Relationships between the Efficiencies of Photosystems I and II and Stromal Redox State in CO(2)-Free Air : Evidence for Cyclic Electron Flow in Vivo

The responses of the efficiencies of photosystems I and II, stromal redox state (as indicated by NADP-malate dehydrogenase activation state), and activation of the Benson-Calvin cycle enzymes ribulose 1,5-bisphosphate carboxylase and fructose 1,6-bisphosphatase to varying irradiance were measured in pea (Pisum sativum L.) leaves operating close to the CO₂ compensation point. A comparison of the relationships among these parameters obtained from leaves in air was made with those obtained when the leaves were maintained in air from which the CO₂ had been removed. P700 was more oxidized at any measured irradiance in CO₂-free air than in air. The relationship between the quantum efficiencies of the photosystems in CO₂-free air was distinctly curvilinear in contrast to the predominantly linear relationship obtained with leaves in air. This nonlinearity may be consistent with the operation of cyclic electron flow around photosystem I because the quantum efficiency of photosystem II was much more restricted than the quantum efficiency of photosystem I. In CO₂-free air, measured NADP-malate dehydrogenase activities varied considerably at low irradiances. However, at high irradiance the activity of the enzyme was low, implying that the stroma was oxidized. In contrast, fructose-1,6-bisphosphatase activities tended to increase with increasing electron flux through the photosystems. Ribulose-1,5-bisphosphate carboxylase activity remained relatively constant with respect to irradiance in CO₂-free air, with an activation state 50% of maximum. We conclude that, at the CO₂ compensation point and high irradiance, low redox states are favored and that cyclic electron flow may be substantial. These two features may be the requirements necessary to trigger and maintain the dissipative processes in the thylakoid membrane.     

Control of the Quantum Efficiencies of Photosystems I and II, Electron Flow, and Enzyme Activation following Dark-to-Light Transitions in Pea Leaves: Relationship between NADP/NADPH Ratios and NADP-Malate Dehydrogenase Activation State

The quantum efficiencies of photosystems I and II (PSI and PSII), [NADP]/[NADPH] ratios, and the activities of chloroplastic fructose-1,6-bisphosphatase and NADP-malate dehydrogenase were measured in intact pea (Pisum sativum L.) leaves in air following the transition from darkness to 750 microeinsteins per square meter per second irradiance. PSII efficiency declined from a low value to a minimum within the first 10 to 15 seconds of irradiance, after which it increased progressively to the steady-state value. The resistance of electron flow between the photosystems was high at this time, but it was not the principal factor limiting electron flow. Oxidation of P700 was restricted by acceptor side processes for approximately the first 60 seconds of illumination. Once the acceptor side limitation was relieved, the oxidation state of P700 was used to estimate the quantum efficiency of electron transport by PSI. This was observed to increase progressively with time. The quantum efficiencies of both photosystems increased in parallel, consistent with a predominant role for noncyclic electron transport. Fructose-1,6-bisphosphatase activity increased in an approximately sigmoidal fashion with time of irradiance, paralleling the changes in the quantum efficiencies of the photosystems. In contrast, the activation of NADP-malate dehydrogenase did not show a lag period but increased with time, reaching a maximum value at about 50 seconds of illumination, after which it declined. The NADP pool was not extensively reduced during the first 10 seconds of illumination, but became so subsequently. It remained in the reduced state until about 60 seconds of illumination and then became relatively oxidized. The empirical relationship between NADP-malate dehydrogenase activity and the reduction state of the NADP pool supports the suggestion that NADP-malate dehydrogenase activity is a useful estimate of the reduction state of the stroma.     

Acclimation of photosynthesis, H2O2 content and antioxidants in maize (Zea mays) grown at sub-optimal temperatures

Maize plants were grown at 14, 18 and 20 °C until the fourth leaf had emerged. Leaves from plants grown at 14 and 18 °C had less chlorophyll than those grown at 20 °C. Maximal extractable ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) activity was decreased at 14 °C compared with 20 °C, but the activation state was highest at 14 °C. Growth at 14 °C increased the abundance (but not the number) of Rubisco breakdown products. Phosphoenolpyruvate carboxylase (PEPC) activity was decreased at 14 °C compared with 20 °C but no chilling-dependent effects on the abundance of the PEPC protein were observed. Maximal extractable NADP-malate dehydrogenase activity increased at 14 °C compared with 20 °C whereas the glutathione pool was similar in leaves from plants grown at both temperatures. Foliar ascorbate and hydrogen peroxide were increased at 14 °C compared with 20 °C. The foliar hydrogen peroxide content was independent of irradiance at both growth temperatures. Plants grown at 14 °C had decreased rates of CO₂ fixation together with decreased quantum efficiencies of photosystem (PS) II in the light, although there was no photo-inhibition. Growth at 14 °C decreased the abundance of the D1 protein of PSII and the PSI psaB gene product but the psaA gene product was largely unaffected by growth at low temperatures. The relationships between the photosystems and the co-ordinate regulation of electron transport and CO₂ assimilation were maintained in plants grown at 14 °C.     

Photosynthetic electron transport and specific photoprotective responses in wheat leaves under drought stress

The photosynthetic responses of wheat (Triticum aestivum L.) leaves to different levels of drought stress were analyzed in potted plants cultivated in growth chamber under moderate light. Low-to-medium drought stress was induced by limiting irrigation, maintaining 20 % of soil water holding capacity for 14 days followed by 3 days without water supply to induce severe stress. Measurements of CO₂ exchange and photosystem II (PSII) yield (by chlorophyll fluorescence) were followed by simultaneous measurements of yield of PSI (by P700 absorbance changes) and that of PSII. Drought stress gradually decreased PSII electron transport, but the capacity for nonphotochemical quenching increased more slowly until there was a large decrease in leaf relative water content (where the photosynthetic rate had decreased by half or more). We identified a substantial part of PSII electron transport, which was not used by carbon assimilation or by photorespiration, which clearly indicates activities of alternative electron sinks. Decreasing the fraction of light absorbed by PSII and increasing the fraction absorbed by PSI with increasing drought stress (rather than assuming equal absorption by the two photosystems) support a proposed function of PSI cyclic electron flow to generate a proton-motive force to activate nonphotochemical dissipation of energy, and it is consistent with the observed accumulation of oxidized P700 which causes a decrease in PSI electron acceptors. Our results support the roles of alternative electron sinks (either from PSII or PSI) and cyclic electron flow in photoprotection of PSII and PSI in drought stress conditions. In future studies on plant stress, analyses of the partitioning of absorbed energy between photosystems are needed for interpreting flux through linear electron flow, PSI cyclic electron flow, along with alternative electron sinks.     

Stoichiometry of Photosystem I, Photosystem II, and Phycobilisomes in the Red Alga Porphyridium cruentum as a Function of Growth Irradiance

Cells of the red alga Porphyridium cruentum (ATCC 50161) exposed to increasing growth irradiance exhibited up to a three-fold reduction in photosystems I and II (PSI and PSII) and phycobilisomes but little change in the relative numbers of these components. Batch cultures of P. cruentum were grown under four photon flux densities of continuous white light; 6 (low light, LL), 35 (medium light, ML), 180 (high light, HL), and 280 (very high light, VHL) microeinsteins per square meter per second and sampled in the exponential phase of growth. Ratios of PSII to PSI ranged between 0.43 and 0.54. About three PSII centers per phycobilisome were found, regardless of growth irradiance. The phycoerythrin content of phycobilisomes decreased by about 25% for HL and VHL compared to LL and ML cultures. The unit sizes of PSI (chlorophyll/P₇₀₀) and PSII (chlorophyll/Q_A) decreased by about 20% with increase in photon flux density from 6 to 280 microeinsteins per square meter per second. A threefold reduction in cell content of chlorophyll at the higher photon flux densities was accompanied by a twofold reduction in β-carotene, and a drastic reduction in thylakoid membrane area. Cell content of zeaxanthin, the major carotenoid in P. cruentum, did not vary with growth irradiance, suggesting a role other than light-harvesting. HL cultures had a growth rate twice that of ML, eight times that of LL, and slightly greater than that of VHL cultures. Cell volume increased threefold from LL to VHL, but volume of the single chloroplast did not change. From this study it is evident that a relatively fixed stoichiometry of PSI, PSII, and phycobilisomes is maintained in the photosynthetic apparatus of this red alga over a wide range of growth irradiance.     

Endogenous Control of Photosynthetic Activity during Progressive Drought: Influence of Final Products of Photosynthesis

Photosynthetic activities and the redox states of photosystem I (PSI) and photosystem II (PSII) in intact leaves of cucumber plants (Cucumis sativus L.), as well as the sucrose and starch contents were examined under conditions of ongoing soil water deficit imposed by the cessation of watering. As the soil drought progressed, the maximum rate of photosynthetic CO₂ fixation was shown to decrease. These changes in the maximum photosynthetic rate occurred synchronously with changes in the maximum quantum yield of photosynthesis. Under soil water deficit, the reduced form of PSII primary acceptor Q _A was accumulated only at photon flux densities of about 100 mol/(m² s). At such photon flux densities, the changes in nonphotochemical quenching (qN) induced by soil water deficit were opposite to changes in photochemical quenching parameter (1 – qP). Irrespective of the duration of soil drought, the relationship between steady-state concentrations of photochemically inactive reaction centers of PSI and PSII (the fractions of P700 and Q _A in the oxidized and reduced state, respectively) was almost linear, which provides evidence for the concerted operation of both photosystems. The conditions of soil water deficit promoted sucrose accumulation in the source leaf, which was paralleled by a substantial decrease in the amount of starch in the same leaf. The highest content of sucrose in the leaf after a 7-day drought was correlated with the largest decrease in photosynthetic activity. It is concluded that the progressive drought triggers an endogenous mechanism that regulates photosynthesis through feedback relations, namely, the inhibition of photosynthesis by its end products.     

Activities of Photosystems I and II in Chlamydomonas segnis Adapted and Adapting to Air and Air-enriched with Carbon Dioxide*

Activities of photosystems I and II were compared at a saturating irradiance in air- and 5% CO₂-adapted and adapting Chlamydomonas segnis at the active phase of photosynthesis during the cell cycle. PSII activity was 200% greater in air- than in 5% CO₂-adapted cells, while PSI activity was similar in both types of cells and matched the level of PSII activity in air-adapted cells. As a result, air- and 5% CO₂-adapted cells were characterized by low and high PSI/PSII ratios, respectively. In air-adapted cells, the greater PSII activity (rate of O₂ evolved) exceeded that of photosynthetic (Ps) O₂ evolution, resulting in a Ps/PSII ratio below unity. This was associated with higher levels of catalase activity, lower l -ascorbate content, and higher dehydro-l -ascorbate content than in 5% CO₂-adapted cells. During adaptation to air or 5% CO₂ for 6 h in light, PSI rather than PSII was sensitive to changes in the concentration of CO₂, and the adapting cells acquired the characteristics of air- and 5% CO₂-adapted cells as indicated by PSI/PSII, Ps/PSII, catalase activity, l -ascorbate and dehydro-l -ascorbate contents. The results are discussed in the light of changes in the molecular organization of the thylakoid membranes and enhanced non-cyclic electron transport coupled with O₂-uptake (Mehler reaction) for the generation of the ATP required for CO₂/HCO^?₃-transport in air-adapted and adapting cells.     

Changes in the photosynthetic characteristics and photosystem stoichiometries in wild-type and Chl <Emphasis Type="Italic">b</Emphasis>-deficient mutant rice seedlings under various irradiances

By using a wild-type rice (Oryza sativa L. cv. Norin No. 8) and the chlorophyll (Chl) b-deficient mutant derived from Norin No. 8 (chlorina 11), the present study monitored the oxygen evolution, contents of Chl a and b, β-carotene, and lutein in leaf and the contents of cytochrome f, and the reaction centres of photosystem I (PSI) and photosystem II (PSII) in thylakoids. The oxygen evolution, maximal quantum yield of PSII (F_v/F_m) and Chl concentration remained constant in both Norin No. 8 and chlorina 11 under 5 and 2% of full sunlight for six days. On the other hand, on the thylakoid level, the PSII reaction centre of chlorina 11 was more stable even under high irradiance, while approximately 40% decrease in levels of the PSII reaction centre occurred under 2% of full sunlight for six days. However, under such conditions, by regulating the stoichiometry of active PSII and PSI centres, the light absorption balance in both rice types was adjusted between the two photosystems. The present study attempted to examine whether the light absorption balance between PSII and PSI is altered to effectively conduct photosynthesis in the wild-type and Chl b-deficient mutant rice seedlings.     

Seasonal Effects on Photosynthetic Electron Transport and Fluorescence Properties in Isolated Chloroplasts of Pinus silvestris

Chloroplasts were isolated from primary needles of 1-year-old seedlings and from secondary needles of a 20-year-old pine tree in a natural stand. In autumn the electron transport capacities of PSII, PSI and PS (II + I) decreased and the electron transport between PSII and PSI became inhibited in October in the 20-year-old tree. This inhibition lasted until May the following year. The partial reactions of PSI and PSII still showed low but fairly constant rates during the whole winter seedlings. Seasonal changes in the electron transport properties of 1-year-old showed the same general trends as observed in the 20-year-old tree, but the changes were less pronounced. However, in snow-covered seedlings the PSI-mediated electron transport and the electron transport from H₂O to NADP increased during the late winter when the seedlings were still covered by snow. The total chlorophyll content of the needles decreased in autumn and winter. Low temperature fluorescence ratios of F692/F680 and F726/F680 indicated more severe destruction of the chlorophyll a antennae closely associated with the two photosystems than of the light harvesting chlorophyll a/b complex. In this case, too, the changes were more pronounced in the 20-year-old tree than in the 1-year-old seedlings. The chlorophyll/P700 ratios indicated a more marked reduction in the reaction centre molecules during autumn than in the antennae chlorophyll molecules. The changes in electron transport and low temperature fluorescence properties which occurred during autumn and winter were mainly reversed during spring.     

Differential Effects of Nitrogen Limitation on Photosynthetic Efficiency of Photosystems I and II in Microalgae

The effects of nitrogen starvation on photosynthetic efficiency were examined in three unicellular algae by measuring changes in the quantum yield of fluorescence with a pump-and-probe method and thermal efficiency (i.e. the percentage of trapped energy stored photochemically) with a pulsed photoacoustic method together with the inhibitor 3-(3,4-dichlorophenyl)-1,1-dimethylurea to distinguish photosystems I and II (PSI and PSII). Measured at 620 nm, maximum thermal efficiency for both photosystems was 32% for the diatom Thalassiosira weissflogii (PSII:PSI ratio of 2:1), 39% for the green alga Dunaliella tertiolecta (PSII:PSI ratio of 1:1), and 29% for the cyanobacterium Synechococcus sp. PCC 7002 (PSII:PSI ratio of 1:2). Nitrogen starvation decreased total thermal efficiency by 56% for T. weissflogii and by 26% for D. tertiolecta but caused no change in Synechococcus. Decreases in the number of active PSII reaction centers (inferred from changes in variable fluorescence) were larger: 86% (T. weissflogii), 65% (D. tertiolecta), and 65% (Synechococcus). The selective inactivation of PSII under nitrogen starvation was confirmed by independent measurements of active PSII using oxygen flash yields and active PSI using P700 reduction. Relatively high thermal efficiencies were measured in all three species in the presence of the PSII inhibitor 3-(3,4-dichlorophenyl)-1,1-dimethylurea, suggesting the potential for significant cyclic electron flow around PSI. Fluorescence or photoacoustic data agreed well; in T. weissflogii, the functional cross-sectional area of PSII at 620 nm was estimated to be the same using both methods (approximately 1.8 x 102 A2). The effects of nitrogen starvation occur mainly in PSII and are well represented by variable fluorescence measurements.     

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.     

Consequences of LHC II deficiency for photosynthetic regulation in chlorina mutants of barley

The light-harvesting chlorophyll a/b proteins associated with PS II (LHC II) are often considered to have a regulatory role in photosynthesis. The photosynthetic responses of four chlorina mutants of barley, which are deficient in LHC II to varying degrees, are examined to evaluate whether LHC II plays a regulatory role in photosynthesis. The efficiencies of light use for PS I and PS II photochemistry and for CO₂ assimilation in leaves of the mutants were monitored simultaneously over a wide range of photon flux densities of white light in the presence and absence of supplementary red light. It is demonstrated that the depletions of LHC II in these mutants results in a severe imbalance in the relative rates of excitation of PS I and PS II in favour of PS I, which cannot be alleviated by preferential excitation of PS II. Analyses of xanthophyll cycle pigments and fluorescence quenching in leaves of the mutants indicated that the major LHC II components are not required to facilitate the light-induced quenching associated with zeaxanthin formation. It is concluded that LHC II is important to balance the distribution of excitation energy between PS I and PS II populations over a wide range of photon flux densities. It appears that LHC II may also be important in determining the quantum efficiency of PS II photochemistry by reducing the rate of quenching of excitation energy in the PS II primary antennae.Abbreviations Fm, Fv maximal and variable fluorescence yields in a light adapted state - LHC II light harvesting chlorophyll a/b protein complex associated with PS II - q_p photochemical quenching - A₈₂₀ light-induced absorbance change at 820 nm - ø_PSI, ø_PSII relative quantum efficiencies of PS I and PS II photochemistry - øCO₂ quantum yield of CO₂ assimilation     

Response of the Photosynthetic Apparatus in Dunaliella salina (Green Algae) to Irradiance Stress

The response of the photosynthetic apparatus in the green alga Dunaliella salina, to irradiance stress was investigated. Cells were grown under physiological conditions at 500 millimoles per square meter per second (control) and under irradiance-stress conditions at 1700 millimoles per square meter per second incident intensity (high light, HL). In control cells, the light-harvesting antenna of photosystem I (PSI) contained 210 chlorophyll a/b molecules. It was reduced to 105 chlorophyll a/b in HL-grown cells. In control cells, the dominant form of photosystem II (PSII) was PSII_α(about 63% of the total PSII) containing >250 chlorophyll a/b molecules. The smaller antenna size PSII_β centers (about 37% of PSII) contained 135 ± 10 chlorophyll a/b molecules. In sharp contrast, the dominant form of PSII in HL-grown cells accounted for about 95% of all PSII centers and had an antenna size of only about 60 chlorophyll a molecules. This newly identified PSII unit is termed PSII_γ. The HL-grown cells showed a substantially elevated PSII/PSI stoichiometry ratio in their thylakoid membranes (PSII/PSI = 3.0/1.0) compared to that of control cells (PSII/PSI = 1.4/1.0). The steady state irradiance stress created a chronic photoinhibition condition in which D. salina thylakoids accumulate an excess of photochemically inactive PSII units. These PSII units contain both the reaction center proteins and the core chlorophyll-protein antenna complex but cannot perform a photochemical charge separation. The results are discussed in terms of regulatory mechanism(s) in the plant cell whose function is to alleviate the adverse effect of irradiance stress.     

Down-regulation of linear and activation of cyclic electron transport during drought

The effects of short-term drought on the regulation of electron transport through photosystems I and II (PSI and PSII) have been studied in Hordeum vulgare L. cv. Chariot. Fluorescence measurements demonstrated that electron flow through PSII decreased in response to both drought and CO₂ limitation. This was due to regulation, as opposed to photoinhibition. We demonstrate that this regulation occurs between the two photosystems—in contrast to PSII, PSI became more oxidised and the rate constant for P700 re-reduction decreased under these conditions. Thus, when carbon fixation is inhibited, electron transport is down-regulated to match the reduced requirement for electrons and minimise reactive oxygen production. At the same time non-photochemical quenching (NPQ) increases, alleviating the excitation pressure placed on PSII. We observe an increase in the proportion of PSI centres that are active (i.e. can be oxidised with a saturating flash and then rapidly re-reduced) under the conditions when NPQ is increased. We suggest that these additional centres are primarily involved in cyclic electron transport, which generates the pH to support NPQ and protect PSII.Abbreviations A assimilation rate - Ci internal CO₂ concentration - ETC electron transport chain - g stomatal conductance - FR far red - k pseudo first-order rate constant for the reduction of oxidised P700 - NPQ non-photochemical quenching - P700 primary electron donor of photosystem I - PSI, PSII photosystem I, II - qP proportion of open PSII centres - ROS reactive oxygen species - pH pH gradient across the thylakoid membrane - PSII quantum yield of photosystem II An erratum to this article can be found at     

In vivo temperature dependence of cyclic and pseudocyclic electron transport in barley

The effect of temperature on the rate of electron transfer through photosystems I and II (PSI and PSII) was investigated in leaves of barley (Hordeum vulgare L.). Measurements of PSI and PSII photochemistry were made in 21% O₂ and in 2% O₂, to limit electron transport to O₂ in the Mehler reaction. Measurements were made in the presence of saturating CO₂ concentrations to suppress photorespiration. It was observed that the O₂ dependency of PSII electron transport is highly temperature dependent. At 10 °C, the quantum yield of PSII (ΦPSII) was insensitive to O₂ concentration, indicating that there was no Mehler reaction operating. At high temperatures (>25 °C) a substantial reduction in ΦPSII was observed when the O₂ concentration was reduced. However, under the same conditions, there was no effect of O₂ concentration on the ΔpH-dependent process of non-photochemical quenching. The rate of electron transport through PSI was also found to be independent of O₂ concentration across the temperature range. We conclude that the Mehler reaction is not important in maintaining a thylakoid proton gradient that is capable of controlling PSII activity, and present evidence that cyclic electron transport around PSI acts to maintain membrane energisation at low temperature. Received: 6 July 2000 / Accepted: 3 August 2000     

Arundo donax L. (Poaceae) — a C3 Species with Unusually High Photosynthetic Capacity

Gas exchange characteristics, chlorophyll a fluorescence and leaf water potential were investigated in the giant reed, Arundo donax, under natural conditions in an estuarine mangrove swamp in Durban, South Africa. Maximum photosynthetic CO₂ uptake ranged between 19.8 and 36.7 μmol m^?2 s^?1, depending on irradiance, and appeared to be regulated by leaf conductance. There was no saturation of CO₂ uptake or electron transport through PSII (ETR) with increasing irradiance up to 2500 μmol photons m^?2 s^?1. A linear relationship between CO₂ uptake, corrected for respiration (A), and ETR has only been reported for C₄ species and C₃ species when photorespiration is eliminated. From this relationship, it was calculated that 8.5 electrons were transported through PSII for the fixation of one mole of CO₂. Predawn leaf water potential was about ?0.5 MPa and decreased to ?1.5 MPa on a cloudy day and to ?2.1 MPa on a clear day. Diurnal change in leaf water potential had little influence on leaf conductance and hence CO₂ uptake. The molar water use efficiency (WUE) ranged between 4.1 and 9.3 μmol mmol^?1. Percentage photorespiration was between 36 and 39%.     

Relationship between the Quantum Efficiencies of Photosystems I and II in Pea Leaves

The irradiance dependence of the efficiencies of photosystems I and II were measured for two pea (Pisum sativum [L.]) varieties grown under cold conditions and one pea variety grown under warm conditions. The efficiencies of both photosystems declined with increasing irradiance for all plants, and the quantum efficiency of photosystem I electron transport was closely correlated with the quantum efficiency of photosystem II electron transport. In contrast to the consistent pattern shown by efficiency of the photosystems, the redox state of photosystem II (as estimated from the photochemical quenching coefficient of chlorophyll fluorescence) exhibited relationships with both irradiance and the reduction of P-700 that varied with growth environment and genotype. This variability is considered in the context of the modulation of photosystem II quantum efficiency by both photochemical and nonphotochemical quenching of excitation energy.     

Effect of light quality on chloroplast-membrane organization and function in pea

The effect of light quality during plant growth of chloroplast membrane organization and function in peas (Pisum sativum L. cv. Alaska) was investigated. In plants grown under photosystem (PS) I-enriched (far-red enriched) illumination both the PSII/PSI stoichiometry and the electrontransport capacity ratios were high, about 1.9. In plants grown under PSII-enriched (far-red depleted) illumination both the PSII/PSI stoichiometry and the electron-transport capacity ratios were significantly lower, about 1.3. In agreement, steady-state electron-transport measurements under synchronous illumination of PSII and PSI demonstrated an excess of PSII in plants grown under far-red-enriched light. Sodium dodecylsulfate polyacrylamide gel electrophoretic analysis of chlorophyll-containing complexes showed greater relative amounts of the PSII reaction center chlorophyll-protein complex in plants grown under farred-enriched light. Additional changes were observed in the ratio of light-harvesting chlorophyll a/b protein to PSII reaction center chlorophyll-protein under the two different light-quality regimes. The results demonstrate the dynamic nature of chloroplast structure and support the notion that light quality is an important factor in the regulation of chloroplast membrane organization and-function.Abbreviations and symbols Chl chlorophyll - CPa PSII reaction center chlorophyll protein complex - CPI PSI chlorophyll protein complex - FR-D light depleted in far-red sensitizing primarily PSII - FR-E light enriched in far-red sensitizing primarily PSI - LHCP PSII light-harvesting chlorophyll a/b protein complex - P ₇₀₀ primary electron donor of PSI - PSI, PSII photosystems I and II, respectively - Q primary electron acceptor of PSII     

Evaluation of photosynthetic activities in thylakoid membranes by means of Fourier transform infrared spectroscopy

Light-induced Fourier transformed infrared (FTIR) difference spectroscopy is a powerful method to study the structures and reactions of redox cofactors involved in the photosynthetic electron transport chain. So far, most of the FTIR studies of the reactions of oxygenic photosynthesis have been performed using isolated photosystem I (PSI) and photosystem II (PSII) preparations, which, however, could be modified during isolation procedures. In this study, we developed a methodology to evaluate the photosynthetic activities of thylakoids using FTIR spectroscopy. FTIR difference spectra upon successive flashes using thylakoids from spinach exhibited signals typical of the S-state cycle at the Mn₄CaO₅ cluster and Q_B reactions in PSII with period-four and -two oscillations, respectively. Similar measurement in the presence of an artificial quinone as an exogenous electron acceptor showed features specific to the S-state cycle. Simulations of the oscillation patterns provided the quantum efficiencies of the S-state cycle and electron transfer in PSII. Moreover, FTIR measurement under continuous illumination on thylakoids in the presence of DCMU showed signals due to Q_A reduction and P700 oxidation simultaneously. From the relative amplitudes of marker bands of Q_A^? and P700⁺, the molar ratio of photoactive PSII and PSI centers in thylakoids was estimated. FTIR analyses of the photo-reactions in thylakoids, which are more intact than isolated photosystems, will be useful in investigations of the photosynthetic mechanism especially by genetic modification of photosystem proteins.     

Effects of O(2) and CO(2) Concentrations on Quantum Yields of Photosystems I and II in Tobacco Leaf Tissue

The interactive effects of irradiance and O₂ and CO₂ levels on the quantum yields of photosystems I and II have been studied under steady-state conditions at 25°C in leaf tissue of tobacco (Nicotiana tabacum). Assessment of radiant energy utilization in photosystem II was based on changes in chlorophyll fluorescence yield excited by a weak measuring beam of modulated red light. Independent estimates of photosystem I quantum yield were based on the light-dark in vivo absorbance change at 830 nanometers, the absorption band of P700⁺. Normal (i.e. 20.5%, v/v) levels of O₂ generally enhanced photosystem II quantum yield relative to that measured under 1.6% O₂ as the irradiance approached saturation. Photorespiration is suspected to mediate such positive effects of O₂ through increases in the availability of CO₂ and recycling of orthophosphate. Conversely, at low intercellular CO₂ concentrations, 41.2% O₂ was associated with lower photosystem II quantum yield compared with that observed at 20.5% O₂. Inhibitory effects of 41.2% O₂ may occur in response to negative feedback on photosystem II arising from a build-up in the thylakoid proton gradient during electron transport to O₂. Covariation between quantum yields of photosystems I and II was not affected by concentrations of either O₂ or CO₂. The dependence of quantum yield of electron transport to CO₂ measured by gas exchange upon photosystem II quantum yield as determined by fluorescence was unaffected by CO₂ concentration.