Carboxylation Capacity Research Articles

Effects of chronic elevated ozone exposure on gas exchange responses of adult beech trees ( Fagus sylvatica) as related to the within-canopy light gradient

The effects of elevated O 3 on photosynthetic properties in adult beech trees ( Fagus sylvatica) were investigated in relation to leaf mass per area as a measure of the gradually changing, within-canopy light availability. Leaves under elevated O 3 showed decreased stomatal conductance at unchanged carboxylation capacity of Rubisco, which was consistent with enhanced δ 13C of leaf organic matter, regardless of the light environment during growth. In parallel, increased energy demand for O 3 detoxification and repair was suggested under elevated O 3 owing to enhanced dark respiration. Only in shade-grown leaves, light-limited photosynthesis was reduced under elevated O 3, this effect being accompanied by lowered F v/ F m. These results suggest that chronic O 3 exposure primarily caused stomatal closure to adult beech trees in the field regardless of the within-canopy light gradient. However, light limitation apparently raised the O 3 sensitivity of photosynthesis and accelerated senescence in shade leaves.

A model for chlorophyll fluorescence and photosynthesis at leaf scale

This paper presents a leaf biochemical model for steady-state chlorophyll fluorescence and photosynthesis of C3 and C4 vegetation. The model is a tool to study the relationship between passively measured steady-state chlorophyll fluorescence and actual photosynthesis, and its evolution during the day. Existing models for chlorophyll fluorescence and photosynthesis are integrated into a relatively simple deterministic model to quantify chlorophyll fluorescence, electron transport, carboxylation and deactivation of antennae in case of light and moisture stress. The model explains the behaviour of the relationship between fluorescence and photosynthesis that has been reported in the literature. Simulations, a sensitivity analysis, and measurements show that variations in total chlorophyll fluorescence correlate well with variations in actual photosynthesis in the late morning and afternoon. Then, photosynthesis is light saturated and limited by stomatal regulation. To calculate the actual photosynthesis rate, an estimate of the maximum carboxylation capacity is needed beside chlorophyll fluorescence.

Elevated CO 2 and water-availability effect on gas exchange and nodule development in N 2-fixing alfalfa plants

N 2-fixing alfalfa plants were grown in controlled conditions at different CO 2 levels (350 μmol mol −1 versus 700 μmol mol −1) and water-availability conditions (WW, watered at maximum pot water capacity versus WD, watered at 50% of control treatments) in order to determine the CO 2 effect (and applied at two water regimes) on plant growth and nodule activity in alfalfa plants. The CO 2 stimulatory effect (26% enhancement) on plant growth was limited to WW plants, whereas no CO 2 effect was observed in WD plants. Exposure to elevated CO 2 decreased Rubisco carboxylation capacity of plants, caused by a specific reduction in Rubisco (EC 4.1.1.39) concentration (11% in WW and 43% in WD) probably explained by an increase in the leaf carbohydrate levels. Plants grown at 700 μmol mol −1 CO 2 maintained control photosynthetic rates (at growth conditions) by diminishing Rubisco content and by increasing nitrogen use efficiency. Interestingly, our data also suggest that reduction in shoot N demand (reflected by the TSP and especially Rubisco depletion) affected negatively nodule activity (malate dehydrogenase, EC 1.1.1.37, and glutamate-oxaloacetate transaminase, EC 2.6.1.1, activities) particularly in water-limited conditions. Furthermore, nodule DM and TSS data revealed that those nodules were not capable to overcome C sink strength limitations.

Investigating a Hierarchy of Eulerian Closure Models for Scalar Transfer Inside Forested Canopies

Modelling the transfer of heat, water vapour, and CO2 between the biosphere and the atmosphere is made difficult by the complex two-way interaction between leaves and their immediate microclimate. When simulating scalar sources and sinks inside canopies on seasonal, inter-annual, or forest development time scales, the so-called well-mixed assumption (WMA) of mean concentration (i.e. vertically constant inside the canopy but dynamically evolving in time) is often employed. The WMA eliminates the need to model how vegetation alters its immediate microclimate, which necessitates formulations that utilize turbulent transport theories. Here, two inter-related questions pertinent to the WMA for modelling scalar sources, sinks, and fluxes at seasonal to inter-annual time scales are explored: (1) if the WMA is to be replaced so as to resolve this two-way interaction, how detailed must the turbulent transport model be? And (2) what are the added predictive skills gained by resolving the two-way interaction vis-a-vis other uncertainties such as seasonal variations in physiological parameters. These two questions are addressed by simulating multi-year mean scalar concentration and eddy-covariance scalar flux measurements collected in a Loblolly pine (P. taeda L.) plantation near Durham, North Carolina, U.S.A. using turbulent transport models ranging from K-theory (or first-order closure) to third-order closure schemes. The multi-layer model calculations with these closure schemes were contrasted with model calculations employing the WMA. These comparisons suggested that (i) among the three scalars, sensible heat flux predictions are most biased with respect to eddy-covariance measurements when using the WMA, (ii) first-order closure schemes are sufficient to reproduce the seasonal to inter-annual variations in scalar fluxes provided the canonical length scale of turbulence is properly specified, (iii) second-order closure models best agree with measured mean scalar concentration (and temperature) profiles inside the canopy as well as scalar fluxes above the canopy, (iv) there are no clear gains in predictive skills when using third-order closure schemes over their second-order closure counterparts. At inter-annual time scales, biases in modelled scalar fluxes incurred by using the WMA exceed those incurred when correcting for the seasonal amplitude in the maximum carboxylation capacity (V cmax, 25) provided its mean value is unbiased. The role of local thermal stratification inside the canopy and possible computational simplifications in decoupling scalar transfer from the generation of the flow statistics are also discussed. “The tree, tilting its leaves to capture bullets of light; inhaling, exhaling; its many thousand stomata breathing, creating the air”. Ruth Stone, 2002, In the Next Galaxy

Elevated CO2 concentration affects leaf photosynthesis-nitrogen relationships in Pinus taeda over nine years in FACE

To investigate whether long-term elevated carbon dioxide concentration ([CO(2)]) causes declines in photosynthetic enhancement and leaf nitrogen (N) owing to limited soil fertility, we measured photosynthesis, carboxylation capacity and area-based leaf nitrogen concentration (N(a)) in Pinus taeda L. growing in a long-term free-air CO(2) enrichment (FACE) facility at an N-limited site. We also determined how maximum rates of carboxylation (V(cmax)) and electron transport (J(max)) varied with N(a) under elevated [CO(2)]. In trees exposed to elevated [CO(2)] for 5 to 9 years, the slope of the relationship between leaf photosynthetic capacity (A(net-Ca)) and N(a) was significantly reduced by 37% in 1-year-old needles, whereas it was unaffected in current-year needles. The slope of the relationships of both V(cmax) and J(max) with N(a) decreased in 1-year-old needles after up to 9 years of growth in elevated [CO(2)], which was accompanied by a 15% reduction in N allocation to the carboxylating enzyme. Nitrogen fertilization (110 kg N ha(-1)) in the ninth year of exposure to elevated [CO(2)] restored the slopes of the relationships of V(cmax) and J(max) with N(a) to those of control trees (i.e., in ambient [CO(2)]). The J(max):V(cmax) ratio was unaffected by either [CO(2)] or N fertilization. Changes in the apparent allocation of N to photosynthetic components may be an important adjustment in pines exposed to elevated [CO(2)] on low-fertility sites. We conclude that fundamental relationships between photosynthesis or its component processes with N(a) may be altered in aging pine needles after more than 5 years of exposure to elevated atmospheric [CO(2)].

Isoprene Emission and Carbon Dioxide Protect Aspen Leaves from Heat Stress

AbstractHigh temperature, especially above 35oC, is known to reduce leaf photosynthetic rate in many tree species. This study investigated the effect of high temperature on isoprene-emitting (aspen) and non- emitting (birch) trees under ambient and elevated CO2 under open field conditions. Aspen trees tolerate heat better than birch trees and elevated CO2 protects both species against moderate heat stress. The increased thermotolerance in aspen trees compared to the birch trees may result from the aspen's ability to produce isoprene. Elevated CO2 increased carboxylation capacity, photosynthetic electron transport capacity and triose phosphate use in both birch and aspen trees. High temperature decreased all of these parameters in birch regardless of CO2 treatment but only photosynthetic electron transport and triose phosphate use at ambient CO2 were reduced in aspen. As temperature rises, non-isoprene-emitting trees will be at a disadvantage and biological diversity and species richness might be lost in some ecosystems. Our results indicate that isoprene emitting tree species will have an advantage over non-isoprene emitting ones under high temperatures.

Response of Leaf Photosynthesis to Vapor Pressure Difference in Rice (Oryza sativa L) Varieties in Relation to Stomatal and Leaf Internal Conductance

In the afternoon when air humidity decreases, leaf photosynthetic rate (Pn) often declines in rice grown under irrigated conditions. To clarify the genotypic difference of Pn in response to humidity, we measured Pn and stomatal conductance (gs) for nine rice varieties with diverse genetic backgrounds, at various vapor pressure differences (VPD) and developmental stages. Pn and gs of all the varieties decreased with VPD increase from 1.0 to 2.3 kPa of VPD. The variety with high gs at low VPD exhibited a greater decline of gs with VPD increase than the variety with low gs, but cv. Takanari showed the highest gs under altered VPD conditions. Significant logarithmic relations were found between the decreased Pn and gs at the respective developmental stages, suggesting that gs is the dominant factor determining Pn and its response to VPD change. To explicate the effect of decreased gs on Pn, we analyzed the relations by using the model that accurately estimated the genotypic difference in Pn at a low VPD with gs and leaf nitrogen content per unit leaf area in the previous study. The model assuming that leaf internal conductance (gw) remains unchanged well explained the decreased Pn at high VPDs by gs change alone. The analysis also suggested the constancy of gw and carboxylation capacity at high VPD. It is concluded that the genotypic difference in the decrease of Pn at a high VPD is brought mainly by that in decreased gs, and the varieties with a high gs always exhibit a high Pn owing to their relatively high gs at either high or low VPD environments.

Carbon partitioning in N2 fixing Medicago sativa plants exposed to different CO2 and temperature conditions.

Many of the studies analysing the CO2 effect on plant development have been conducted in optimal growth conditions. Furthermore, although some of those studies suggest that legumes might show a steady productivity increase with rising CO2, the role of nodule activity on the plant responsiveness to predicted atmospheric CO2 enhancement is not well understood. In this study, C (metabolism and allocation) and N (nodule activity) interaction between the plant and the bacterial symbiont during the photosynthetic acclimation of N2-fixing alfalfa (Medicago sativa L. cv. Aragón) plants exposed to elevated CO2 and temperature conditions was analysed. The plants were grown in temperature gradient greenhouses (TGG) where, in the case of elevated CO2 treatments, the isotopic 13C/12C composition (δ13C) inside the TGG was modified. Compared with the corresponding temperature treatment, exposure to 700 μmol mol-1 CO2 enhanced dry mass (DM) of plants in elevated temperature treatments (26%), whereas no significant effect was detected in ambient temperature treatments. The δ13C data revealed that although all the carbon corresponding to leaf total organic matter (TOM) came from newly assimilated C, plants exposed to elevated CO2 did not develop strong sink activity (especially in ambient temperature conditions). Leaf carbohydrate build-up induced reduction in the Rubisco (E.C. 4.1.1.39) carboxylation capacity of plants. Despite this reduction in Rubisco content, plants exposed to elevated CO2 conditions maintained (at ambient temperature) or increased (at elevated temperature) photosynthetic rates (measured at growth conditions) by increasing N use efficiency. The larger C sink strength of nodules in plants grown at elevated CO2 and temperature conditions did not contribute towards overcoming photosynthetic acclimation. Further, the inhibitory effect of CO2 on nodule total activity was caused by a large depletion in total soluble protein (TSP) of nodules. Depletion of leaf N demand, together with the reduction in nodule carbohydrate availability (as reflected by the nodule starch concentration), negatively affected the nodule TSP content and enzymatic activity.

Elevated growth temperatures reduce the carbon gain of black spruce [Picea mariana (Mill.) B.S.P.

AbstractWe explored the effect of high‐growth temperatures on a dominant North American boreal tree, black spruce [Picea mariana (Mill.) B.S.P.]. In 2004 and 2005, we grew black spruce at either 22 °C/16 °C day/night temperatures [low temperature (LT)] or 30°/24 °C [high temperature (HT)] and determined how temperature affected growth, leaf morphology, photosynthesis, respiration and thermotolerance. HT spruce were 20% shorter, 58% lighter, and had a 58% lower root : shoot ratio than LT trees. Mortality was negligible in the LT treatment, but up to 14% of HT seedlings died by the end of the growing season. HT seedlings had a higher photosynthetic temperature optimum, but net photosynthesis at growth temperatures was 19–35% lower in HT than LT trees. HT seedlings had both a lower apparent maximum ribulose‐1,5‐bisphosphate carboxylation capacity (Vcmax) and a lower apparent maximum electron transport rate (Jmax) than LT trees, indicating reduced allocation to photosynthetic components. Consistently, HT needles had 26% lower leaf nitrogen content than LT needles. At each measurement temperature, HT seedlings had 20–25% lower respiration rates than LT trees; however, this did not compensate for reduced photosynthetic rates at growth temperature, leading to a greater ratio of dark respiration to net carbon dioxide assimilation rate in HT trees. HT needles had 16% lower concentrations of soluble sugars than LT needles, but similar starch content. Growth at high temperatures increased the thermotolerance of black spruce. HT trees showed less PSII inhibition than LT seedlings and no increase in electrolyte leakage when briefly exposed to 40–57 °C. While trees that develop at high temperatures have enhanced tolerance for brief, extreme heat events, the reduction in root allocation indicates that seedlings will be more susceptible to episodic soil drying and less competitive for belowground resources in future climates of the boreal region.

Chill-induced decrease in capacity of RuBP carboxylation and associated H2O2 accumulation in cucumber leaves are alleviated by grafting onto figleaf gourd.

Chilling results in a significant decrease in Rubisco content and increased generation of reactive oxygen species (ROS) in cucumber (Cucumis sativus), a chilling-sensitive species. The role of roots in the regulation of the tolerance is unknown. Here, cucumber plants grafted onto figleaf gourd (Cucurbita ficifolia), a chilling-tolerant species were used to study the role of roots in the regulation of shoot functioning and the associated root-to-shoot communication. Gas exchange and chlorophyll fluorescence were measured using an infrared gas analyser combined with a pulse amplitude fluorimeter during chilling at 14 degrees C or 7 degrees C and subsequent recovery. At the same time, Rubisco content and activity and ROS generation were spectrophotometrically assayed. Abscisic acid and cytokinin concentrations in xylem sap were also determined by enzyme-linked immunosorbent assay. Grafted plants showed a significantly higher light-saturated rate of CO(2) assimilation (A(sat)) than own-rooted plants when roots were gradually cooled, but no differences were detected when shoots were cooled. Chill at 7 degrees C irreversibly reduced A(sat), and significantly decreased maximum carboxylation activity, Rubisco content and initial Rubisco activity. However, grafted plants showed weaker inhibition, together with decreased electron flux in the water-water cycle. Higher activity of antioxidant enzymes with less ROS production was found in grafted plants. In addition, ABA concentration increased by 48.4-fold whilst cytokinin concentration decreased by 91.5% in the xylem sap of own-rooted plants after exposure to a 7 degrees C chill. In comparison, ABA and cytokinin concentrations increased by 10.5-fold and 36.9%, respectively, for the grafted plants. Improved plant growth was also observed in grafted plants after the chill. These results suggest that some signals coming from chilling-resistant roots (i.e. ABA and cytokinins) protect leaf photosynthesis in shoots of chilling-sensitive plants.

Growth and photosynthetic responses to salinity of the salt-marsh shrub Atriplex portulacoides.

Atriplex (Halimione) portulacoides is a halophytic, C(3) shrub. It is virtually confined to coastal salt marshes, where it often dominates the vegetation. The aim of this study was to investigate its growth responses to salinity and the extent to which these could be explained by photosynthetic physiology. The responses of young plants to salinity in the range 0-700 mol m(-3) NaCl were investigated in a glasshouse experiment. The performance of plants was examined using classical growth analysis, measurements of gas exchange (infrared gas analysis), determination of chlorophyll fluorescence characteristics (modulated fluorimeter) and photosynthetic pigment concentrations; total ash, sodium, potassium and nitrogen concentrations, and relative water content were also determined. Plants accumulated Na(+) approximately in proportion to external salinity. Salt stimulated growth up to an external concentration of 200 mol m(-3) NaCl and some growth was maintained at higher salinities. The main determinant of growth response to salinity was unit leaf rate. This was itself reflected in rates of CO(2) assimilation, which were not affected by 200 mol m(-3) but were reduced at higher salinities. Reductions in net photosynthetic rate could be accounted for largely by lower stomatal conductance and intercellular CO(2) concentration. Apart from possible effects of osmotic shock at the beginning of the experiment, salinity did not have any adverse effect on photosystem II (PSII). Neither the quantum efficiency of PSII (Phi(PSII)) nor the chlorophyll fluorescence ratio (F(v)/F(m)) were reduced by salinity, and lower mid-day values recovered by dawn. Mid-day F(v)/F(m) was in fact depressed more at low external sodium concentration, by the end of the experiment. The growth responses of the hygro-halophyte A. portulacoides to salinity appear largely to depend on changes in its rate of photosynthetic gas exchange. Photosynthesis appears to be limited mainly through stomatal conductance and hence intercellular CO(2) concentration, rather than by effects on PSII; moderate salinity might stimulate carboxylation capacity. This is in contrast to more extreme halophytes, for which an ability to maintain leaf area can partially offset declining rates of carbon assimilation at high salinity.

Seasonal changes in the temperature response of photosynthesis in canopy leaves of Quercus crispula in a cool-temperate forest

Understanding seasonal changes in photosynthetic characteristics of canopy leaves is indispensable for modeling the carbon balance in forests. We studied seasonal changes in gas exchange characteristics that are related to the temperature dependence of photosynthesis in canopy leaves of Quercus crispula Blume, one of the most abundant species in cool-temperate forests in Japan. Photosynthetic rate and ribulose-1,5-bisphosphate (RuBP) carboxylation capacity (V(cmax)) at 20 degrees C increased from June to August and then decreased in September. The activation energy of V(cmax), a measure of the temperature dependence of V(cmax), was highest in summer, indicating that V(cmax) was most sensitive to leaf temperature at this time. The activation energy of V(cmax) was significantly correlated with growth temperature. Other parameters related to the temperature dependence of photosynthesis, such as intercellular CO(2) partial pressure and temperature dependence of RuBP regeneration capacity, showed no clear seasonal trend. It was suggested that leaf senescence affected the balance between carboxylation and regeneration of RuBP. The model simulation showed that photosynthetic rate and its optimal temperature were highest in summer.

Mesophyll conductance to CO2 in Arabidopsis thaliana

The close rosette growth form, short petioles and small leaves of Arabidopsis thaliana make measurements with commercial gas exchange cuvettes difficult. This difficulty can be overcome by growing A. thaliana plants in 'ice-cream cone-like' soil pots. This design permitted simultaneous gas exchange and chlorophyll fluorescence measurements from which the first estimates of mesophyll conductance to CO(2) (g(m)) in Arabidopsis were obtained and used to determine photosynthetic limitations during plant ageing from c. 30-45 d. Estimations of g(m) showed maximum values of 0.2 mol CO(2) m(-2) s(-1) bar(-1), lower than expected for a thin-leaved annual species. The parameterization of the response of net photosynthesis (A(N)) to chloroplast CO(2) concentrations (C(c)) yielded estimations of the maximum velocity of carboxylation (V(c,max_Cc)) which were also lower than those reported for other annual species. As A. thaliana plants aged from 30 to 45 d, there was a 40% decline of A(N) that was entirely the result of increased diffusional limitations to CO(2) transfer, with g(m) being the largest. The results suggest that in A. thaliana A(N) is limited by low g(m) and low capacity for carboxylation. Decreased g(m) is the main factor involved in early age-induced photosynthetic decline.

Interaction of drought and elevated CO2 concentration on photosynthetic down-regulation and susceptibility to photoinhibition in Japanese white birch seedlings grown with limited N availability

The interaction of drought and elevated carbon dioxide concentration ([CO(2)]) on carboxylation capacity of Rubisco (V(cmax)) and susceptibility to photoinhibition may be an important determinant of plant responses to seasonal fluctuations in precipitation in an anticipated elevated [CO(2)] environment. Japanese white birch (Betula platyphylla var. japonica) leaves that developed wholly during a period of drought showed an increase in leaf nitrogen and a decrease in leaf carbohydrates that could ameliorate photosynthetic down-regulation, defined as a decrease in V(cmax) in response to elevated [CO(2)]. Photochemical quenching (q(P)) was decreased by elevated [CO(2)] but increased by drought when compared at a given intercellular [CO(2)] (C(i)), indicating that elevated [CO(2)] could increase the risk of photoinhibition, whereas long-term drought could alleviate the risk of photoinhibition. However, only a small variation in q(P) was measured among seedlings in the various water availability x [CO(2)] treatment combinations, consistent with the small treatment differences in chronic photoinhibition among the seedlings, as indicated by the ratio of variable to maximum chlorophyll fluorescence after overnight dark-adaptation. Our results suggest that the offsetting responses-reduced V(cmax) plus increased C(i) at elevated [CO(2)] and increased V(cmax) plus reduced C(i) under drought conditions-resulted in a narrow range of susceptibility to photoinhibition at the growth [CO(2)] in Japanese white birch seedlings grown in various water availability x [CO(2)] treatment combinations.

Analysis of leakage in IRGA's leaf chambers of open gas exchange systems: quantification and its effects in photosynthesis parameterization

The measurement of the response of net photosynthesis to leaf internal CO2 (i.e. A-Ci curves) is widely used for ecophysiological studies. Most studies did not consider CO2 exchange between the chamber and the surrounding air, especially at the two extremes of A-Ci curves, where large CO2 gradients are created, leading to erroneous estimations of A and Ci. A quantitative analysis of CO2 leakage in the chamber of a portable open gas exchange system (Li-6400, LI-COR Inc., NE, USA) was performed. In an empty chamber, the measured CO2 leakage was similar to that calculated using the manufacturer's equations. However, in the presence of a photosynthetically inactive leaf, the magnitude of leakage was substantially decreased, although still significant. These results, together with the analysis of the effects of chamber size, tightness, flow rate, and gasket material, suggest that the leakage is larger at the interface between the gaskets than through the gaskets. This differential leakage rate affects the parameterization by photosynthesis models. The magnitude of these errors was assessed in tobacco plants. The results showed that leakage results in a 10% overestimation of the leaf maximum capacity for carboxylation (Vc,max) and a 40% overestimation of day respiration (Rl). Using the manufacturer's equations resulted in larger, non-realistic corrections of the true values. The photosynthetic response to CO2 concentrations at the chloroplast (i.e. A-Cc curves) was significantly less affected by leakage than A-Ci curves. Therefore, photosynthetic parameterization can be improved by: (i) correcting A and Ci values for chamber leakage estimated using a photosynthetically inactive leaf; and (ii) using A-Cc instead of A-Ci curves.

The response of photosynthesis and stomatal conductance to rising [CO2]: mechanisms and environmental interactions

This review summarizes current understanding of the mechanisms that underlie the response of photosynthesis and stomatal conductance to elevated carbon dioxide concentration ([CO2]), and examines how downstream processes and environmental constraints modulate these two fundamental responses. The results from free-air CO2 enrichment (FACE) experiments were summarized via meta-analysis to quantify the mean responses of stomatal and photosynthetic parameters to elevated [CO2]. Elevation of [CO2] in FACE experiments reduced stomatal conductance by 22%, yet, this reduction was not associated with a similar change in stomatal density. Elevated [CO2] stimulated light-saturated photosynthesis (Asat) in C3 plants grown in FACE by an average of 31%. However, the magnitude of the increase in Asat varied with functional group and environment. Functional groups with ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco)-limited photosynthesis at elevated [CO2] had greater potential for increases in Asat than those where photosynthesis became ribulose-1,5-bisphosphate (RubP)-limited at elevated [CO2]. Both nitrogen supply and sink capacity modulated the response of photosynthesis to elevated [CO2] through their impact on the acclimation of carboxylation capacity. Increased understanding of the molecular and biochemical mechanisms by which plants respond to elevated [CO2], and the feedback of environmental factors upon them, will improve our ability to predict ecosystem responses to rising [CO2] and increase our potential to adapt crops and managed ecosystems to future atmospheric [CO2].

Decrease in the capacity for RuBP carboxylation and regeneration with the progression of cold-induced photoinhibition during winter in evergreen broadleaf tree species in a temperate forest.

We measured the photosynthetic capacity (RuBP carboxylation and electron transport capacity at 25°C) and the maximum photochemical efficiency (Fv/Fm) from autumn to spring in saplings of two evergreen broadleaf tree species and examined the negative effects of photoinhibition on the photosynthetic capacity. Saplings were grown in pots under three simulated natural light environments typical of temperate forests: an open site, deciduous understorey and evergreen understorey. During winter, the photosynthetic capacity and Fv/Fm synchronously decreased in leaves in the sun, but not those in the shade. We found large differences in Fv/Fm and photosynthetic capacity, along with a positive correlation between Fv/Fm and the photosynthetic capacity among leaves in different light environments. In photoinhibited leaves that were transferred to the shade in mid-winter, photosynthetic capacity increased synchronously with the increment of Fv/Fm. The decrease in photosynthetic capacity in photoinhibited leaves and the synchronous recovery of photosynthetic capacity with photoinhibition supported the hypothesis that photoinhibition depressed the photosynthetic capacity during winter. We showed that difference in the degree of photoinhibition was responsible for the different winter photosynthetic capacity among leaves exposed to different light environments.

Influence of N form on growth photosynthesis of Phaseolus vulgaris L. plants

AbstractMany plant species are characterized by pronounced sensitivity to sole ammonium supply and exhibit growth depression and particularly reduced leaf growth rates. Stress symptoms under sole ammonium supply may be related to perturbation of photosynthetic processes, e.g., low rates of net CO2 assimilation, low quantum yield, reduced stomatal conductance, and carboxylation capacity. The results of three experiments with French bean plants supplied with an N concentration of 5 mM illustrate significantly lower dry mass and specific leaf area, reduced leaf expansion, and higher chlorophyll and N content of ammonium‐ compared to nitrate‐supplied plants. Light‐saturated rates of CO2 assimilation (Amax) per unit leaf area were higher under ammonium compared to nitrate supply while no significant effects of N form on quantum yield and Amax per unit leaf weight and chlorophyll were found. Maximal carboxylation (Vcmax) and electron‐transport (JMax) rates were significantly higher under ammonium supply only in one of three experiments. Vcmax was linearly related to total leaf N, the slope of the regression was similar with both N forms, the x‐axis intercept was significantly higher for ammonium‐ compared to nitrate‐supplied plants. The ratio Vcmax : JMax was not affected by N form. It is concluded that ammonium supply had no negative effects on the operation of photosynthetic protein‐enzyme complexes.

Photosynthetic stimulation under long‐term CO2 enrichment and fertilization is sustained across a closed Populus canopy profile (EUROFACE)

The long-term response of leaf photosynthesis to rising CO2 concentrations [CO2] depends on biochemical and morphological feedbacks. Additionally, responses to elevated [CO2] might depend on the nutrient availability and the light environment, affecting the net carbon uptake of a forest stand. After 6 yr of exposure to free-air CO2 enrichment (EUROFACE) during two rotation cycles (with fertilization during the second cycle), profiles of light, leaf characteristics and photosynthetic parameters were measured in the closed canopy of a poplar (Populus) short-rotation coppice. Net photosynthetic rate (A(growth)) was 49% higher in poplars grown in elevated [CO2], independently of the canopy position. Jmax significantly increased (15%), whereas leaf carboxylation capacity (Vcmax), leaf nitrogen (N(a)) and chlorophyll (Chl(a)) were unaffected in elevated [CO2]. Leaf mass per unit area (LMA) increased in the upper canopy. Fertilization created more leaves in the top of the crown. These results suggest that the photosynthetic stimulation by elevated [CO2] in a closed-canopy poplar coppice might be sustained in the long term. The absence of any down-regulation, given a sufficient sink capacity and nutrient availability, provides more carbon for growth and storage in this bioenergy plantation.

Modeling photosynthesis in olive leaves under drought conditions

We quantified parameters for a model of leaf-level photosynthesis for olive, and tested the model against an independent dataset. Specific temperature-dependence parameters of the model for olive leaves were measured, as well as the relationship of the model parameters with area-based leaf nitrogen (N) content. The effect of soil water deficit on leaf photosynthesis was examined by applying two irrigation treatments to 29-year-old trees growing in a plantation: drip irrigation sufficient to meet the crop water requirements (I) and dry-farming (D). In both treatments, leaves had a higher photosynthetic capacity in April than in August. In August, photosynthetic capacity was lower in D trees than in I trees. Leaf photosynthetic capacity was linearly and positively related to leaf N content on an area basis (N(a)) and to leaf mass per unit area (LMA), and the regression slope varied with irrigation treatment. The seasonal reduction in N(a) was used in the model to predict photosynthesis under drought conditions. Olive leaves showed a clear limitation of photosynthesis by triose phosphate utilization (TPU) even at 40 degrees C, and the data suggest that olive invests fewer resources in TPU than other species. The seasonal decrease in photosynthetic capacity moderated the stomatal limitation to carbon dioxide (CO(2)) fixation as soil water deficit increased. Further, it enabled leaves to operate close to the transition point between photosynthetic limitation due to RuBP carboxylation capacity and that due to RuBP regeneration capacity, and resulted in a near constant value of internal CO(2) concentration from April to August. Under well watered conditions, N-use efficiency of the olive leaves was enhanced at the expense of reduced water-use efficiency.