes in vulnerability to xylem cavitation or in environmental conditions in which runaway embolism is likely to occur. Further study is needed to resolve these important issues. Methodology and other CO₂ effects on plant water relations are discussed.Abbreviations A net photosynthesis - C_a ambient [CO₂] - C_i internal [CO₂] - E evaporative flux density - g₁ leaf conductance - g_s stomatal conductance - LSC leaf specific conductivity - IRGA infrared gas analyzer - LAI leaf area index - PAR photosynthetically active radiation - total plant water potential - _soil soil water potential - _s solute potential - _pt turgor pressure potential - _px xylem pressure potential - RH relative humidity - R : S root to shoot ratio - RWC relative water content - SLA specific leaf area - SLW specific leaf weight - SPAC soil-plant-atmosphere-continuum - SWC soil water content - VPD vapor pressure deficit - WUE water use efficiency     

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     

Photosynthesis,water use and growth of a C4 grass stand at high CO2 concentration

Leaf photosynthesis rate of the C₄ species Paspalum plicatulum Michx was virtually CO₂-saturated at normal atmospheric CO₂ concentration but transpiration decreased as CO₂ was increased above normal concentrations thereby increasing transpiration efficiency. To test whether this leaf response led growth to be CO₂-sensitive when water supply was restricted, plants were grown in sealed pots of soil as miniature swards. Water was supplied either daily to maintain a constant water table, or at three growth restricting levels on a 5-day drying cycle. Plants were either in a cabinet with normal air (340 mol (CO₂) mol^-1 (air)) or with 250 mol mol^-1 enrichment. Harvesting was by several cycles of defoliation.With abundant water supply high CO₂ concentration did not cause increased growth, but it did not cause an increase in growth over a wide range of growth-limiting water supplies either. Only when water supply was less than 30–50% of the amount used by the stand with a water-table was there evidence that dry weight growth was enhanced by high CO₂. In addition, with successive regrowth, the enhancing effect under a regime of minimal water allocations, became attenuated. Examination of leaf gas exchange, growth and water use data showed that in the long term stomatal conductance responses were of little significance in matching plant water use to low water allocation; regulation of leaf area was the mechanism through which consumption matched supply. Since high CO₂ effects operate principally via stomatal conductance in C₄ species, we postulate that for this species higher CO₂ concentrations expected globally in future will not have much effect on long term growth.     

Leaf and canopy responses to elevated CO2 in a pine forest under free-air CO2 enrichment

Physiological responses to elevated CO₂ at the leaf and canopy-level were studied in an intact pine (Pinus taeda) forest ecosystem exposed to elevated CO₂ using a free-air CO₂ enrichment (FACE) technique. Normalized canopy water-use of trees exposed to elevated CO₂ over an 8-day exposure period was similar to that of trees exposed to current ambient CO₂ under sunny conditions. During a portion of the exposure period when sky conditions were cloudy, CO₂-exposed trees showed minor (7%) but significant reductions in relative sap flux density compared to trees under ambient CO₂ conditions. Short-term (minutes) direct stomatal responses to elevated CO₂ were also relatively weak (5% reduction in stomatal aperture in response to high CO₂ concentrations). We observed no evidence of adjustment in stomatal conductance in foliage grown under elevated CO₂ for nearly 80 days compared to foliage grown under current ambient CO₂, so intrinsic leaf water-use efficiency at elevated CO₂ was enhanced primarily by direct responses of photosynthesis to CO₂. We did not detect statistical differences in parameters from photosynthetic responses to intercellular CO₂ (A _net-C _i curves) for Pinus taeda foliage grown under elevated CO₂ (550 mol mol^–1) for 50–80 days compared to those for foliage grown under current ambient CO₂ from similar-sized reference trees nearby. In both cases, leaf net photosynthetic rate at 550 mol mol^–1 CO₂ was enhanced by approximately 65% compared to the rate at ambient CO₂ (350 mol mol^–1). A similar level of enhancement under elevated CO₂ was observed for daily photosynthesis under field conditions on a sunny day. While enhancement of photosynthesis by elevated CO₂ during the study period appears to be primarily attributable to direct photosynthetic responses to CO₂ in the pine forest, longer-term CO₂ responses and feedbacks remain to be evaluated.     

Seasonal trends in leaf photosynthesis and stomatal conductance of drought stressed and nonstressed pearl millet as associated to vapor pressure deficit

Single leaf photosynthesis (Pn) and stomatal conductance (Cg) of drought stressed and nonstressed pearl millet [Pennisetum americanum (L.) Leeke] were measured across growth stages to determine if a pattern exists in Pn and Cg during the growing season and to evaluate the influence of air vapor pressure deficit (VPD_a) on the seasonal variations of Pn and Cg. Leaf photosynthesis and Cg were measured independently on pearl millet plants grown at the driest (drought stressed) and wettest (nonstressed) ends of a line-source irrigation gradient system. Well defined and predictable variations in both Pn and Cg were found across two growing seasons. Leaf photosynthesis of the nonstressed plants declined from a maximumof 25.8 mol m^–2 s^–1 at the flag leaf emergence (48 days after planting, DAP) to a minimum of 14.5 mol m^–2 s^–1 at physiological maturity. Stomatal conductance of the nonstressed plants peaked at the flowering and early grain fill stages and declined as plants approached maturity. In contrast, Pn and Cg of the stressed plants declined from a maximum at flag leaf emergence to a minimum at flowering and increased as plants approached maturity. High VPD_a during the flowering and grain fill stages induced stomatal closure and decreased Pn in the stressed plants. High mid-season VPD_a did not induce stomatal closure and did not reduce leaf photosynthesis in nonstressed plants. The lack of sensitivity of Pn to VPD_a in the nonstressed treatment suggests large air VPD such as that prevalent in southern Arizona does not limit the growth of irrigated pearl millet by limiting CO₂ assimilation.Abbreviations Cg stomatal conductance - DAP days after planting - Pn leaf photosynthesis - VPD_a air vapor pressure deficit - VPD_1-a leaf to air vapor pressure deficit Contribution of the Arizona Agricultural Experimental Station. Research supported in part by INTSORMIL/USAID.     

Elevated CO2 alleviates the impact of drought on barley improving water status by lowering stomatal conductance and delaying its effects on photosynthesis

We analysed the impact of elevated CO₂ on water relations, water use efficiency and photosynthetic gas exchange in barley (Hordeum vulgare L.) under wet and drying soil conditions. Soil moisture was less depleted under elevated compared to ambient [CO₂]. Elevated CO₂ had no significant effect on the water relations of irrigated plants, except on whole plant hydraulic conductance, which was markedly decreased at elevated compared to ambient CO₂ concentrations. The values of relative water content, water potential and osmotic potential were higher under elevated CO₂ during the entire drought period. The better water status of water-limited plants grown at elevated CO₂ was the result of stomatal control rather than of osmotic adjustment. Despite the low stomatal conductance produced by elevated CO₂, net photosynthesis was higher under elevated than ambient CO₂ concentrations. With water shortage, photosynthesis was maintained for longer at higher rates under elevated CO₂. The reduction of stomatal conductance and therefore transpiration, and the enhancement of carbon assimilation by elevated CO₂, increased instantaneous and whole plant water use efficiency in both irrigated and droughted plants. Thus, the metabolism of barley plants grown under elevated CO₂ and moderate or mild water deficit conditions is benefited by increased photosynthesis and lower transpiration. The reduction in plant water use results in a marked increase in soil water content which delays the onset and severity of water deficit.     

Do Stomata Respond to CO(2) Concentrations Other than Intercellular?

Most studies on stomatal responses to CO₂ assume that guard cells respond only to intercellular CO₂ concentration and are insensitive to the CO₂ concentrations in the pore and outside the leaf. If stomata are sensitive to the CO₂ concentration at the surface of the leaf or in the stomatal pore, the stomatal response to intercellular CO₂ concentration will be incorrect for a `normally' operating leaf (where ambient CO₂ concentration is a constant). In this study asymmetric CO₂ concentrations for the two surfaces of amphistomatous leaves were used to vary intercellular and leaf surface CO₂ concentrations independently in Xanthium strumarium L. and Helianthus annuus L. The response of stomata to intercellular CO₂ concentration when the concentration at the leaf surface was held constant was found to be the same as the response when the surface concentration was varied. In addition, stomata did not respond to changes in leaf surface CO₂ concentration when the intercellular concentration for that surface was held constant. It is concluded that stomata respond to intercellular CO₂ concentration and are insensitive to the CO₂ concentration at the surface of the leaf and in the stomatal pore.     

Effect of increased salinity on CO2 assimilation,O2 evolution and the δ13C values of leaves of Plantago maritima L. developed at low and high NaCl levels

The effect of increased salinity on photosynthesis was studied in leaves of Plantago maritima L. that developed while plants were at low and high NaCl levels. In leaves that developed while plants were grown at 50 mol·m^-3, exposure to 200 and 350 mol·m^-3 NaCl resulted in reductions in net CO₂ assimilation and stomatal conductance. The decline in CO₂ assimilation in plants at 200 and 350 mol·m^-3 NaCl occurred almost exclusively at high intercellular CO₂ concentrations. The initial slope of the CO₂ assimilation-intercellular CO₂ (A-C _i) curve, determined after salinity was increased, was identical or very similar to that measured initially. In contrast to the reductions observed in CO₂ assimilation, there were no significant differences in O₂ evolution rates measured at 5% CO₂ among leaves from plants exposed to higher salinity and plants remaining at low salinity.Leaves that developed while plants were at increased salinity levels also had significantly lower net CO₂ assimilation rates than plants remaining at 50 mol·m^-3 NaCl. The lower CO₂ assimilation rates in plants grown at 200 and 350 mol·m^-3 NaCl were a result of reduced stomatal conductance and low intercellular CO₂ concentration. There were no significant differences among treatments for O₂ evolution rates measured at high CO₂ levels. The increased stomatal limitation of photosynthesis was confirmed by measurements of the ¹³C/¹²C composition of leaf tissue. Water-use efficiency was increased in the plants grown at high salinity.Abbreviations and symbols A net CO₂ assimilation rate - C _a ambient CO₂ concentration - C _i intercellular CO₂ concentration - ¹³C isotopic ratio (¹³C/¹²C) expressed relative to a standard - RuBP ribulose-1,5-bisphosphate     

Gas-exchange analysis of chloroplastic fructose-1,6-bisphosphatase antisense potatoes at different air humidities and at elevated CO2

Gas-exchange measurements were performed to analyze the leaf conductances and assimilation rates of potato (Solanum tuberosum L. cv. Desireé) plants expressing an antisense construct against chloroplastic fructose-1,6-bisphosphatase (FBPase, EC 3.1.3.11) in response to increasing photon flux densities, different relative air humidities and elevated CO₂ concentrations. Assimilation rates (A) and transpiration rates (E) were observed during a stepwise increase of photon flux density. These experiments were carried out under atmospheric conditions and in air containing 500 μmol mol⁻¹ CO₂. In both gas atmospheres, two levels of relative air humidity (60–70% and 70–80%) were applied in different sets of measurements. Intercellular CO₂ concentration, leaf conductance, air-to-leaf vapour pressure deficit, and instantaneous water-use efficiency (A/E) were determined. As expected, assimilation rates of the FBPase antisense plants were significantly reduced as compared to the wild type. Saturation of assimilation rates in transgenic plants occurred at a photon flux density of 200 μmol m⁻² s⁻¹, whereas saturation in wild type plants was observed at 600 μmol m⁻² s⁻¹. Elevated ambient CO₂ levels did not effect assimilation rates of transgenic plants. At 70–80% relative humidity and atmospheric CO₂ concentration the FBPase antisense plants had significantly higher leaf conductances than wild-type plants while no difference emerged at 60–70%. These differences in leaf conductance vanished at elevated levels of ambient CO₂. Stomatal response to different relative air humidities was not affected by mesophyll photosynthetic activity. It is suggested that the regulation of stomatal opening upon changes in photon flux density is merely mediated by a signal transmitted from mesophyll cells, whereas the intercellular CO₂ concentration plays a minor role in this kind of stomatal response. The results are discussed with respect to stomatal control by environmental parameters and mesophyll photosynthesis. Received: 24 September 1998 / Accepted: 9 February 1999     

Water stress in Eucalyptus pauciflora: comparison of effects on stomatal conductance with effects on the mesophyll capacity for photosynthesis,and investigation of a possible involvement of photoinhibition

Seedlings of Eucalyptus pauciflora Sieb. ex Spreng., grown in 4-1 pots, were stressed by withholding water while relationships between net assimilation rate (A) and intercellular partial pressure of CO₂ (p_i) in selected leaves were obtained repeatedly throughout the stress cycle. Water stress at first caused stomatal closure without any decline in the A(p_i) relationship. As stress became more severe, the A(p_i) relationship was affected as well. This always affected assimilation rate at both high and low intercellular partial pressures of CO₂. It was then tested whether water-stressed leaves were more prone to photoinhibition than unstressed ones. Plants were water-stressed while at the same time subjected to strong photon flux area density (2000 mol quanta·m^-2·s^-1). A possible light-induced inhibition was assessed by comparing quantum yields of photosynthesis with light directed onto one or the other surface of the leaf. A decline in quantum yield was observed, and the decline on the previously irradiated side was more pronounced than on the previously shaded side, but the effect was small and disappeared entirely within 1 d of rewatering the plants. It is concluded that photoinhibition can play a role, but not an important one, in the effect of water stress on the A(p_i) relationship in leaves of E. pauciflora.Abbreviations and symbols RuBP ribulose-1,5-bisphosphate - A net assimilation rate - p_i intercellular partial pressure of CO₂ - quantum yield of photosynthesis (net assimilation or RuBP-regeneration rate) - w difference in water content between air saturated at leaf temperature and the actual vapor content of the air, expressed as mole fraction     

Partitioning of photosynthetic electron flow between CO2 and O2 reduction in a C3 leaf (Phaseolus vulgaris L.) at different CO2 concentrations and during drought stress

Photosystem II chlorophyll fluorescence and leaf net gas exchanges (CO₂ and H₂O) were measured simultaneously on bean leaves (Phaseolus vulgaris L.) submitted either to different ambient CO₂ concentrations or to a drought stress. When leaves are under photorespiratory conditions, a simple fluorescence parameter F/ F_m (B. Genty et al. 1989, Biochem. Biophys. Acta 990, 87–92; F = difference between maximum, F_m, and steady-state fluorescence emissions) allows the calculation of the total rate of photosynthetic electron-transport and the rate of electron transport to O₂. These rates are in agreement with the measurements of leaf O₂ absorption using ¹⁸O₂ and the kinetic properties of ribulose-1,5bisphosphate carboxylase/oxygenase. The fluorescence parameter, F/F_m, showed that the allocation of photosynthetic electrons to O₂ was increased during the desiccation of a leaf. Decreasing leaf net CO₂ uptake, either by decreasing the ambient CO₂ concentration or by dehydrating a leaf, had the same effect on the partitioning of photosynthetic electrons between CO₂ and O₂ reduction. It is concluded that the decline of net CO₂ uptake of a leaf under drought stress is only due, at least for a mild reversible stress (causing at most a leaf water deficit of 35%), to stomatal closure which leads to a decrease in leaf internal CO₂ concentration. Since, during the dehydration of a leaf, the calculated internal CO₂ concentration remained constant or even increased we conclude that this calculation is misleading under such conditions.Abbreviations Ca, Ci ambient, leaf internal CO₂ concentrations - F_m, F_o, F_s maximum, minimal, steady-state fluorescence emission - F_v variable fluorescence emission - PPFD photosynthetic photon flux density - q_p, q_N photochemical, non-photochemical fluorescence quenching - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase     

Performance of two Picea abies (L.) Karst. stands at different stages of decline

Summary Photosynthetic rates and nutrient contents of spruce needles were measured in a region with high levels of air pollution in NE Bavaria, Germany (FRG), and compared to spruce grown under clean air conditions at Craigieburn, in the South Island of New Zealand (NZ). The absolute rates of CO₂ uptake, the slope of the CO₂ response curve at 240 l l^–1 internal CO₂ concentration, and the change of photosynthetic rates with needle age at ambient and saturated CO₂ concentrations were virtually identical at both measuring sites. These results confirm an earlier conclusion, that there is no long-term effect of atmospheric pollutants directly on photosynthetic CO₂ uptake rates with persistent exposure at the FRG site to high levels of anthropogenic air pollution. Photosynthetic capacity at saturating CO₂ concentration was three times higher in the NZ spruce. Needles with high photosynthetic capacity in NZ had lower nitrogen and higher calcium concentrations per unit dry weight but higher concentrations of nitrogen, phosphorus, potassium, magnesium and calcium per unit leaf area, and twice the specific leaf weight.     

Anatomy of non-uniform leaf photosynthesis

Since 1986, non-uniform photosynthesis over the leaf area that may be attributed to patchy stomatal closure, has been an important issue in stress physiology of photosynthesis. In this review, I first outline the gaseous environment within the intercellular spaces, because this is the most fundamental background of this problem. Then, recent studies approaching non-uniform photosynthesis are reviwed. After examining techniques for the detection of non-uniform photosynthesis or non-uniform stomatal aperture, results of the relevant studies are discussed for respective stress factors, seeking causes and consequences of non-uniform photosynthesis. From these, mechanisms responsible for, and consequences of non-uniform photosynthesis, are considered. The problems which should be challenged are also pointed out.Abbreviations A rate of photosynthetic CO₂ fixation (assimilation rate) - ABA abscisic acid - g conductance for CO₂ or H₂O - p_a partial pressure of CO₂ in the ambient air - p_i partial pressure of CO₂ in the intercellular space - p_i p_i estimated by the conventional gas exchange techniques - q_N non-photochemical quenching - q_p photochemical quenching     

Stomatal conductance in mature deciduous forest trees exposed to elevated CO<Subscript>2</Subscript>

Stomatal conductance (g _s) of mature trees exposed to elevated CO₂ concentrations was examined in a diverse deciduous forest stand in NW Switzerland. Measurements of g _s were carried out on upper canopy foliage before noon, over four growing seasons, including an exceptionally dry summer (2003). Across all species reductions in stomatal conductance were smaller than 25% most likely around 10%, with much variation among species and trees. Given the large heterogeneity in light conditions within a tree crown, this signal was not statistically significant, but the responses within species were surprisingly consistent throughout the study period. Except during a severe drought, stomatal conductance was always lower in trees of Carpinus betulus exposed to elevated CO₂ compared to Carpinus trees in ambient air, but the difference was only statistically significant on 2 out of 15 days. In contrast, stomatal responses in Fagus sylvatica and Quercus petraea varied around zero with no consistent trend in relation to CO₂ treatment. During the 2003 drought in the third treatment year, the CO₂ effect became reversed in Carpinus, resulting in higher g _s in trees exposed to elevated CO₂ compared to control trees, most likely due to better water supply because of the previous soil water savings. This was supported by less negative predawn leaf water potential in CO₂ enriched Carpinus trees, indicating an improved water status. These findings illustrate (1) smaller than expected CO₂-effects on stomata of mature deciduous forest trees, and (2) the possibility of soil moisture feedback on canopy water relations under elevated CO₂.     

Gas exchange of Ginkgo biloba leaves at different CO2 concentration levels

One and a half year-old Ginkgo saplings were grown for 2 years in 7 litre pots with medium fertile soil at ambient air CO₂ concentration and at 700 μmol mol⁻¹ CO₂ in temperature and humidity-controlled cabinets standing in the field. In the middle of the 2nd season of CO₂ enrichment, CO₂ exchange and transpiration in response to CO₂ concentration was measured with a mini-cuvette system. In addition, the same measurements were conducted in the crown of one 60-year-old tree in the field. Number of leaves/tree was enhanced by elevated CO₂ and specific leaf area decreased significantly.CO₂ compensation points were reached at 75–84 μmol mol⁻¹ CO₂. Gas exchange of Ginkgo saplings reacted more intensively upon CO₂ than those of the adult Ginkgo. On an average, stomatal conductance decreased by 30% as CO₂ concentration increased from 30 to 1000 μmol mol⁻¹ CO₂. Water use efficiency of net photosynthesis was positively correlated with CO₂ concentration levels. Saturation of net photosynthesis and lowest level of stomatal conductance was reached by the leaves of Ginkgo saplings at >1000 μmol mol⁻¹ CO₂. Acclimation of leaf net CO₂ assimilation to the elevated CO₂ concentration at growth occurred after 2 years of exposure. Maximum of net CO₂ assimilation was 56% higher at ambient air CO₂ concentration than at 700 μmol mol⁻¹ CO₂.