Zeaxanthin Synthesis, Energy Dissipation, and Photoprotection of Photosystem II at Chilling Temperatures

When leaves of a mangrove, Rhizophora mangle, were exposed to an excess of light at chilling temperatures, synthesis of zeaxanthin through violaxanthin de-epoxidation as well as nonphotochemical fluorescence quenching were markedly reduced. The results suggest a protective role of energy dissipation against the adverse effects of high light and chilling temperatures: leaves of R. mangle that had been preilluminated in 2% O₂, 0% CO₂ at low photon flux density and showed a high level of zeaxanthin, and leaves that had been kept in the dark and contained no zeaxanthin, were both exposed to high light and chilling temperatures (5°C leaf temperature) in air and then held under control conditions in low light in air at 25°C. Measurements of chlorophyll a fluorescence at room temperature showed that the photochemical efficiency of PSII and the yield of maximum fluorescence of the preilluminated leaf recovered completely within 1 to 3 hours under the control conditions. In contrast, the fluorescence responses of the predarkened leaf in high light at 5°C did not recover at all. During a dark/light transient in 2% O₂, 0% CO₂ in low light at 5°C, nonphotochemical fluorescence quenching increased linearly with an increase in the zeaxanthin content in leaves of R. mangle. In soybean (Glycine max) leaves, which contained a background level of zeaxanthin in the dark, a similar treatment with excess light induced a level of nonphotochemical fluorescence quenching that was not paralleled by an increase in the zeaxanthin content.     

Photosynthetic performance and resistance to photoinhibition of Zea mays L. leaves grown at sub-optimal temperature

The performance of the photosynthetic apparatus was examined in the third leaves of Zea mays L. seedlings grown at near-optimal (25 °C) or at suboptimal (15 °C) temperature by measuring chlorophyll (ChI) a fluorescence parameters and oxygen evolution in different temperature and light conditions. In leaf tissue grown at 25 and 15 °C, the quantum yield of PSII electron transport (ψ_PSII) and the rate of O₂ evolution decreased with decreasing temperature (from 25 to 4 °C) at a photon flux density of 125 μmol m^?2 s^?1. In leaves grown at 25 °C, the decrease of ψ_PSII correlated with a decrease of photochemical ChI fluorescence quenching (q_p), whereas in leaves crown at 15 °C q_p was largely insensitive to the temperature decrease. Compared with leaves grown at 25 °C, leaves grown at 15 °C were also able to maintain a higher fraction of oxidized to reduced Q_A (greater q_p) at high photon flux densities (up to 2000 μmol m^?2 s^?1), particularly when the measurements were performed at high temperature (25 °C). With decreasing temperature and/or increasing light intensity, leaves grown at 15 °C exhibited a substantial quenching of the dark level of fluorescence F₀ (q₀) whereas this type of quenching was virtually absent in leaves grown at 25 °C. Furthermore, leaves grown at 15 °C were able to recover faster from photo inhibition of photosynthesis after a photoinhibitory treatment (1200 μmol m^?2 s^?1 at 25, 15 or 6 °C for 8 h) than leaves grown at 25 °C. The results suggest that, in spite of having a low photosynthetic capacity, Z. mays leaves grown at sub optimal temperature possess efficient mechanisms of energy dissipation which enable them to cope better with photoinhibition than leaves grown at near-optimal temperature. It is suggested that the resistance of Z. mays leaves grown at 15 °C to photoinhibition is related to the higher content of carotenoids of the xanthophyll cycle (violaxanthin + antheraxanthin + zeaxanthin) measured in these leaves than in leaves grown at 25 °C.     

Inhibition of zeaxanthin formation and of rapid changes in radiationless energy dissipation by dithiothreitol in spinach leaves and chloroplasts

Dithiothreitol, which completely inhibits the de-epoxidation of violaxanthin to zeaxanthin, was used to obtain evidence for a causal relationship between zeaxanthin and the dissipation of excess excitation energy in the photochemical apparatus in Spinicia oleracea L. In both leaves and chloroplasts, inhibition of zeaxanthin formation by dithiothreitol was accompanied by inhibition of a component of nonphotochemical fluorescence quenching. This component was characterized by a quenching of instantaneous fluorescence (F_o) and a linear relationship between the calculated rate constant for radiationless energy dissipation in the antenna chlorophyll and the zeaxanthin content. In leaves, this zeaxanthin-associated quenching, which relaxed within a few minutes upon darkening, was the major component of nonphotochemical fluorescence quenching determined in the light, i.e. it represented the `high-energy-state' quenching. In isolated chloroplasts, the zeaxanthin-associated quenching was a smaller component of total nonphotochemical quenching and there was a second, rapidly reversible high-energy-state component of fluorescence quenching which occurred in the absence of zeaxanthin and was not accompanied by F_o quenching. Leaves, but not chloroplasts, were capable of maintaining the electron acceptor, Q, of photosystem II in a low reduction state up to high degrees of excessive light and thus high degrees of nonphotochemical fluorescence quenching. When ascorbate, which serves as the reductant for violaxanthin de-epoxidation, was added to chloroplast suspensions, zeaxanthin formation at low photon flux densities was stimulated and the relationship between nonphotochemical fluorescence quenching and the reduction state in chloroplasts then became more similar to that found in leaves. We conclude that the inhibition of zeaxanthin-associated fluorescence quenching by dithiothreitol provides further evidence that there exists a close relationship between zeaxanthin and potentially photoprotective dissipation of excess excitation energy in the antenna chlorophyll.     

Light Response of CO(2) Assimilation, Dissipation of Excess Excitation Energy, and Zeaxanthin Content of Sun and Shade Leaves

Intact attached sun leaves of Helianthus annuus and shade leaves of Monstera deliciosa and Hedera helix were used to obtain light response curves of CO₂ uptake, the content of the carotenoid zeaxanthin (formed by violaxanthin de-epoxidation), as well as nonphotochemical quenching (q_NP), and the rate constant of radiationless energy dissipation (k_D). The latter two parameters were calculated from the decrease of chlorophyll a fluorescence at closed photosystem II traps in saturating pulses in the light. Among the three species, the light-saturated capacity of CO₂ uptake differed widely and light saturation of CO₂ uptake occurred at very different photon flux densities. Fluorescence quenching and zeaxanthin content exhibited features which were common to all three species: below light-saturation of CO₂ uptake nonphotochemical quenching occurred in the absence of zeaxanthin and was not accompanied by a decrease in the yield of instantaneous fluorescence. Nonphotochemical quenching, q_NP, increased up to values which ranged between 0.35 and 0.5 when based on a control value of the yield of variable fluorescence determined after 12 hours of darkness. As light saturation of CO₂ uptake was approached, q_NP showed a secondary increase and the zeaxanthin content of the leaves began to rise. This was also the point from which the yield of instantaneous fluorescence began to decrease. The increase in zeaxanthin was paralleled by an increase in the rate constant for radiationless energy dissipation k_D, which opens the possibility that zeaxanthin is related to the rapidly relaxing “high-energy-state quenching” in leaves.     

Adaptation of the Photosynthetic Apparatus in Maize Leaves as a Result of Nitrogen Limitation : Relationships between Electron Transport and Carbon Assimilation

In maize (Zea mays L., cv Contessa), nitrogen (NO₃⁻) limitation resulted in a reduction in shoot growth and photosynthetic capacity and in an increase in the leaf zeaxanthin contents. Nitrogen deficiency had only a small effect on the quantum yield of CO₂ assimilation but a large effect on the light-saturated rate of photosynthesis. Linear relationships persisted between the quantum yield of CO₂ assimilation and that of photosystem II photochemistry in all circumstances. At high irradiances, large differences in photochemical quenching and nonphotochemical quenching of Chl a fluorescence as well as the ratio of variable to maximal fluorescence (Fv/Fm) were apparent between nitrogen-deficient plants and nitrogen-replete controls, whereas at low irradiances these parameters were comparable in all plants. Light intensity-dependent increases in nonphotochemical quenching were greatest in nitrogen-deficient plants as were the decreases in Fv/Fm ratio. In nitrogen-deficient plants, photochemical quenching decreased with increasing irradiance but remained higher than in controls at high irradiances. Thermal dissipative processes were enhanced as a result of nitrogen deficiency (nonphotochemical quenching was elevated and Fv/Fm was lowered) allowing PSII to remain relatively oxidised even when carbon metabolism was limited via nitrogen limitation.     

Reduction State of Q and Nonradiative Energy Dissipation during Photosynthesis in Leaves of a Crassulacean Acid Metabolism Plant, Kalanchoë daigremontiana Hamet et Perr

Fluorescence was measured in leaves of the CAM plant Kalanchoë daigremontiana using a pulse modulation technique at room temperature. During a 12-h light period at 500 micromole photons per square meter per second (400-700 nanometers) in air containing 350 microbar CO₂, the component of fluorescence quenching related to the reduction state of Q, the primary electron transport acceptor of PSII, remained fairly constant and showed that only 20% of Q were in the reduced form. The reduction state was slightly increased at the onset and at the end of the light period. By contrast, the nonphotochemical component of fluorescence quenching which is a measure of the fraction of nonradiative deexcitation underwent marked diurnal changes. Nonradiative energy conversion was low during the phase of most active malic acid decarboxylation in the middle of the light period when uptake of atmospheric CO₂ was negligible, and when internal CO₂ partial pressures were higher than in air; this allowed for high rates of CO₂ reduction in the chloroplasts. Nonradiative energy conversion was high during the early and the late light period when atmospheric CO₂ was taken up and internal CO₂ partial pressures were below air level. Manipulation of the internal CO₂ partial pressure during the late light period by increasing or decreasing the external CO₂ partial pressure to 1710 and 105 microbar, respectively, led to changes in the magnitude of energy dependent fluorescence quenching which were consistent with the relationship between nonradiative energy dissipation and internal CO₂ partial pressure observed during the diurnal cycle. Again, the reduction state of Q was hardly affected by these treatments. Thus, changes in electron transport rate during the diurnal CAM cycle at a given photon flux density lead primarily to alterations in the rate of nonradiative energy dissipation, with the reduction state of Q being maintained at a relatively low and constant level. Conditions are described under which nonphotochemical dissipation of excitation energy reaches a maximum value and the reduction state of Q is increased.     

The mechanism(s) involved in the photoprotection of PSII at elevated CO2 in nodulated alfalfa plants

In a previous study, we found that enhanced CO₂ subjected to nodulated alfalfa plants grown at different temperatures (ambient and ambient + 4 °C) and water availability regimes could protect PSII from photodamage. The main objective of this study was to determine the mechanism(s) involved in the photoprotection of PSII at elevated CO₂ levels in this plant. Elevated CO₂ reduced carboxylation capacity-induced photosynthetic acclimation and reduced enzymatic and/or nonenzymatic antioxidant activities, suggesting that changes in electron flow did not cause any photooxidative damage (which was also confirmed by H₂O₂ and lipid peroxidation analyses). Enhanced nonphotochemical quenching and xanthophyll cycle pigments revealed that plants grown at 700 μmol mol⁻¹ CO₂ compensated for the reduction in energy sink with a larger capacity for nonphotochemical dissipation of excitation energy as heat, i.e., modulating the status of the VAZ components. Elevated CO₂ induced the de-epoxidation of violaxanthin to zeaxanthin, facilitating thermal dissipation and protecting the photosynthetic apparatus against the deleterious effect of excess excitation energy.     

The mechanism(s) involved in the photoprotection of PSII at elevated CO2 in nodulated alfalfa plants

In a previous study, we found that enhanced CO₂ subjected to nodulated alfalfa plants grown at different temperatures (ambient and ambient + 4 °C) and water availability regimes could protect PSII from photodamage. The main objective of this study was to determine the mechanism(s) involved in the photoprotection of PSII at elevated CO₂ levels in this plant. Elevated CO₂ reduced carboxylation capacity-induced photosynthetic acclimation and reduced enzymatic and/or nonenzymatic antioxidant activities, suggesting that changes in electron flow did not cause any photooxidative damage (which was also confirmed by H₂O₂ and lipid peroxidation analyses). Enhanced nonphotochemical quenching and xanthophyll cycle pigments revealed that plants grown at 700 μmol mol^?1 CO₂ compensated for the reduction in energy sink with a larger capacity for nonphotochemical dissipation of excitation energy as heat, i.e., modulating the status of the VAZ components. Elevated CO₂ induced the de-epoxidation of violaxanthin to zeaxanthin, facilitating thermal dissipation and protecting the photosynthetic apparatus against the deleterious effect of excess excitation energy.     

Zeaxanthin and the Induction and Relaxation Kinetics of the Dissipation of Excess Excitation Energy in Leaves in 2% O(2), 0% CO(2)

The relationship between the carotenoid zeaxanthin, formed by violaxanthin de-epoxidation, and nonphotochemical fluorescence quenching (q_NP) in the light was investigated in leaves of Glycine max during a transient from dark to light in 2% O₂, 0% CO₂ at 100 to 200 micromoles of photons per square meter per second. (a) Up to a q_NP (which can vary between 0 and 1) of about 0.7, the zeaxanthin content of leaves was linearly correlated with q_NP as well as with the rate constant for radiationless energy dissipation in the antenna chlorophyll (k_D). Beyond this point, at very high degrees of fluorescence quenching, only k_D was directly proportional to the zeaxanthin content. (b) The relationship between zeaxanthin and k_D was quantitatively similar for the rapidly relaxing quenching induced in 2% O₂, 0% CO₂ at 200 micromoles of photons per square meter per second and for the sustained quenching induced by long-term exposure of Nerium oleander to drought in high light (B Demmig, K Winter, A Krüger, F-C Czygan [1988] Plant Physiol 87: 17-24). These findings suggest that the same dissipation process may be induced by very different treatments and that this particular dissipation process can have widely different relaxation kinetics. (c) A rapid induction of strong nonphotochemical fluorescence quenching within about 1 minute was observed exclusively in leaves which already contained a background level of zeaxanthin.     

Photochemical heat-shock response in common bean leaves as affected by previous water deficit

The heat sensitivity of photochemical processes was evaluated in the common bean (Phaseolus vulgaris) cultivars A222, A320, and Carioca grown under well-watered conditions during the entire plant cycle (control treatment) or subjected to a temporal moderate water deficit at the preflowering stage (PWD). The responses of chlorophyll fluorescence to temperature were evaluated in leaf discs excised from control and PWD plants seven days after the complete recovery of plant shoot hydration. Heat treatment was done in the dark (5 min) at the ambient CO₂ concentration. Chlorophyll fluorescence was assessed under both dark and light conditions at 25, 35, and 45°C. In the dark, a decline of the potential quantum efficiency of photosystem II (PSII) and an increase in minimum chlorophyll fluorescence were observed in all genotypes at 45°C, but these responses were affected by PWD. In the light, the apparent electron transport rate and the effective quantum efficiency of PSII were reduced by heat stress (45°C), but no change due to PWD was demonstrated. Interestingly, only the A222 cultivar subjected to PWD showed a significant increase in nonphotochemical fluorescence quenching at 45°C. The common bean cultivars had different photochemical sensitivities to heat stress altered by a previous water deficit period. Increased thermal tolerance due to PWD was genotype-dependent and associated with an increase in potential quantum efficiency of PSII at high temperature. Under such conditions, the genotype responsive to PWD treatment enhanced its protective capacity against excessive light energy via increased nonphotochemical quenching.     

Yellowing and photosynthetic decline of barley primary leaves in response to atmospheric CO2 enrichment

The photosynthetic response of barley (Hordeum vulgare L. cv. Brant) primary leaves was studied as a function of chlorosis induced by CO₂ enrichment. Leaf yellowing, measured as changes of chlorophyll a and b, was more extensive in controlled environments at elevated (680 ± 17 µl l^?1) than at ambient (380 ± 21 µl l^?1) CO₂. Stomatal conductance of primary leaves was decreased by growth in elevated CO₂ between 11 and 18 days after sowing (DAS) when measured at both 380 and 680 µl l^?1 CO₂. Internal leaf CO₂ concentration (C_i) was also lower for elevated- compared to ambient-CO₂-grown primary leaves between 11 and 14 DAS. Results suggest that non-stomatal factors were responsible for the decreased photosynthetic rates of elevated- compared to ambient-CO₂-grown primary leaves 18 DAS. Various photochemical measurements, including quantum absorptance (α), minimal (F₀), maximal (F_m), and variable (F_v) chlorophyll fluorescence, as well as the F_v/F_m ratio, were significantly decreased 18 DAS in the elevated- compared to ambient-CO₂ treatment. Photochemical (qP) and nonphotochemical (qN) chlorophyll fluorescence quenching coefficients of 18-day-old primary leaves did not differ between CO₂ treatments. Photosynthetic electron transport rates of photosystem II were slightly lower for elevated- compared to ambient-CO₂-grown primary leaves 18 DAS. Concentrations of α-amino N (i.e. free amino acids) in barley primary leaves were increased by CO₂ enrichment 10 DAS, but subsequently, α-amino N decreased in association with photosynthetic decline. Total acid protease activity was greater in elevated- than in ambient-CO₂-grown leaves 18 DAS. The above findings suggest that photoinhibition and premature senescence were factors in the CO₂-dependent yellowing of barley primary leaves.     

The temperature acclimation of photosynthetic responses to CO2 in Zea mays and its relationship to the activities of photosynthetic enzymes and the CO2-concentrating mechanism of C4 photosynthesis

Abstract Associations between photosynthetic responses to CO₂ at rate-saturating light and photosynthetic enzyme activities were compared for leaves of maize grown under constant air temperatures of 19, 25 and 31°C. Key photosynthetic enzymes analysed were ribulose bisphosphatc (RuBP) carboxylase, phosphoenolpyruvate (PEP) carboxylase, NADP-malic enzyme and pyruvate, P_i dikinasc. Rates of CO₂-saturated photosynthesis were similar in leaves developed at 19°C and 25°C but were decreased significantly by growth at 31°C. In contrast, carboxylation efficiency differed significantly between all three temperature regimes. Carboxylation efficiency was greatest in leaves developed at 19°C and decreased with increasing temperature during growth. The changes of carboxylation efficiency were highly correlated with changes in the activity of pyruvate, P_i dikinase (r= 0.95), but not with other photosynthetic enzyme activities. The activities of these latter enzymes, including that of RuBP carboxylase, were relatively insensitive to temperature during growth. The sensitivity of quantum yield to O₂ concentration was lower in leaves grown at 19°C than in leaves grown at 31°C. These observations support the novel hypothesis that variation in the capacity for CO₂ delivery to the bundle sheath by the C₄ cycle, relative to the capacity for net assimilation by the C₂ cycle, can be a principal determinant of C₄ photosynthetic responses to CO₂.     

Influence of irradiance on photosynthesis and PSII photochemical efficiency in maize during short-term exposure at high CO<Subscript>2</Subscript> concentration

This work aimed to evaluate if gas exchange and PSII photochemical activity in maize are affected by different irradiance levels during short-term exposure to elevated CO₂. For this purpose gas exchange and chlorophyll a fluorescence were measured on maize plants grown at ambient CO₂ concentration (control CO₂) and exposed for 4 h to short-term treatments at 800 μmol(CO₂) mol⁻¹ (high CO₂) at a photosynthetic photon flux density (PPFD) of either 1,000 μmol m⁻² s⁻¹ (control light) or 1,900 μmol m⁻² s⁻¹ (high light). At control light, high-CO₂ leaves showed a significant decrease of net photosynthetic rate (P _N) and a rise in the ratio of intercellular to ambient CO₂ concentration (C _i/C _a) and water-use efficiency (WUE) compared to control CO₂ leaves. No difference between CO₂ concentrations for PSII effective photochemistry (Φ_PSII), photochemical quenching (q_p) and nonphotochemical quenching (NPQ) was detected. Under high light, high-CO₂ leaves did not differ in P _N, C _i/C _a, Φ_PSII and NPQ, but showed an increase of WUE. These results suggest that at control light photosynthetic apparatus is negatively affected by high CO₂ concentration in terms of carbon gain by limitations in photosynthetic dark reaction rather than in photochemistry. At high light, the elevated CO₂ concentration did not promote an increase of photosynthesis and photochemistry but only an improvement of water balance due to increased WUE.     

Relative contributions of zeaxanthin-related and zeaxanthin-unrelated types of ;high-energy-state' quenching of chlorophyll fluorescence in spinach leaves exposed to various environmental conditions

We have identified two rapidly relaxing components of non-photochemical fluorescence quenching which suggests that dissipative processes occur in two different sites in the photochemical system of leaves. Under a variety of treatment conditions involving different leaf temperatures, photon flux densities (PFD), exposure times, and in the presence of 5% CO₂ or 2% O₂, no CO₂, the components of nonphotochemical fluorescence quenching were characterized with respect to their sensitivity to dithiothreitol (DTT, which completely inhibits zeaxanthin formation), the effect on instantaneous fluorescence, and the rapidity of relaxation upon darkening. Under most circumstances the DTT-sensitive component (associated with a quenching of instantaneous fluorescence and correlated with zeaxanthin) represented the majority of the rapidly relaxing portion of fluorescence quenching. A DTT-insensitive (zeaxanthin-independent) component, which also relaxed rapidly upon darkening but was not associated with a quenching of instantaneous fluorescence, became proportionally greater in an atmosphere of 2% O₂ and no CO₂, at elevated leaf temperatures, and to some degree during the induction of photosynthesis (1 minute after the onset of illumination). A third component which was also DTT-insensitive and was sustained upon darkening, was largely suppressed in 2% O₂, O% CO₂. We conclude that, under conditions favorable for photosynthesis, energy dissipation occurred mainly in the chlorophyll antennae whereas, under conditions less favorable for photosynthesis, a second dissipation process, probably in or around the reaction center of photosystem II, also developed. Furthermore, evidence is presented that the zeaxanthin-associated dissipation process prevents sustained inactivation of photochemistry by excessive light.     

Chlorophyll Fluorescence Quenching During Photosynthetic Induction in Leaves of Abutilon striatum Dicks. Infected with Abutilon Mosaic Virus,Observed with a Field-Portable Imaging System

A portable Chi fluorescence imaging system was used to characterise nonuniform Chi fluorescence quenching in Abutilon striatum leaves infected with phloem-localised abutilon mosaic virus. The instrument was used to observe fluorescence emission at intervals during induction transients, and to map nonphotochemical quenching during saturating pulses applied in the course of these transients. Two symptom types were distinguished: yellow vein-associated motifs that showed lower maximum Chi fluorescence than nearby green tissues, but virtually zero nonphotochemical quenching, and vein-defined mosaics (pale green) that initially showed normal maximum Chi fluorescence but strongly impaired nonphotochemical quenching. Mature vein-defined mosaics (yellow to white areas) resembled vein-associated symptoms with zero nonphotochemical quenching. Islands of apparently healthy green tissue enclosed by mosaic symptoms showed slower nonphotochemical quenching than controls. Possible effects of localised carbohydrate accumulation, thought to follow from infection by the phloem-limited virus, on photosynthetic processes as well as the synthesis and stability of chloroplast protein complexes, are discussed in the context of symptom development.     

The role of the intra- and extracellular protons in the photosynthetic response induced by the variation potential in pea seedlings

Local burning induces generation and propagation of variation potential (VP) in higher plants. VP induces transient inactivation of photosynthesis, which is possibly connected with proton signal in plant cell. Analysis of the role of changes in intracellular and extracellular pH in the VP-induced photosynthetic response in pea seedlings was the aim of this work. It was shown that local burning induced VP propagation, which was accompanied with a decrease of intracellular pH and increase of extracellular pH. VP induced photosynthesis inactivation that included an increase in the nonphotochemical fluorescence quenching and a decrease in the CO₂ assimilation rate. Analysis of photosynthetic responses under control and low external CO₂ concentration and changes in pH showed that there were two components in the responses. The first component appeared as a fast decrease of the CO₂ assimilation and increase of nonphotochemical quenching. It depended on the activity of the dark stage of photosynthesis and was connected with apoplast alkalization. The second component was presented as a slow increase of nonphotochemical quenching. It weakly depended on a dark stage and was connected with a decrease of intracellular pH.     

Growth and photosynthesis of two eucalypt species during high temperature stress under ambient and elevated [CO2]

Two species of eucalypt (Eucalyptus macrorhyncha and E. rossii) were grown under conditions of high temperatures (45 °C, maximum) and high light (1500 μmol m^?2 s^?1, maximum) at either ambient (350 μL L^?1) or elevated (700 μL L^?1) CO₂ concentrations for 8 weeks. The growth enhancement, in terms of total dry weight, was 41% and 103% for E. macrorhyncha and E. rossii, respectively, when grown in elevated [CO₂]. A reduction in specific leaf area and increased concentrations of non-structural carbohydrates were observed for leaves grown in elevated [CO₂]. Plants grown in elevated [CO₂] had an overall increase in photosynthetic CO₂ assimilation rate of 27%; however, when measured at the same CO₂ concentration a down-regulation of photosynthesis was evident especially for E. macrorhyncha. During the midday period when temperatures and irradiances were maximal, photosynthetic efficiency as measured by chlorophyll fluorescence (F_v/F_m) was lower in E. macrorhyncha than in E. rossii. Furthermore, F_v/F_m was lower in leaves of E. macrorhyncha grown under elevated than under ambient [CO₂]. These reductions in F_v/F_m were accompanied by increases in both photochemical (qP) and nonphotochemical quenching (qN and NPQ), and by increases in the concentrations of xanthophyll cycle pigments with an increased proportion of the total xanthophyll cycle pool comprising of antheraxanthin and zeaxanthin. Thus, increased atmospheric [CO₂] may enhance photoinhibition when environmental stresses such as high temperatures limit the capacity of a plant to respond with growth to elevated [CO₂].     

Estimation of the whole-plant CO2 compensation point of tobacco (Nicotiana tabacum L.)

The whole-plant CO₂ compensation point (Γ_plant) is the minimum atmospheric CO₂ level required for sustained growth. The minimum CO₂ requirement for growth is critical to understanding biosphere feedbacks on the carbon cycle during low CO₂ episodes; however, actual values of Γ_plant remain difficult to calculate. Here, we have estimated Γ_plant in tobacco by measuring the relative leaf expansion rate at several low levels of atmospheric CO₂, and then extrapolating the leaf growth vs. CO₂ response to estimate CO₂ levels where no growth occurs. Plants were grown under three temperature treatments, 19/15, 25/20 and 30/25°C day/night, and at CO₂ levels of 100, 150, 190 and 270 μmol CO₂ mol⁻¹ air. Biomass declined with growth CO₂ such that Γ_plant was estimated to be approximately 65 μmol mol⁻¹ for plants grown at 19/15 and 30/25°C. In the first 19 days after germination, plants grown at 100 μmol mol⁻¹ had low growth rates, such that most remained as tiny seedlings (canopy size <1 cm²). Most seedlings grown at 150 μmol mol⁻¹ and 30/25°C also failed to grow beyond the small seedling size by day 19. Plants in all other treatments grew beyond the small seedling size within 3 weeks of planting. Given sufficient time (16 weeks after planting) plants at 100 μmol mol⁻¹ eventually reached a robust size and produced an abundance of viable seed. Photosynthetic acclimation did not increase Rubisco content at low CO₂. Instead, Rubisco levels were unchanged except at the 100 and 150 μmol mol⁻¹ where they declined. Chlorophyll content and leaf weight per area declined in the same proportion as Rubisco, indicating that leaves became less expensive to produce. From these results, we conclude that the effects of very low CO₂ are most severe during seedling establishment, in large part because CO₂ deficiency slows the emergence and expansion of new leaves. Once sufficient leaf area is produced, plants enter the exponential growth phase and acquire sufficient carbon to complete their life cycle, even under warm conditions (30/25°C) and CO₂ levels as low as 100 μmol mol⁻¹.     

Photosynthetic downregulation in leaves of the Japanese white birch grown under elevated CO2 concentration does not change their temperature‐dependent susceptibility to photoinhibition

To determine the effects of elevated CO₂ concentration ([CO₂]) on the temperature‐dependent photosynthetic properties, we measured gas exchange and chlorophyll fluorescence at various leaf temperatures (15, 20, 25, 30, 35 and 40°C) in 1‐year‐old seedlings of the Japanese white birch (Betula platyphylla var. japonica), grown in a phytotron under natural daylight at two [CO₂] levels (ambient: 400 µmol mol^?1 and elevated: 800 µmol mol^?1) and limited N availability (90 mg N plant^?1). Plants grown under elevated [CO₂] exhibited photosynthetic downregulation, indicated by a decrease in the carboxylation capacity of Rubisco. At temperatures above 30°C, the net photosynthetic rates of elevated‐CO₂‐grown plants exceeded those grown under ambient [CO₂] when compared at their growth [CO₂]. Electron transport rates were significantly lower in elevated‐CO₂‐grown plants than ambient‐CO₂‐grown ones at temperatures below 25°C. However, no significant difference was observed in the fraction of excess light energy [(1 ? q_P)× F_v′/F_m′] between CO₂ treatments across the temperature range. The quantum yield of regulated non‐photochemical energy loss was significantly higher in elevated‐CO₂‐grown plants than ambient, when compared at their respective growth [CO₂] below 25°C. These results suggest that elevated‐CO₂‐induced downregulation might not exacerbate the temperature‐dependent susceptibility to photoinhibition, because reduced energy consumption by electron transport was compensated for by increased thermal energy dissipation at low temperatures.     

The effects of development at sub-optimal growth temperatures on photosynthetic capacity and susceptibility to chilling-dependent photoinhibition in Zea mays

When plants of Zea mays L. cv. LG11 that have been grown at optimal temperatures are transferred to chilling temperatures (0–12°C) photoinhibition of photosynthetic CO₂ assimilation can occur. This study examines how growth at sub-optimal temperatures alters both photosynthetic capacity and resistance to chilling-dependent photoinhibition. Plants of Z. mays cv. LG11 were grown in controlled environments at 14, 17, 20 and 25°C. As a measure of the capacity for photosynthesis under light limiting conditions, the maximum quantum yields of CO₂ assimilation (φ^a.c) and O₂ evolution (φ^a.o) were determined for the laminae of the second leaves at photon fluxes of 50–150 μmol m^-2s^-1. To determine photosynthetic capacity at photon fluxes approaching light saturation, rates of CO₂ uptake (A₁₅₀₀) and O₂ evolution (A₁₅₀₀) were determined in a photon flux of 1500 μmol m^-2s^-1. In leaves developed at 14°C, φ and φ were 26 and 43%, respectively, of the values for leaves grown at 25°C. Leaves grown at 17°C showed intermediate reductions in φ and φ, whilst leaves developed at 20°C showed no significant differences from those grown at 25°C. Similar patterns of decrease were observed for A₁₅₀₀ and A_1500.0 with decreasing growth temperature. Leaves developed at 25°C showed higher rates of CO₂ assimilation at all light levels and measurement temperatures in comparison to leaves developed at 17 and 14°C. A greater reduction in A₁₅₀₀ relative to A_1500.0 with decreasing growth temperature was attributed to increased stomatal limitation. Exposure of leaves to 800–1000 μmol m^-2 s^-1 when plant temperature was depressed to ca 6.5°C produced a photoinhibition of photosynthetic CO₂ assimilation in all leaves. However, in leaves developed at 17°C the decrease in A₁₅₀₀ following this chilling treatment was only 25% compared to 90% in leaves developed at 25°C. Recovery following chilling was completed earlier in leaves developed at 17°C. The results suggest that growth at sub-optimal temperatures induces increased tolerance to exposure to high light at chilling temperatures. This is offset by the large loss in photosynthetic capacity imposed by leaf development at sub-optimal temperatures.