Interactive effects of elevated CO2 and drought on photosynthetic capacity and PSII performance in maize

Elevated atmospheric CO₂ concentration [CO₂] and the change of water distribution in arid and semiarid areas affect plant physiology and ecosystem processes. The interaction of elevated [CO₂] and drought results in the complex response such as changes in the energy flux of photosynthesis. The performance of photosystem (PS) II and the electron transport were evaluated by using OJIP induction curves of chlorophyll a fluorescence and the P _N-C _i curves in the two-factor controlled experiment with [CO₂] of 380 (AC) or 750 (EC) [μmol mol^?1] and water stress by 10% polyethylene glycol 6000. Compared to water-stressed maize (Zea mays L.) under AC, the EC treatment combined with water stress decreased the number of active reaction centers but it increased the antenna size and the energy flux (absorbed photon flux, trapping flux, and electron transport flux) of each reaction center in PSII. Thus, the electron transport rate was enhanced, despite the indistinctively changed quantum yield of the electron transport and energy dissipation. The combination of EC and the water-stress treatment resulted in the robust carboxylation rate without elevating the saturated photosynthetic rate (P _max). This study demonstrated that maize was capable of transporting more electrons into the carboxylation reaction, but this could not be used to increase P _max under EC.     

Coupled cyclic electron transport in intact chloroplasts and leaves of C3 plants: Does it exist? if so,what is its function?

Transthylakoid proton transport based on Photosystem I-dependent cyclic electron transport has been demonstrated in isolated intact spinach chloroplasts already at very low photon flux densities when the acceptor side of Photosystem I (PS I) was largely closed. It was under strict redox control. In spinach leaves, high intensity flashes given every 50 s on top of far-red, but not on top of red background light decreased the activity of Photosystem II (PS II) in the absence of appreciable linear electron transport even when excitation of PS II by the background light was extremely weak. Downregulation of PS II was a consequence of cyclic electron transport as shown by differences in the redox state of P700 in the absence and the presence of CO₂ which drained electrons from the cyclic pathway eliminating control of PS II. In the presence of CO₂, cyclic electron transport comes into play only at higher photon flux densities. At H⁺/e=3 in linear electron transport, it does not appear to contribute much ATP for carbon reduction in C3 plants. Rather, its function is to control the activity of PS II. Control is necessary to prevent excessive reduction of the electron transport chain. This helps to protect the photosynthetic apparatus of leaves against photoinactivation under light stress.     

Kinetics of urea uptake by Melosira italica (Ehr.) Kütz at different luminosity conditions

The kinetic parameters K_m and V_max for urea uptake by Melosira italica were determined at 160 μeinsteins m⁻² s⁻¹ and in the dark. The transport systems showed an affinity for the substrate and a storing capacity in the dark (K_m = 65.07 μM; V_max = 2.18 nmoles 10⁵ cells ⁻¹ h⁻¹) greater than under 160 μE m⁻² s ⁻¹ (K_m = 111.2 μM; V_max = 1.11 nmoles 105 cells⁻¹ h⁻¹). Similarly, a reduction in consumption rate of urea under increasing photon flux densities was observed. The use of an inhibitor (potassium cyanide) indicated that the uptake process requires metabolic energy. That urea transport is more important in darkness, may constitute a survival strategy in which this compound is utilized by cells mainly during heterotrophic growth.     

Differential Control of Xanthophylls and Light-Induced Stress Proteins, as Opposed to Light-Harvesting Chlorophyll a/b Proteins, during Photosynthetic Acclimation of Barley Leaves to Light Irradiance

Barley (Hordeum vulgare L.) plants were grown at different photon flux densities ranging from 100 to 1800 μmol m⁻² s⁻¹ in air and/or in atmospheres with reduced levels of O₂ and CO₂. Low O₂ and CO₂ partial pressures allowed plants to grow under high photosystem II (PSII) excitation pressure, estimated in vivo by chlorophyll fluorescence measurements, at moderate photon flux densities. The xanthophyll-cycle pigments, the early light-inducible proteins, and their mRNA accumulated with increasing PSII excitation pressure irrespective of the way high excitation pressure was obtained (high-light irradiance or decreased CO₂ and O₂ availability). These findings indicate that the reduction state of electron transport chain components could be involved in light sensing for the regulation of nuclear-encoded chloroplast gene expression. In contrast, no correlation was found between the reduction state of PSII and various indicators of the PSII light-harvesting system, such as the chlorophyll a-to-b ratio, the abundance of the major pigment-protein complex of PSII (LHCII), the mRNA level of LHCII, the light-saturation curve of O₂ evolution, and the induced chlorophyll-fluorescence rise. We conclude that the chlorophyll antenna size of PSII is not governed by the redox state of PSII in higher plants and, consequently, regulation of early light-inducible protein synthesis is different from that of LHCII.     

A simple calibrated model of Amazon rainforest productivity based on leaf biochemical properties

A simple ‘big leaf’ ecosystem gas exchange model was developed, using eddy covariance data collected at an undisturbed tropical rainforest in south-western Amazonia (Brazil). The model used mechanistic equations of canopy biochemistry combined with an empirical stomatal model describing responses to light, temperature and humidity. After calibration, the model was driven using hourly data from a weather station at the top of the tower at the measurement site, yielding an estimate of gross primary productivity (annual photosynthesis) in 1992/1993 of about 200 mol C m^?2 year ^?. Although incoming photon flux density emerged as the major control on photosynthesis in this forest, at a given PAR CO₂ assimilation rates were higher in the mornings than in the afternoons. This was attributable to stomatal closure in the afternoon in response to increasing canopy-to-air vapour pressure differences. Although most morning gas exchange was clearly limited by the rate of electron transport, afternoon gas exchange was generally observed to be very nearly co-limited by both Rubisco activity (V_max) and electron transport rate. The sensitivity of the model to changes in nitrogen allocation showed that the modelled ratio of V_max to electron transport (J_max) served nearly to maximize the annual carbon gain, and indeed, would have resulted in almost maximum annual carbon gain at the pre-industrial revolution atmospheric CO₂ concentration of 27 Pa. Modelled gross primary productivity (GPP) was somewhat lower at 27 Pa, being about 160 mol C m^?2 year^?1. The model suggests that, in the absence of any negative feedbacks on GPP, future higher concentrations of atmospheric CO₂ will continue to increase the GPP of this rainforest, up to about 230 mol C m^?2 year^?1 at 70 Pa.     

Participation of Photosynthesis in Floral Induction of the Long Day Plant Anagallis arvensis L

The saturating photon flux density (400 to 700 nanometers) for induction of flowering of the long day plant Anagallis arvensis L. was 1,900 micromoles per square meter per second (6,000 foot-candles) when an 8-hour daylength was extended to 24 hours by a single period of supplementary irradiation. The saturating photon flux density for photosynthetic CO₂ uptake during the same single supplementary light period was lower, at about 1,000 to 650 micromoles per square meter per second (3,000 to 2,000 foot-candles).

The per cent flowering and mean number of floral buds per plant were significantly reduced when the light extension treatment was given in CO₂-free air, and glucose (10 kilograms per cubic meter in water) relieved this effect. Glucose solution also significantly increased flowering of plants given supplementary light treatment in atmospheric air under a photon flux density of 80 micromoles per square meter per second. Increasing the CO₂ concentration to 1.27 grams per cubic meter of CO₂ in air during the supplementary light period did not increase flowering.

It is concluded that high photon flux densities promote flowering of Anagallis through both increased photosynthesis and the photomorphogenic action of high irradiance.

     

The Contents of Abscisic Acid and Cytokinins in Wild-Growing Species Using Different Ecological Strategies

The biomass structure and the contents of (ABA) and cytokinins (zeatin plus zeatin riboside) were studied in flowering plants of ten herbaceous species of the Middle Urals representing stress-tolerant (S) and ruderal (R) ecological strategies. The plants, which explicitly manifested S- and R-strategies, differed in ABA and cytokinin concentrations and phytohormone partitioning in particular organs. Higher ABA accumulation in the leaves and especially in the reproductive organs (here the ABA concentration exceeded by several times that of cytokinins) was characteristic of the R-plants as compared to the S-plants. In addition, the R-plants manifested higher cytokinin contents in the roots, whereas, in the S-plants, cytokinins dominated in the leaves and the reproductive organs (in the latter, the content of cytokinins exceeded that of ABA). The diverse hormonal status of the R- and S-species is discussed in relation to the patterns of plant growth and development and the photosynthate transport and partitioning characteristic of two ecological strategies. The authors conclude that plant ability to maintain the typical hormonal balance is instrumental in acquiring specific ecological strategies.     

The uptake,distribution, and translocation of86Rb in alfalfa plants susceptible and resistant to the bacterial wilt and the effect ofCorynebacterium insidiosum upon these processes

Alfalfa (Medicago sativa; L.) plants susceptible (S) and resistant (R) to the bacterial wilt were fedvia roots with a nutrient solution labelled with⁸⁶Rb⁺, at different times after inoculating them withCorynebacterium insidiosum (McCull.) H. L. Jens. The infection did not influence⁸⁶Rb⁺ uptake per plant in the course of a 14-day-period following inoculation, however it did affect its distribution differentially in the S- and the R-plants.⁸⁶Rb⁺ uptake was significantly decreased due to the infection in the S-plants on the day 49 after the inoculation (a 4-h-exposure to⁸⁶Rb⁺), with the iona also being more slowly translocated to the shoots in diseased S-plants than in diseased R-plants. Likely factors causing these effects and their relationship to alfalfa resistance to the bacterial wilt are discussed.     

Crassulacean Acid Metabolism and Photochemical Efficiency of Photosystem II in the Adaxial and Abaxial Parts of the Succulent Leaves of Kalanchoë daigremontiana Grown at Four Photon Flux Densities

Kalanchoë daigremontiana, a species possessing crassulacean acid metabolism, was grown at four photon flux densities (1300, 400, 60, and 25 micromole photons per square meter per second). In leaves which had developed at 1300 and 400 micromole photons per square meter per second, CO₂ was mainly incorporated through the lower, shaded leaf surfaces, and the chlorenchyma adjacent to the lower surfaces showed a higher degree of nocturnal acid synthesis than the chlorenchyma adjacent to the upper surfaces. In leaves acclimated to 60 and 25 micromole photons per square meter per second, the gradient in CAM activity was reversed, i.e. more CO₂ was taken up through the upper than through the lower surfaces and nocturnal acidification was higher in the tissue next to the upper surfaces. Total net carbon gain and total nocturnal acid synthesis were highest in leaves which had developed at 400 micromole photons per square meter per second. Chlorophyll content was markedly reduced in leaves which had developed at 1300 micromole photons per square meter per second, especially in the exposed adaxial parts. There was also a sustained reduction in photosystem II photochemical efficiency as indicated by measurements of the ratio of variable over maximum chlorophyll a fluorescence. These findings suggest that, at high growth photon flux densities, the reduced activity of the exposed portions of these succulent leaves is caused by (a) the adverse effects of excess light, (b) together with a genotypic component which favors CO₂ uptake and acid synthesis in the abaxial (lower) leaf parts even when light is not or only marginally excessive. This latter component is predominant at medium photon flux densities, e.g. at 400 micromole photons per square meter per second. It becomes overridden, however, under conditions of deep shade when strongly reduced light levels in the abaxial parts of the leaf chlorenchyma severely limit photosynthesis.     

Oxygen-dependent electron transport and protection from photoinhibition in leaves of tropical tree species

The roles of photorespiration and the Mehlerperoxidase pathway in sustaining electron transport and protection from photoinhibition were studied in outer canopy leaves of two species of tropical trees: the drought-deciduous Pseudobombax septenatum (Jacq.) Dug. and the evergreen Ficus insipida Willd. Ficus had a higher photosynthetic capacity than Pseudobombax and also a greater capacity for light-dependent electron transport under photorespiratory conditions (in the absence of CO₂). As a consequence, in the absence of CO₂, Ficus was able to maintain a largely oxidized electron-transport chain at higher photon flux densities than Pseudobombax. Under the same light conditions, photoinhibition (reduction in F_v/F_m) was always greater in Pseudobombax than Ficus, was increased when leaves were exposed to 2% O₂ in nitrogen compared to 21% O₂ in CO₂-free air, but was not increased by the absence of CO₂. Rates of electron transport due to the Mehler-peroxidase pathway (assessed in 2% O₂ in nitrogen) ranged between 16–40 mol · m^–2·s^–1 in both species. As the dry season approached and Pseudobombax neared leaf senescence there was a decline in the capacity for photorespiratory flux to maintain electron transport in Pseudobombax, but not in Ficus. Ratios of light-dependent electron transport to net CO₂ fixation for Pseudobombax, Ficus and two other species in the field, Luehea seemannii Tr. & Planch, and Didymopanax morototoni (Aubl.) Dec. & Planch., ranged from 6.2 (Ficus) to 16.7 (Pseudobombax). High in-situ rates of photorespiration combined with the decreased capacity of Pseudobombax for photorespiratory flux as the dry season approached indicates a decreased capacity to protect against photooxidative damage. This may contribute to the promotion of leaf senescence in Pseudobombax during the transition from wet to dry season.Abbreviations F_v/F_m ratio of variable to maximum chlorophyll a fluorescence - NPQ nonphotochemical fluorescence quenching - PFD photon flux density - Q_A primary electron acceptor of PSII This research was supported by a grant from the Mellon Foundation. We thank Monica Mejia and Juan Posada for assisting with the fluorescence measurements and Aurelio Virgo for assisting with the field CO₂-exchange measurements.     

Seasonal variations of CO2 and water vapour exchange rates over a temperate deciduous forest

Long-term and direct measurements of CO₂ and water vapour exchange are needed over forested ecosystems to determine their net annual fluxes of carbon dioxide and water. Such measurements are also needed to parameterize and test biogeochemical, ecological and hydrological assessment models. Responding to this need, eddy covariance measurements of CO₂ and water vapour were made ever a deciduous forest growing near Oak Ridge, TN, between April 1993 and April 1994. Periodic measurements were made of leaf area index, stomatal resistance, soil moisture and pre-dawn leaf water potential to characterize the gas exchange capacity of the canopy. Four factors had a disproportionate influence on the seasonal variation of CO₂ flux densities. These factors were photon flux densities (during the growing season), temperature (during the dormant season), leaf area index and the occurrence of drought The drought period occurred during the peak of the growing season and caused a significant decline in daily and hourly CO₂ flux densities, relative to observations over the stand when soil moisture was plentiful. The annual net uptake of carbon was calculated by integrating flux measurements and filling missing and spurious data with the relations obtained between measured CO₂ fluxes and environmental forcing variables. The net flux of carbon for the period between April 1993 and April 1994 was -525 g C m^?2 y^?1. This value represents a net flux of carbon from the atmosphere and into the forest. The net annual carbon exchange of this southern temperate broadleaved forest exceeded values measured over a northern temperate forest (which experiences a shorter growing season and has less leaf area) by 200 g C m^?2 y^?1 (cf. Wofsy et al 1993). The seasonal variation of canopy evaporation (latent heat flux) was controlled mostly by changes in leaf area and net radiation. A strong depression in evaporation rates was not observed during the drought Over a broadleaved forest large vapour pressure deficits promote evaporation and trees in a mixed stand are able to tap a variety of deep and shallow water sources.     

The effect of leaf-level spatial variability in photosynthetic capacity on biochemical parameter estimates using the Farquhar model: a theoretical analysis

Application of the widely used Farquhar model of photosynthesis in interpretation of gas exchange data assumes that photosynthetic properties are homogeneous throughout the leaf. Previous studies showed that heterogeneity in stomatal conductance (g_s) across a leaf could affect the shape of the measured leaf photosynthetic CO₂ uptake rate (A) versus intercellular CO₂ concentration (C_i) response curve and, in turn, estimation of the critical biochemical parameters of this model. These are the maximum rates of carboxylation (V_c,max), whole-chain electron transport (J_max), and triose-P utilization (V_TPU). The effects of spatial variation in V_c,max, J_max, and V_TPU on estimation of leaf averages of these parameters from A-C_i curves measured on a whole leaf have not been investigated. A mathematical model incorporating defined degrees of spatial variability in V_c,max and J_max was constructed. One hundred and ten theoretical leaves were simulated, each with the same average V_c,max and J_max, but different coefficients of variation of the mean (CV_VJ) and varying correlation between V_c,max and J_max (Ω). Additionally, the interaction of variation in V_c,max and J_max with heterogeneity in V_TPU, g_s, and light gradients within the leaf was also investigated. Transition from V_c,max- to J_max-limited photosynthesis in the A-C_i curve was smooth in the most heterogeneous leaves, in contrast to a distinct inflection in the absence of heterogeneity. Spatial variability had little effect on the accuracy of estimation of V_c,max and J_max from A-C_i curves when the two varied in concert (Ω = 1.0), but resulted in underestimation of both parameters when they varied independently (up to 12.5% in V_c,max and 17.7% in J_max at CV_VJ = 50%; Ω = 0.3). Heterogeneity in V_TPU also significantly affected parameter estimates, but effects of heterogeneity in g_s or light gradients were comparatively small. If V_c,max and J_max derived from such heterogeneous leaves are used in models to project leaf photosynthesis, actual A is overestimated by up to 12% at the transition between V_c,max- and J_max-limited photosynthesis. This could have implications for both crop production and Earth system models, including projections of the effects of atmospheric change.     

Kinetics and Apparent K(m) of Oxygen Cycle under Conditions of Limiting Carbon Dioxide Fixation

A mass spectrometer with a membrane inlet was used to monitor light-driven O₂ evolution, O₂ uptake, and CO₂ uptake in suspensions of algae (Scenedesmus obliquus). We observed the following. (a) The rate of O₂ uptake, which, in the presence of iodoacetamide, replaces the uptake of CO₂, showed a distinct plateau (V_max) beyond ~30% O₂ and was half-maximal at ~8% O₂. We concluded that this light-driven O₂ uptake process, which does not involve carbon compounds, is saturated at lower O₂ concentrations than are photorespiration and glycolate formation. (b) In the absence of inhibitor, O₂ evolution was relatively unaffected by the presence or absence of CO₂. During the course of CO₂ depletion, electron flow to CO₂ was replaced by an equivalent flow to O₂. (c) There was a distinct delay between the cessation of CO₂ uptake and the increase in O₂ uptake. We ascribe this delay to the transient utilization of another electron acceptor—possibly bicarbonate or another bound form of CO₂.     

Estimation of photosynthesis parameters for a modified Farquhar–von Caemmerer–Berry model using simultaneous estimation method and nonlinear mixed effects model

The aims of this paper was to modify the photosynthesis model of Farquhar, von Caemmerer and Berry (FvCB) to be able to predict light dependency of the carboxylation capacity (V_c) and to improve the prediction of temperature dependency of the maximum carboxylation capacity (V_cmax) and the maximum electron transport rate (J_max). The FvCB model was modified by adding a sub-model for Ribulose-1,5-bisphosphate carboxylase (Rubisco) activation and validating the parameters for temperature dependency of V_cmax and J_max. Values of parameters for temperature dependency of V_cmax and J_max were validated and adjusted based on data of the photosynthesis response to temperature. Parameter estimation was based on measurements under a wide range of environmental conditions, providing parameters with broad validity. The simultaneous estimation method and the nonlinear mixed effects model were applied to ensure the accuracy of the parameter estimation. The FvCB parameters, V_cmax, J_max, α (the efficiency of light energy conversion), θ (the curvature of light response of electron transport), and R_d (the non-photorespiratory CO₂ release) were estimated and validated on a dataset from two other years. Observations and predictions matched well (R² = 0.94). We conclude that incorporating a sub-model of Rubisco activation improved the FvCB model through predicting light dependency of carboxylation rate; and that estimating V_cmax, J_max, α, θ, and R_d requires data sets of both CO₂ and light response curves.     

Membrane function in differentiating skeletal muscle cells: I. Kinetic analysis of amino acid transport

Primary cultures of mononucleated myoblasts from 12-day-old chick embryos have a twofold higher rate of α-aminoisobutyric acid (AIB) transport before fusion occurs to form multinucleated myotubes. Several lines of evidence indicate that the uptake of AIB observed in both myoblasts and myotubes is primarily carrier-mediated by a membrane transport system. Increasing the temperature from 24 to 37°C results in a threefold increase in the rate of AIB uptake; both methionine and glycine inhibit AIB uptake by more than 85%; and 2,4-dinitrophenol inhibits AIB uptake by approximately 50%. In addition, the energies of activation (14.5 and 14.0 kcal/mole for myoblasts and myotubes, respectively) are characteristic of carrier-mediated transport. Resolution of AIB uptake into a saturable, carrier-mediated component and a nonsaturable, diffusion component shows that at concentrations of AIB≤1.5 mM over 97% of total AIB uptake is carriermediated in both myoblasts and myotubes. Kinetic analysis of carrier-mediated AIB uptake indicates that myoblasts and myotube membrane carriers have the same affinity for AIB (K_m values = 1.73 and 1.31 mM, respectively). However, the V_max for myoblasts is 23.7 nmole/mg/min while myotubes have a V_max of 12.6 nmole/mg/min. The twofold difference in V_max is shown to be due to a twofold difference in the quantity of membrane transport sites per milligram of protein.     

Protection against photoinhibition in the alpine plant Geum montanum

Geum montanum L. is an alpine plant usually found at altitudes between 1700 and 2600 m. Its wintergreen leaves can be subjected to very low temperatures and at the same time receive high photon flux densities at the beginning of the growth season when the snow melts. We report results of a study, performed with classical methods of biophysics, showing that leaves of G. montanum were remarkably tolerant to sunlight even at low temperatures. This tolerance results from the interplay of photorespiration and CO₂ photosassimilation. When temperatures approach 0°C, responses include stomatal opening and CO₂ uptake even under desiccation stress. This permits linear electron transport that is sufficient to avoid the excessive reduction of the electron transport chain which is known to lead to photodamage. In addition, excitation energy was shifted from photosystem (PS)II to PSI which is a very efficient energy quencher. Sensitivity of P700 in PSI to oxidation by far-red light was decreased and rates of dark reduction of photooxidized P700 were increased by actinic illumination, suggesting activation of cyclic electron transport. Consistent with this, far-red light was able to decrease the quantum yield of PSII (measured by the F _v/F _m ratio of chlorophyll fluorescence). We suggest that cyclic electron transport decreases the lumenal pH under strong light. In the presence of zeaxanthin, this increases energy dissipation at the PSII level. At low temperatures, P700 remained strongly oxidized under high irradiation while the primary electron acceptor of PSII, Q_A, was largely reduced. This shows efficient control of electron transport presumably at the level of the cytochrome b/f complex and suggests formation of a protective transthylakoid proton gradient even when linear electron transport is much reduced in the cold. Thus, several mechanisms cooperate to effectively protect the photosynthetic apparatus of G. montanum from photodamage. We see no indication of destructive “photostress” in this species during the growth season under alpine low-temperature and drought conditions. Received: 2 March 1998 / Accepted: 7 January 1999     

Daytime and nighttime carbon balance and assimilate export in soybean leaves at different photon flux densities

To evaluate daytime and nighttime carbon balance and assimilate export in soybean (Glycine max [L.] Merrill) leaves at different photon flux densities, rates of CO₂ exchange, specific leaf weights, and concentrations of sucrose and starch were measured at intervals in leaves of pod-bearing `Amsoy 71' and `Wells II' plants grown in a controlled environment room. Assimilate export was estimated from CO₂ exchange and change in specific leaf weight. Total diurnal assimilate export was similar for both cultivars. Large cultivar differences existed, however, in the partitioning of carbon into starch reserves and the relative amounts of assimilate exported during the day and the night. Total amounts of both daytime and nighttime export increased with increasing photon flux density, as did sucrose and starch concentrations, specific leaf weight, and rate of respiratory carbohydrate loss at night. Cultivar differences in nighttime rate of export were more closely related to the differences in amount of assimilate available at the end of the day than to differences in daytime rate of net CO₂ assimilation. Daytime rates of export, however, were closely related to daytime rates of net CO₂ assimilation within each cultivar. The total amount of starch depleted during the 10-hour night increased as starch concentration at the beginning of the night increased.     

Mechanisms underlying leaf photosynthetic acclimation to warming and elevated CO2 as inferred from least‐cost optimality theory

The mechanisms responsible for photosynthetic acclimation are not well understood, effectively limiting predictability under future conditions. Least‐cost optimality theory can be used to predict the acclimation of photosynthetic capacity based on the assumption that plants maximize carbon uptake while minimizing the associated costs. Here, we use this theory as a null model in combination with multiple datasets of C₃ plant photosynthetic traits to elucidate the mechanisms underlying photosynthetic acclimation to elevated temperature and carbon dioxide (CO₂). The model‐data comparison showed that leaves decrease the ratio of the maximum rate of electron transport to the maximum rate of Rubisco carboxylation (J_max/V_cmax) under higher temperatures. The comparison also indicated that resources used for Rubisco and electron transport are reduced under both elevated temperature and CO₂. Finally, our analysis suggested that plants underinvest in electron transport relative to carboxylation under elevated CO₂, limiting potential leaf‐level photosynthesis under future CO₂ concentrations. Altogether, our results show that acclimation to temperature and CO₂ is primarily related to resource conservation at the leaf level. Under future, warmer, high CO₂ conditions, plants are therefore likely to use less nutrients for leaf‐level photosynthesis, which may impact whole‐plant to ecosystem functioning.     

Response of Photosynthesis to Radiation and Intercellular CO2 Concentration in Sun and Shade Shoots of Norway Spruce

Functional differentiation of assimilation activity of sun versus shade foliage was analysed in a Norway spruce monoculture stand (age 15 years). The investigated stand density (leaf area index 8.6) and crown structure led to variation in the photosynthetically active photon flux density (PPFD) within the crowns of the sampled trees. At the saturating PPFD, the maximum rate of CO₂ uptake (P _Nmax) of exposed shoots (E-shoots) was 1.7 times that of the shaded shoots (S-shoots). The apparent quantum yield (α) of E-shoots was 0.9 times that of the S-shoots. A lower ability to use excess energy at high PPFD in photosynthesis was observed in the S-layer. The CO₂- and PPFD-saturated rate of CO₂ uptake (P _Nsat) of the E-shoots was 1.12 times and the carboxylation efficiency (τ) 1.6 times that of the S-shoots. The CO₂-saturated rate of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) carboxylation (V_Cmax) and of actual electron transport (J_amax) in the S-needles amounted to 89 and 95 % of V_Cmax and J_amax in the E-needles. Thus, in addition to the irradiation conditions and thus limitation by low J_a, the important limitation of photosynthesis in shade needles is due to carboxylation. This limitation of photosynthesis is accompanied by lower stomatal conductance. This revised version was published online in June 2006 with corrections to the Cover Date.     

Continuous ECS-indicated recording of the proton-motive charge flux in leaves

Technical features and examples of application of a special emitter–detector module for highly sensitive measurements of the electrochromic pigment absorbance shift (ECS) via dual-wavelength (550–520 nm) transmittance changes (P515) are described. This device, which has been introduced as an accessory of the standard, commercially available Dual-PAM-100 measuring system, not only allows steady-state assessment of the proton motive force (pmf) and its partitioning into ΔpH and ΔΨ components, but also continuous recording of the overall charge flux driven by photosynthetic light reactions. The new approach employs a double-modulation technique to derive a continuous signal from the light/dark modulation amplitude of the P515 signal. This new, continuously measured signal primarily reflects the rate of proton efflux via the ATP synthase, which under quasi-stationary conditions corresponds to the overall rate of proton influx driven by coupled electron transport. Simultaneous measurements of charge flux and CO₂ uptake as a function of light intensity indicated a close to linear relationship in the light-limited range. A linear relationship between these two signals was also found for different internal CO₂ concentrations, except for very low CO₂, where the rate of charge flux distinctly exceeded the rate of CO₂ uptake. Parallel oscillations in CO₂ uptake and charge flux were induced by high CO₂ and O₂. The new device may contribute to the elucidation of complex regulatory mechanisms in intact leaves.