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.     

Response of Photochemical Processes of Photosynthesis to Dinitrogen Fixation in Soybean

Symbiotic N2 fixation activity brings about changes in the photochemical processes of photosynthesis in soybean (Glycine max L. Merr.). For a potential photochemical efficiency ([phi]Po) similar to that obtained with an exclusively mineral nutrition, soybean, at full bloom stage (R2) with a moderate N2 fixation activity, had a better electron transfer quantum yield ([phi]PSII) and a higher photochemical quenching. At the beginning seed stage (R5), corresponding to more intense N2 fixation, the same phenomena were enhanced; in addition, an effect on the photochemical (k2b) and nonphotochemical (Kn-k22) transfer rates and an earlier activation of the electron transfer chain were characterized using a new parameter, the relative induction time of PSII fluorescence (Ap/Fm). The response of the photochemical parameters was related to the N2 fixation level (performance of the host plant-microsymbiont association): the energetic cost of symbiotic N2 fixation appeared to be met by a better photochemical efficiency of photosynthesis coupled with a decrease in thermal dissipation (kn - k22), by faster thylakoid energization, and by faster reopening of photosystem II centers at the time of fluorescence induction, as shown by decreased Ap/Fm.     

D,L-甘油醛处理引起的小麦叶片表观光合量子效率的降低（英文）

D,L-甘油醛(磷酸核酮糖激酶抑制剂,10mmol/L)处理小麦旗叶1 h可降低叶片净光合速率和表观量子效率.同时,光系统Ⅱ光化学效率(△F/Fm′)、电子传递速率(ETR)和单位叶面积ATP含量均降低,而胞间二氧化碳浓度(Ci)和叶绿素荧光非光化学猝灭(NPQ)增加.这些结果说明,D,L-甘油醛引起的小麦旗叶表观量子效率降低是由于光合碳同化受阻对光合电子传递的反馈抑制.     

Modifications in photosystem II photochemistry in senescent leaves of maize plants

Analyses of CO2 exchange and chlorophyll fluorescence were carried out to assess photosynthetic performance during senescence of maize leaves. Senescent leaves displayed a significant decrease in CO2 assimilatory capacity accompanied by a decrease in stomatal conductance and an increase in intercellular CO2 concentration. The analyses of fluorescence quenching under steady-state photosynthesis showed that senescence resulted in an increase in non-photochemical quenching and a decrease in photo-chemical quenching. It also resulted in a decrease in the efficiency of excitation energy capture by open PSII reaction centres and the quantum yield of PSII electron transport, but had very little effect on the maximal efficiency of PSII photochemistry. The results determined from the fast fluorescence induction kinetics indicated an increase in the proportion of QB-non-reducing PSII reaction centres and a decrease in the rate of QA reduction in senescent leaves. Theoretical analyses of fluorescence parameters under steady-state photosynthesis suggest that the increase in the non-photochemical quenching was due to an increase in the rate constant to thermal dissipation of excitation energy by PSII and that the decrease in the quantum yield of PSII electron transport was associated with a decrease in the rate constant of PSII photochemistry. Based on these results, it is suggested that the decrease in the quantum yield of PSII electron transport in senescent leaves was down-regulated by an increase in the proportion of QB-non-reducing PSII reaction centres and in the non-photochemical quenching. The photosynthetic electron transport would thus match the decreased demand for ATP and NADPH in carbon assimilation which was inhibited significantly in senescent leaves.Key words: Chlorophyll fluorescence, gas exchange, maize (Zea mays L.), photochemical and non-photochemical quenching, photosystem II photochemistry.      

Chlamydomonas Xanthophyll Cycle Mutants Identified by Video Imaging of Chlorophyll Fluorescence Quenching

The photosynthetic apparatus in plants is protected against oxidative damage by processes that dissipate excess absorbed light energy as heat within the light-harvesting complexes. This dissipation of excitation energy is measured as nonphotochemical quenching of chlorophyll fluorescence. Nonphotochemical quenching depends primarily on the [delta]pH that is generated by photosynthetic electron transport, and it is also correlated with the amounts of zeaxanthin and antheraxanthin that are formed from violaxanthin by the operation of the xanthophyll cycle. To perform a genetic dissection of nonphotochemical quenching, we have isolated npq mutants of Chlamydomonas by using a digital video-imaging system. In excessive light, the npq1 mutant is unable to convert violaxanthin to antheraxanthin and zeaxanthin; this reaction is catalyzed by violaxanthin de-epoxidase. The npq2 mutant appears to be defective in zeaxanthin epoxidase activity, because it accumulates zeaxanthin and completely lacks antheraxanthin and violaxanthin under all light conditions. Characterization of these mutants demonstrates that a component of nonphotochemical quenching that develops in vivo in Chlamydomonas depends on the accumulation of zeaxanthin and antheraxanthin via the xanthophyll cycle. However, observation of substantial, rapid, [delta]pH-dependent nonphotochemical quenching in the npq1 mutant demonstrates that the formation of zeaxanthin and antheraxanthin via violaxanthin de-epoxidase activity is not required for all [delta]pH-dependent nonphotochemical quenching in this alga. Furthermore, the xanthophyll cycle is not required for survival of Chlamydomonas in excessive light.     

Inorganic Carbon Accumulation Stimulates Linear Electron Flow to Artificial Electron Acceptors of Photosystem I in Air-Grown Cells of the Cyanobacterium Synechococcus UTEX 625

The effect of inorganic carbon (Ci) transport and accumulation on photosynthetic electron transport was studied in air-grown cells of the cyanobacterium Synechococcus UTEX 625. When the cells were depleted of Ci, linear photosynthetic electron flow was almost completely inhibited in the presence of the photosystem I (PSI) acceptor N,N-dimethyl-p-nitrosoaniline (PNDA). The addition of Ci to these cells, in which CO2 fixation was inhibited with glycolaldehyde, greatly stimulated linear electron flow and resulted in increased levels of photochemical quenching and O2 evolution. In aerobic conditions substantial quenching resulted from methyl viologen (MV) addition and further quenching was not observed upon the addition of Ci. In anaerobic conditions MV addition did not result in quenching until Ci was added. Intracellular Ci pools were formed when MV was present in aerobic or anaerobic conditions or PNDA was present in aerobic conditions. There was no inhibitory effect of Ci depletion on electron flow to 2,6-dimethylbenzoquinone and oxidized diaminodurene, which accept electrons from photosystem II. The degree of stimulation of PNDA-dependent O2 evolution varied with the Ci concentration. The extracellular Ci, concentration required for a half-maximum rate (K1/2) was 3.8 [mu]M and the intracellular K1/2 was 1.4 mM for the stimulation of PNDA reduction. These values agreed closely with the K1/2 values of extracellular and intracellular Ci for O2 photoreduction. Linear electron flow to artificial electron acceptors of PSI was enhanced by intracellular Ci, which appeared to exert an effect on PSI or on the intersystem electron transport chain.     

Relationship between Steady-State Fluorescence Yield and Photosynthetic Efficiency in Spinach Leaf Tissue

The relationship between steady-state photosynthetic efficiency, as moles CO₂ per mole of incident visible photons under 2% O₂, and chlorophyll fluorescence quenching has been investigated in intact leaf tissue of Spinacia oleracia. Fluorescence yield was measured using a pulse amplitude modulation technique that permitted rapid and sensitive resolution and quantitation of photochemical and nonphotochemical quenching coefficients. A highly linear relationship was observed between photosynthetic efficiency and the ratio of photochemical:nonphotochemical quenching coefficients for values of the latter less than 1.6. This relationship applied whether irradiance or CO₂ concentration was varied. The observed relationships between photochemical yield and fluorescence yield were compatible with the photosystem II model proposed by Butler and Kitajima (1975 Biochim Biophys Acta 376: 116-125). The results are discussed with respect to the proposed role of nonphotochemical quenching in regulating radiant energy utilization and also the applicability of fluorescence measurements as a means of estimation of the rate of photosynthetic electron transport.     

Photochemical and Nonphotochemical Fluorescence Quenching Processes in the Diatom Phaeodactylum tricornutum

Nonphotochemical fluorescence quenching was found to exist in the dark-adapted state in the diatom Phaeodactylum tricornutum. Pretreatment of cells with the uncoupler carbonylcyanide m-chlorophenylhydrazone (CCCP) or with nigericin resulted in increases in dark-adapted minimum and maximum fluorescence yields. This suggests that a pH gradient exists across the thylakoid membrane in the dark, which serves to quench fluorescence levels nonphotochemically. The physiological processes involved in establishing this proton gradient were sensitive to anaerobiosis and antimycin A. Based on these results, it is likely that this energization of the thylakoid membrane is due in part to chlororespiration, which involves oxygen-dependent electron flow through the plastoquinone pool. Chlororespiration has been shown previously to occur in diatoms. In addition, we observed that cells treated with 3-(3,4-dichlorophenyl)-1,1-dimethylurea exhibited very strong nonphotochemical quenching when illuminated with actinic light. The rate and extent of this quenching were light-intensity dependent. This quenching was reversed upon addition of CCCP or nigericin and was thus due primarily to the establishment of a pH gradient across the thylakoid membrane. Preincubation of cells with CCCP or nigericin or antimycin A completely abolished this quenching. Cyclic electron transport processes around photosystem I may be involved in establishing this proton gradient across the thylakoid membrane under conditions where linear electron transport is inhibited. At steady state under normal physiological conditions, the qualitative changes in photochemical and nonphotochemical fluorescence quenching at increasing photon flux densities were similar to those in higher plants. However, important quantitative differences existed at limiting and saturating intensities. Dissimilarities in the factors that regulate fluorescence quenching mechanisms in these organisms may account for these differences.     

CO2 fixation and chlorophyll a fluorescence in leaves of Ramonda serbica during a dehydration-rehydration cycle

Changes in CO(2) photo-assimilation and PSII photochemical efficiency in Ramonda serbica leaves during a dehydration-rehydration cycle were examined. The rate of CO(2) photo-assimilation was greatly reduced during dehydration, but recovery was complete with rehydration when the relative water content of leaves reached values similar to those of well-hydrated, control leaves. The results showed that the response of R. serbica leaves to severe water stress involves two different mechanisms. In the first, CO(2) assimilation is limited by stomata closure that creates an excess proton concentration in the lumen and activates non-photochemical quenching. This plays an important role in the mechanism of photoprotection by dissipation of excitation energy. When dehydration became severe and leaf RWC reached very low values, the electron transport rate (ETR) decreased markedly, while the capacity for regulatory mechanisms such as q(NP) (non-photochemical quenching) was greatly reduced. For severely dehydrated leaves of R. serbica, it appears that reactive oxygen species (ROS) formation is better prevented by mechanisms that quench chlorophyll triplet formation via lutein.     

Chloroplastic NAD(P)H dehydrogenase in tobacco leaves functions in alleviation of oxidative damage caused by temperature stress

In this study, the function of the NAD(P)H dehydrogenase (NDH)-dependent pathway in suppressing the accumulation of reactive oxygen species in chloroplasts was investigated. Hydrogen peroxide accumulated in the leaves of tobacco (Nicotiana tabacum) defective in ndhC-ndhK-ndhJ (DeltandhCKJ) at 42 degrees C and 4 degrees C, and in that of wild-type leaves at 4 degrees C. The maximum quantum efficiency of PSII decreased to a similar extent in both strains at 42 degrees C, while it decreased more evidently in DeltandhCKJ at 4 degrees C. The parameters linked to CO(2) assimilation, such as the photochemical efficiency of PSII, the decrease of nonphotochemical quenching following the initial rise, and the photosynthetic O(2) evolution, were inhibited more significantly in DeltandhCKJ than in wild type at 42 degrees C and were seriously inhibited in both strains at 4 degrees C. While cyclic electron flow around PSI mediated by NDH was remarkably enhanced at 42 degrees C and suppressed at 4 degrees C. The proton gradient across the thylakoid membranes and light-dependent ATP synthesis were higher in wild type than in DeltandhCKJ at either 25 degrees C or 42 degrees C, but were barely formed at 4 degrees C. Based on these results, we suggest that cyclic photophosphorylation via the NDH pathway might play an important role in regulation of CO(2) assimilation under heat-stressed condition but is less important under chilling-stressed condition, thus optimizing the photosynthetic electron transport and reducing the generation of reactive oxygen species.     

Characteristics of photosynthesis and functions of the water-water cycle in rice (Oryza sativa) leaves in response to potassium deficiency

The mechanisms of photoprotection of photosynthesis and dissipation of excitation energy in rice leaves in response to potassium (K) deficiency were investigated. Net photosynthetic rate and the activity of ribulose-1,5-bisphosphate carboxylase/oxygenase decreased under K deficiency. Compared with the control, non-photochemical quenching of Chl fluorescence increased in K-deficient plant, whereas the efficiency of excitation transfer (F'(v)/F'(m)) and the photochemical quenching coefficient (q(P)) decreased. Thus, thermal dissipation of excitation energy increased as more excess electrons were accumulated in the photosynthetic chain. The electron transport rate through PSII (J(f)) was more sensitive to O2 concentration, and the fraction of electron transport rate required to sustain CO2 assimilation and photorespiration (J(g)/J(f)) was significantly decreased under K deficiency compared with the control. Furthermore, the alternative electron transport (J(a)/J(f)) was increased, indicating that a considerable amount of electrons had been transported to O2 during the water-water cycle in the K-deficient leaves. Although the fraction of electron transport to photorespiration (J(o)/J(f)) was also increased in the K-deficient leaves, it was less sensitive than that of the water-water cycle. With the generation of reactive oxygen species level, the activities of superoxide dismutase and ascorbate peroxidase, two of the key enzymes involved in scavenging of active oxygen species in the water-water cycle, also increased in K-deficient rice. Therefore, it is likely that a series of photoprotective mechanisms were initiated in rice plants in response to K deficiency and the water-water cycle might be critical for protecting photosynthetic apparatus under K deficiency in rice.     

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.     

硫对烟草叶片光合特性和叶绿素荧光参数的影响

通过液培试验,测定了不同硫营养水平(0～32 mmol/L)下烟草叶片的叶绿素含量、气体交换参数和叶绿素荧光参数.结果表明,随着硫浓度的升高,烟草叶片的Chl.a、Chl.b、总Chl的含量和Chl.a/b逐渐增加;类胡萝卜素呈先下降后升高的趋势;烟草叶片的净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)、胞间CO2浓度(Ci)下降显著,气孔限制值(Ls)明显升高,这表明高硫对烟草叶片的气体交换参数影响较大.随着硫浓度的升高,烟草叶片光合有效量子产量(EQY)、光合电子传递速率(ETR)、光化学淬灭(qP)都表现为先升高后降低,一般是4 mmol/L和8 mmol/L处理相对较高,浓度超过8 mmol/L以后,以上参数显著下降.非光化学淬灭(NPQ) 0,16 mmol/L和32 mmol/L处理表现相对较高,2、4、8 mmol/L处理的相对较低,2～8 mmol/L浓度的处理最适宜烟草的生长,高硫或低硫都会导致烟草EQY、ETR、qP降低和NPQ的升高.     

Mehler-peroxidase reaction mediates zeaxanthin formation and zeaxanthin-related fluorescence quenching in intact chloroplasts

Induction of zeaxanthin formation and the associated nonphotochemical quenching in iodoacetamide-treated, non-CO₂-fixing intact chloroplasts of Lactuca sativa L. cv Romaine is reported. The electron transport needed to generate the required ΔpH for zeaxanthin formation and nonphotochemical quenching are ascribed to the Mehler-ascorbate peroxidase reaction. KCN, an inhibitor of ascorbate peroxidase, significantly affected these activities without affecting linear electron transport to methyl viologen or violaxanthin deepoxidase activity. At 1 millimolar KCN, zeaxanthin formation and ΔpH were inhibited 60 and 55%, respectively, whereas ascorbate peroxidase activity was inhibited almost totally. The KCN-resistant activity, which apparently was due to electron transport mediated by the Mehler reaction alone, however, was insufficient to support a high level of nonphotochemical quenching. We suggest that in vivo, as CO₂ fixation becomes limiting, the Mehler-peroxidase reaction protects photosystem II against the excess light by supporting the electron transport needed for zeaxanthin-dependent nonphotochemical quenching and concomitantly scavenging H₂O₂. Ascorbate is essential for this process to occur.     

Effect of Elevated CO2 on the Primary Conversion of Light Energy of Alfalfa Photosynthesis

In studying the mechanism of increase in alfalfa (Medicago sativa L. ) photosynthesis under elevated atmospheric CO2, it was found that the capacity of chloroplasts for light absorption was greater, the potential activity and efficiency of primary conversion of light enlergy of PS Ⅱ , quantum yield of PS Ⅱ electron transport, and activation capacity of PS Ⅰ were stimulated, photochemical quenching coefficient was increased and non-photochemical quenching coefficient was decreased under elevated atmospheric CO2.     

Theoretical study of the pulse method for the quenching analysis of photosystem II fluorescence

The kinetics of photochemical (qQ) and nonphotochemical (qE) fluorescence quenching during induction was studied in terms of the earlier developed theoretical model of photosynthesis. Photochemical and nonphotochemical quenching was calculated from the parameters of fluorescence induction upon simultaneous excitation with continuous light and by high-intensity light pulses by the method used in a PAM fluorometer. It is shown that the fraction of closed reaction centers during the pulse can change in the course of induction. In consequence, even when none of supposed mechanisms of photochemical quenching is taken into account, the value of qE during induction varies and is not equal to zero.     

Photosystem II cycle and alternative electron flow in leaves

Sunflower (Helianthus annuus L.) and tobacco (Nicotiana tabacum L.) were grown in the laboratory and leaves were taken from field-grown birch trees (Betula pendula Roth). Chlorophyll fluorescence, CO2 uptake and O2 evolution were measured and electron transport rates were calculated, J(C) from the CO2 uptake rate considering ribulose-1,5-bisphosphate (RuBP) carboxylation and oxygenation, J(O) from the O2 evolution rate, and J(F) from Chl fluorescence parameters. Mesophyll diffusion resistance, r(md), used for the calculation of J(C), was determined such that the in vivo Rubisco kinetic curve with respect to the carboxylation site CO2 concentration became a rectangular hyperbola with Km(CO2) of 10 microM at 22.5 degrees C. In sunflower, in the absence of external O2, J(O) = 1.07 J(C) when absorbed photon flux density (PAD) was varied, showing that the O2-independent components of the alternative electron flow to acceptors other than CO2 made up 7% of J(C). Under saturating light, J(F), however, was 20-30% faster than J(C), and J(F)-J(C) depended little on CO2 and O2 concentrations. The inter-relationship between J(F)-J(C) and non-photochemical quenching (NPQ) was variable, dependent on the CO2 concentration. We conclude that the relatively fast electron flow J(F)-J(C) appearing at light saturation of photosynthesis contains a minor component coupled with proton translocation, serving for nitrite, oxaloacetate and oxygen reduction, and a major component that is mostly cyclic electron transport around PSII. The rate of the PSII cycle is sufficient to release the excess excitation pressure on PSII significantly. Although the O2-dependent Mehler-type alternative electron flow appeared to be under the detection threshold, its importance is discussed considering the documented enhancement of photosynthesis by oxygen.     

The relationship between photosynthesis and a mastoparan-induced hypersensitive response in isolated mesophyll cells

The G-protein activator mastoparan (MP) was found to elicit the hypersensitive response (HR) in isolated Asparagus sprengeri mesophyll cells at micromolar concentrations. The HR was characterized by cell death, extracellular alkalinization, and an oxidative burst, indicated by the reduction of molecular O₂ to O₂⁻. To our knowledge, this study was the first to monitor photosynthesis during the HR. MP had rapid and dramatic effects on photosynthetic electron transport and excitation energy transfer as determined by variable chlorophyll a fluorescence measurements. A large increase in nonphotochemical quenching of chlorophyll a fluorescence accompanied the initial stages of the oxidative burst. The minimal level of fluorescence was also quenched, which suggests the origin of this nonphotochemical quenching to be a decrease in the antenna size of photosystem II. In contrast, photochemical quenching of fluorescence decreased dramatically during the latter stages of the oxidative burst, indicating a somewhat slower inhibition of photosystem II electron transport. The net consumption of O₂ and the initial rate of O₂ uptake, elicited by MP, were higher in the light than in the dark. These data indicate that light enhances the oxidative burst and suggest a complex relationship between photosynthesis and the HR.     

Low-Temperature Effects on Photosynthesis and Correlation with Freezing Tolerance in Spring and Winter Cultivars of Wheat and Rye

Winter cultivars of rye (Secale cereale L., cv Musketeer) and wheat (Triticum aestivum L. cvs Kharkov and Monopol), but not a spring cultivar of wheat (Glenlea), grown at cold-hardening temperatures showed, at high irradiances, a higher proportion of oxidized to reduced primary, stable quinone receptor (QA) than did the same cultivars grown under nonhardening conditions. In addition, there was a positive correlation between the effects of low-growth temperature on this increased proportion of oxidized QA, and a concomitant increase in the capacity for photosynthesis, and LT50, the temperature at which 50% of the seedlings are killed, in cultivars showing different freezing tolerances. This suggests that low-temperature modulation of the photosynthetic apparatus may be an important factor during the induction of freezing resistance in cereals. Finally, the control of photosystem II photochemistry by nonphotochemical quenching of excitation energy was identical for nonhardened and cold-hardened winter rye. However, examination of measuring temperature effects per se revealed that, irrespective of growth temperature, nonphotochemical quenching exerted a stronger control on photosystem II photochemistry at 10[deg] C rather than at 20[deg] C.     

Carbon Dioxide-Induced Oscillations in Fluorescence and Photosynthesis: Role of Thylakoid Membrane Energization in Regulation of Photosystem II Activity

The response of CO₂ fixation to a sudden increase in ambient CO₂ concentration has been investigated in intact leaf tissue from spinach (Spinacia oleracea) using a dual channel infrared gas analyzer. Simultaneous with these measurements, changes in fluorescence emission associated with a weak, modulated measuring beam were recorded. Application of brief (2-3 seconds) dark intervals enabled estimation of the dark fluorescence level (F_o) under both steady state and transient conditions. The degree of suppression of F_o level fluorescence in the light was strongly correlated with nonphotochemical quenching under all conditions. During CO₂-induced oscillations in photosynthesis under 2% O₂ the changes in nonphotochemical quenching anticipate changes in the rate of uptake of CO₂. At such low levels of O₂ and constant illumination, changes in the relative quantum efficiency of open photosystem II units were estimated as the ratio of the rate of CO₂ uptake and the photochemical quenching coefficient. Under the same conditions the relative quantum efficiency of photosystem II was found to vary inversely with the degree of nonphotochemical quenching. The relationship between changes in the rate of CO₂ uptake: photochemical quenching coefficient and nonphotochemical quenching was altered somewhat when the same experiment was conducted under 20% O₂. The results suggest that electron transport coupled to reduction of O₂ occurs to varying degrees with time during oscillations, especially when ambient O₂ concentrations are high.