Pseudocyclic Electron Transport Research Articles

Molecular Genetic Dissection of the Regulatory Network of Proton Motive Force in Chloroplasts.

The proton motive force (pmf) generated across the thylakoid membrane rotates the Fo-ring of ATP synthase in chloroplasts. The pmf comprises two components: membrane potential (∆Ψ) and proton concentration gradient (∆pH). Acidification of the thylakoid lumen resulting from ∆pH downregulates electron transport in the cytochrome b6f complex. This process, known as photosynthetic control, is crucial for protecting photosystem I (PSI) from photodamage in response to fluctuating light. To optimize the balance between efficient photosynthesis and photoprotection, it is necessary to regulate pmf. Cyclic electron transport around PSI and pseudo-cyclic electron transport involving flavodiiron proteins contribute to the modulation of pmf magnitude. By manipulating the ratio between the two components of pmf, it is possible to modify the extent of photosynthetic control without affecting the pmf size. This adjustment can be achieved by regulating the movement of ions (such as K+ and Cl-) across the thylakoid membrane. Since ATP synthase is the primary consumer of pmf in chloroplasts, its activity must be precisely regulated to accommodate other mechanisms involved in pmf optimization. Although fragments of information about each regulatory process have been accumulated, a comprehensive understanding of their interactions is lacking. Here, I summarize current knowledge of the network for pmf regulation, mainly based on genetic studies.

Revealing the significance of chlorophyll b in the moss Physcomitrium patens by knocking out two functional chlorophyllide a oxygenase.

The chlorophyllide a oxygenase (CAO) plays a crucial role in the biosynthesis of chlorophyll b (Chl b). In the moss Physcomitrium patens (P. patens), two distinct gene copies, PpCAO1 and PpCAO2, are present. In this study, we investigate the differential expression of these CAOs following light exposure after a period of darkness (24h) and demonstrate that the accumulation of Chl b is only abolished when both genes are knocked out. In the ppcao1cao2 mutant, most of the antenna proteins associated with both photosystems (PS) I and II are absent. Despite of the existence of LHCSR proteins and zeaxanthin, the mutant exhibits minimal non-photochemical quenching (NPQ) capacity. Nevertheless, the ppcao1cao2 mutant retains a certain level of pseudo-cyclic electron transport to provide photoprotection for PSI. These findings shed light on the dual dependency of Chl b synthesis on two CAOs and highlight the distinct effects of Chl b deprival on PSI and PSII core complexes in P. patens, a model species for bryophytes.

The water-water cycle facilitates photosynthetic regulation under fluctuating light in the epiphytic orchid Dendrobium officinale

Fluctuating light can cause selective photoinhibition of photosystem I (PSI) in angiosperms. However, the strategies for photosynthetic regulation under fluctuating light in wild epiphytic plants are little known. We here compared PSI and PSII performances under fluctuating light in the epiphytic orchid Dendrobium officinale and the terrestrial orchid Bletilla striata. Upon dark-to-light transition, D. officinale showed a rapid re-oxidation of P700, but this rapid re-oxidation of P700 was not observed in B. striata. Within the first 40 s after transition from dark to actinic light, B. striata showed much higher activation of cyclic electron transport (CET) than D. officinale. However, this strong activation of CET could not prevent the over-reduction of PSI. Therefore, the rapid re-oxidation of P700 in D. officinale was attributed to pseudo-cyclic electron transport (water-water cycle). Within the first 60 s after transition from low to high light, PSI was highly oxidized in D. officinale but was highly reduced in B. striata. Concomitantly, B. striata showed much higher stimulation of CET. These results indicate that water-water cycle significantly regulates PSI redox state under fluctuating light in the epiphytic orchid D. officinale. By comparison, the terrestrial orchid B. striata employs CET to cope with fluctuating light, similar to the phenotype of Arabidopsis thaliana. Taking together, these two orchids used different strategies for photosynthetic regulation to cope with fluctuating light.

Temperature-dependent regulation of electron transport and ATP synthesis in chloroplasts in vitro and in silico.

The significance of temperature-dependent regulation of photosynthetic apparatus (PSA) is determined by the fact that plant temperature changes with environmental temperature. In this work, we present a brief overview of temperature-dependent regulation of photosynthetic processes in class B chloroplasts (thylakoids) and analyze these processes using a computer model that takes into account the key stages of electron and proton transport coupled to ATP synthesis. The rate constants of partial reactions were parametrized on the basis of experimental temperature dependences of partial photosynthetic processes: (1) photosystem II (PSII) turnover and plastoquinone (PQ) reduction, (2) the plastoquinol (PQH2) oxidation by the cytochrome (Cyt) b6f complex, (3) the ATP synthase activity, and (4) the proton leak from the thylakoid lumen. We consider that PQH2 oxidation is the rate-limiting step in the intersystem electron transport. The parametrization of the rate constants of these processes is based on earlier experimental data demonstrating strong correlations between the functional and structural properties of thylakoid membranes that were probed with the lipid-soluble spin labels embedded into the membranes. Within the framework of our model, we could adequately describe a number of experimental temperature dependences of photosynthetic reactions in thylakoids. Computer modeling of electron and proton transport coupled to ATP synthesis supports the notion that PQH2 oxidation by the Cyt b6f complex and proton pumping into the lumen are the basic temperature-dependent processes that determine the overall electron flux from PSII to molecular oxygen and the net ATP synthesis upon variations of temperature. The model describes two branches of the temperature dependence of the post-illumination reduction of [Formula: see text] characterized by different activation energies (about 60 and ≤ 3.5kJmol-1). The model predicts the bell-like temperature dependence of ATP formation, which arises from the balance of several factors: (1) the thermo-induced acceleration of electron transport through the Cyt b6f complex, (2) deactivation of PSII photochemistry at sufficiently high temperatures, and (3) acceleration of the passive proton outflow from the thylakoid lumen bypassing the ATP synthase complex. The model describes the temperature dependence of experimentally measured parameter P/2e, determined as the ratio between the rates of ATP synthesis and pseudocyclic electron transport (H2O → PSII → PSI → O2).

The chloroplast NADH dehydrogenase-like complex influences the photosynthetic activity of the moss Physcomitrella patens

Alternative electron pathways contribute to regulation of photosynthetic light reactions to adjust to metabolic demands in dynamic environments. The chloroplast NADH dehydrogenase-like (NDH) complex mediates the cyclic electron transport pathway around PSI in different cyanobacteria, algae, and plant species, but it is not fully conserved in all photosynthetic organisms. In order to assess how the physiological role of this complex changed during plant evolution, we isolated Physcomitrella patens lines knocked out for the NDHM gene that encodes a subunit fundamental for the activity of the complex. ndhm knockout mosses indicated high PSI acceptor side limitation upon abrupt changes in illumination. In P. patens, pseudo-cyclic electron transport mediated by flavodiiron proteins (FLVs) was also shown to prevent PSI over-reduction in plants exposed to light fluctuations. flva ndhm double knockout mosses had altered photosynthetic performance and growth defects under fluctuating light compared with the wild type and single knockout mutants. The results showed that while the contribution of NDH to electron transport is minor compared with FLV, NDH still participates in modulating photosynthetic activity, and it is critical to avoid PSI photoinhibition, especially when FLVs are inactive. The functional overlap between NDH- and FLV-dependent electron transport supports PSI activity and prevents its photoinhibition under light variations.

PH-Dependent Regulation of Electron and Proton Transport in Chloroplasts In Situ and In Silico

The analysis of electron and proton transport in chloroplasts of higher plants has been carried out on the basis of a mathematical model that takes into account the pH-dependent regulation of electron transport and the operation of the ATP synthase. Numerical experiments aimed at simulation of these processes under pseudocyclic electron transport (water–water cycle) have shown good agreement with experimental data on the kinetics of electron transfer to photosystem 1 (PS1) in class B chloroplasts in metabolic states corresponding to high (state 3) and low (state 4) ATP synthase activity. The simulation of electron transport processes that took into account the Calvin–Benson cycle (CBC), cyclic electron transport around PS1, pH-dependent heat dissipation of energy in photosystem 2 (PS2), and nonphotochemical quenching (NPQ) made it possible to estimate the contribution of these factors to the kinetics of induction phenomena in chloroplasts in situ. It has been shown that the multiphase kinetics of the photooxidation of P700 (a primary electron donor in PS1) reflects the redistribution of electron flows between cyclic and non-cyclic electron transfer pathways, caused by the activation of CBC due to the alkalization of the stroma, as well as the change of the limiting stage in the electron transport chain, induced by a decrease in the intrathylakoid pH (pHin). The electron flux between PS2 and PS1 decelerates with pHin decrease, which may be caused by the reduced rate of plastoquinol oxidation and attenuated activity of PS2 due to NPQ.

Role of cyclic and pseudo-cyclic electron transport in response to dynamic light changes in Physcomitrella patens.

Photosynthetic organisms support cell metabolism by harvesting sunlight and driving the electron transport chain at the level of thylakoid membranes. Excitation energy and electron flow in the photosynthetic apparatus is continuously modulated in response to dynamic environmental conditions. Alternative electron flow around photosystem I plays a seminal role in this regulation contributing to photoprotection by mitigating overreduction of the electron carriers. Different pathways of alternative electron flow coexist in the moss Physcomitrella patens, including cyclic electron flow mediated by the PGRL1/PGR5 complex and pseudo-cyclic electron flow mediated by the flavodiiron proteins FLV. In this work, we generated P.patens plants carrying both pgrl1 and flva knock-out mutations. A comparative analysis of the WT, pgrl1, flva, and pgrl1 flva lines suggests that cyclic and pseudo-cyclic processes have a synergic role in the regulation of photosynthetic electron transport. However, although both contribute to photosystem I protection from overreduction by modulating electron flow following changes in environmental conditions, FLV activity is particularly relevant in the first seconds after a light change whereas PGRL1 has a major role upon sustained strong illumination.

Artificial remodelling of alternative electron flow by flavodiiron proteins in Arabidopsis

In photosynthesis, linear electron transport from water to nicotinamide adenine dinucleotide phosphate (NADP(+)) cannot satisfy the ATP/NADPH production stoichiometry required by the Calvin-Benson cycle. Cyclic electron transport (CET) around photosystem I (PSI) and pseudocyclic electron transport (pseudoCET) can produce ATP without the accumulation of NADPH. Flavodiiron proteins (Flv) are the main mediator of pseudoCET in photosynthetic organisms, spanning cyanobacteria to gymnosperms. However, their genes are not conserved in angiosperms. Here we explore the possibility of complementing CET with Flv-dependent pseudoCET in the angiosperm Arabidopsis thaliana. We introduced FlvA and FlvB genes from the moss Physcomitrella patens into both wild-type (WT) Arabidopsis and the proton gradient regulation 5 (pgr5) mutant, which is defective in the main pathway of CET. We measured rates of pseudoCET using membrane inlet mass spectrometry, along with several photosynthetic parameters. Flv expression significantly increased rates of pseudoCET in the mutant plants, particularly at high light intensities, and partially restored the photosynthetic phenotype. In WT plants, Flv did not compete with PGR5-dependent CET during steady-state photosynthesis, but did form a large electron sink in fluctuating light. We conclude that flavodiiron proteins can help to protect the photosystems in Arabidopsis under fluctuating light, even in the presence of CET.

Reactive oxygen intermediates produced by photosynthetic electron transport are enhanced in short-day grown plants

Leaves of tobacco plants grown in short days (8h light) generate more reactive oxygen species in the light than leaves of plants grown in long days (16h light). A two fold higher level of superoxide production was observed even in isolated thylakoids from short day plants. By using specific inhibitors of photosystem II and of the cytochrome b6f complex, the site of O2 reduction could be assigned to photosystem I. The higher rate of O2 reduction led to the formation of a higher proton gradient in thylakoids from short day plants. In the presence of an uncoupler, the differences in O2 reduction between thylakoids from short day and long day plants were abolished. The pigment content and the protein content of the major protein complexes of the photosynthetic electron transport chain were unaffected by the growth condition. Addition of NADPH, but not of NADH, to coupled thylakoids from long day plants raised the level of superoxide production to the same level as observed in thylakoids from short day plants. The hypothesis is put forward that the binding of an unknown protein permits the higher rate of pseudocyclic electron flow in thylakoids from short-day grown plants and that this putative protein plays an important role in changing the proportions of linear, cyclic and pseudocyclic electron transport in favour of pseudocyclic electron transport. This article is part of a Special Issue entitled: Photosynthesis Research for Sustainability: from Natural to Articifical.

Abolition of photosystem I cyclic electron flow in Arabidopsis thaliana following thermal-stress

Heat tolerance of Arabidopsis thaliana (WT) and its mutants, crr2-2, lacking NADPH-dehydrogenase (Ndh-pathway), and pgr5, deficient in proton gradient regulation and/or ferredoxin-quinone-reductase (FQR-pathway), was studied from 30 to 46°C. Chlorophyll fluorescence revealed that thermal damage to photosystem II (PSII) was maximal in WT plants following short-term exposure of leaves to moderate or high temperature stress. Thermal stress impaired the photosynthetic electron flow at oxidizing and reducing sides of PSII. This was deduced from the transformation of temperature dependent OJIP to OKP patterns, changes in the relative amplitudes of K-step fluorescence rise and F(v)/F(o) ratio. The amplitude of the K-peak that corresponds to the magnitude of damage to the oxygen evolving complex (OEC) in crr2-2 mutants was about 50% of that observed in WT plants exposed to 46°C. The damage to OEC in pgr5 mutants was relatively smaller and thus their PSII complexes were more heat tolerant. P700 oxidation-reduction kinetics following heat-stress revealed that photosystem I (PSI) complexes remained oxidizable either with 10-ms multiple turn-over flashes or far-red illumination but the complementary cyclic electron flow around PSI (CEF) was abolished in both mutants. With further increase in incubation temperature, CEF was fully suppressed even in WT. Thus, P700 turn-over was not enhanced following thermal stress. Furthermore, the experimental data predicts the onset of pseudocyclic electron transport with molecular oxygen as terminal acceptor in crr2-2 and pgr5 mutants but not in wild type Arabidopsis subjected to severe thermal-stress.

Light‐dependent oxygen consumption in nitrogen‐fixing cyanobacteria plays a key role in nitrogenase protection1

All colonial diazotrophic cyanobacteria are capable of simultaneously evolving O2 through oxygenic photosynthesis and fixing nitrogen via nitrogenase. Since nitrogenase is irreversibly inactivated by O2, accommodation of the two metabolic pathways has led to biochemical and/or structural adaptations that protect the enzyme from O2. In some species, differentiated cells (heterocysts) are produced within the filaments. PSII is absent in the heterocysts, while PSI activity is maintained. In other, nonheterocystous species, however, a “division of labor” occurs whereby individual cells within a colony appear to ephemerally fix nitrogen while others evolve oxygen. Using membrane inlet mass spectrometry (MIMS) in conjunction with tracer 18O2 and inhibitors of photosynthetic and respiratory electron transport, we examined the light dependence of O2 consumption in Trichodesmium sp. IMS 101, a nonheterocystous, colonial cyanobacterium, and Anabaena flos‐aquae (Lyngb.) Bréb. ex Bornet et Flahault, a heterocystous species. Our results indicate that in both species, intracellular O2 concentrations are maintained at low levels by the light‐dependent reduction of oxygen via the Mehler reaction. In N2‐fixing Trichodesmium colonies, Mehler activity can consume ∼75% of gross O2 production, while in Trichodesmium utilizing nitrate, Mehler activity declines and consumes ∼10% of gross O2 production. Moreover, evidence for the coupling between N2 fixation and Mehler activity was observed in purified heterocysts of Anabaena, where light accelerated O2 consumption by 3‐fold. Our results suggest that a major role for PSI in N2‐fixing cyanobacteria is to effectively act as a photon‐catalyzed oxidase, consuming O2 through pseudocyclic electron transport while simultaneously supplying ATP in both heterocystous and nonheterocystous taxa.

Extension of a biochemical model for the generalized stoichiometry of electron transport limited C3 photosynthesis

ABSTRACTThe widely used steady‐state model of Farquhar et al. (Planta 149: 78–90, 1980) for C3 photosynthesis was developed on the basis of linear whole‐chain (non‐cyclic) electron transport. In this model, calculation of the RuBP‐regeneration limited CO2‐assimilation rate depends on whether it is insufficient ATP or NADPH that causes electron transport limitation. A new, generalized equation that allows co‐limitation of NADPH and ATP on electron transport is presented herein. The model is based on the assumption that other thylakoid pathways (the Q‐cycle, cyclic photophosphorylation, and pseudocyclic electron transport) interplay with the linear chain to co‐contribute to a balanced production of NADPH and ATP as required by stromal metabolism. The original model assuming linear electron transport limited either by NADPH or by ATP, predicts quantum yields for CO2 uptake that represent the highest and the lowest values, respectively, of the range given by the new equation. The applicability of the new equation is illustrated for a number of C3 crop species, by curve fitting to gas exchange data in the literature. In comparison with the original model, the new model enables analysis of photosynthetic regulation via the electron transport pathways in response to environmental stresses.

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 degrees C, the quantum yield of PSII (phi PSII) was insensitive to O2 concentration, indicating that there was no Mehler reaction operating. At high temperatures (> 25 degrees C) a substantial reduction in phi PSII was observed when the O2 concentration was reduced. However, under the same conditions, there was no effect of O2 concentration on the delta pH-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.

Light dependence of quantum yields for PSII charge separation and oxygen evolution in eucaryotic algae

Quantum yields of photosystcm II (PSII) charge separation (ΦP and oxygen production () were determined by simultaneous measurements of oxygen production and variable fluorescence in four different aquatic microalgae representing three different taxonomic groups: the freshwater alga Scenedesmus protuberans (Chlorophyceae) and the marine algae Phaeocystis globosa (Prymnesiophyceae), Emiliania huxleyi (Prymnesiophyceae), and Phaeodactylum tricornutum (Bacillariophyceae). In S. protuberans, P. tricornutum, and E. huxleyi, light‐dependent variability was observed in the ratio of to ΦP i.e. in the number of oxygen molecules produced per electron generated by PSII. The ratio :ΦP was highly variable at low light intensities (E < 0.5 Ek), and at higher light intensities (E > 0.5 EK) :ΦP showed a nonlinear decrease with increasing light intensity. In contrast, in P. globosa a trend in :ΦP could not be distinguished, and this species showed a decrease in :ΦP during the day, indicating a dependency of :ΦP on light history. Additionally, considerable interspecific quantitative differences in :ΦP were observed. Two possible interpretations to explain the variability in :ΦP are discussed. Assuming that ΦP is a reliable measure of the quantum yield for charge separation at PSII, one interpretation is that net oxygen production is influenced by processes that consume oxygen or affect linear electron transport (e.g. cyclic electron transport around PSII, pseudocyclic electron transport in the Mehler reaction, Rubisco oxygenase activity, and lightdependent mitochondrial respiration). A second interpretation, however, suggests that at saturating light, changes in photosynthesis turnover time occur, such that ΦP does not predict the steady‐state O2 yield.

Induction of two types of non-photochemical chlorophyll fluorescence quenching in carbon-assimilating intact spinach chloroplasts: the effects of ascorbate, de-epoxidation, and dibucaine

We show with intact carbon-assimilating chloroplasts the induction of two types of rapidly reversible non-photochemical fluorescence quenching, distinguishable by the effects of ascorbate, de-epoxidation and dibucaine. In the presence of ascorbate, fluorescence quenching is antenna type and obligatory on violaxanthin de-epoxidation; in the absence of ascorbate, fluorescence quenching is reaction-center type and unrelated to violaxanthin de-epoxidation. The latter quenching type is inhibited by ascorbate, characterizing it as donor-side Photosystem II (PS II) inhibition. Consistent with current concepts of down-regulation of PS II efficiency by non-radiative energy dissipation, both types of quenching vary inversely to the putative ATP requirement of the carbon-substrate acceptors, although less clearly in the presence of ascorbate, presumably due to pseudocyclic electron transport by the Mehler-ascorbate-peroxidase reaction. It is concluded from the effects of dibucaine that reaction-center and antenna types of quenching are related to protons in different thylakoid compartments, the former to domains de-localized in the lumen and the latter to domains localized in the membrane. Antenna quenching, however, also requires lumen acidification for the obligatory violaxanthin de-epoxidase activity. No antenna quenching that is unrelated to de-epoxidation was observed. Since ascorbate is usually present in leaves, these results suggest that the prevailing mechanism for energy-dependent non-photochemical fluorescence quenching in vivo is de-epoxidation-dependent antenna quenching.

Ferredoxin–NADP+ reductase is the site of oxygen reduction in pseudocyclic electron transport

The effects of ferredoxin (Fd) and Fd–NADP+ reductase (FNR) on the oxygen photoreduction by photosystem I (PS I) in spinach (Spinacia oleracea L.) thylakoid membranes were investigated using a unique photoelectrochemical cell. This cell was previously shown to monitor the Mehler reaction products of photosynthetic oxygen reduction and represents an excellent tool for studying pseudocyclic electron transport. The magnitude of the photocurrent produced by the thylakoids was increased by as much as 40% in the presence of 60 μM Fd. If thylakoids were supplemented by both Fd and FNR, an additional increase of photocurrent was observed. All these reactions were inhibited by catalase, an enzyme that degrades H2O2, to demonstrate that O2 reduction was involved in all the photoreactions studied. The fact that more O2 was consumed in the presence of FNR was interpreted as evidence that the most effective site of oxygen reduction on the acceptor side of PS I is on FNR and not on Fd. The in vivo implication is that during pseudocyclic electron transport, NADP+ and oxygen directly compete for PS I electrons, with the former having significantly faster reaction kinetics. The advantageous physiological consequences of such a competition are (i) pseudocyclic electron transport would represent a true attenuating mechanism of the redox state of the NADP+–NADPH pool, (ii) oxygen would be a contingent acceptor under high illumination stress, helping to cope with the resultant elevated electron transport rates, and (iii) this mechanism is indisputably a faster response to stress than cyclic electron transport. Key words: Spinacia oleracea, photosystem I, thylakoid membranes, ferredoxin–NADP+ reductase, pseudocyclic electron transport, photoelectrochemistry.

Effects of selenite, CO 2 and illumination on the induction of selenium-dependent glutathione peroxidase in Chlamydomonas reinhardtii

When sodium selenite was added to Chlamydomonas reinhardtii cells grown in the absence of sodium selenite under illumination (10 000 lux) and ordinary air (0.03% CO 2 concentration), glutathione peroxidase activity increased to a peak after 24 h, and ascorbate peroxidase activity concomitantly disappeared after 6 h. The effect of cycloheximide and the immunochemical titration with antibody directed against bovine erythrocyte glutathione peroxidase showed that the increase in glutathione peroxidase activity was due to protein synthesis. Transfer of Chlamydomonas cells either from low CO 2 concentration (ordinary air: 0.03% CO 2) to high CO 2 concentration (5% CO 2) or from light to dark together with the addition of sodium selenite stopped the increase in the glutathione peroxidase activity. The effect of illumination was saturated at a light intensity of about 6000 lux. The results reported here indicate that low CO 2 concentration in the atmosphere and illumination are required for the induction of glutathione peroxidase, and suggest that selenium-induced glutathione peroxidase as well as catalase can play an important role in the scavenging of H 2O 2 generated in pseudocyclic electron transport of Chlamydomonas chloroplasts.

Generation of photocurrents by a chloroplast photosystem I submembrane fraction using the Mehler reaction

Abstract A photosystem I submembrane fraction isolated from spinach leaves is used to generate photocurrents under potentiostatic conditions in a small (80 μl) one compartment electrochemical cell. The photocurrent was generated via a Mehler-type reaction pathway where superoxide radicals are formed during the reduction of dissolved oxygen by photosystem I. The superoxide ions dismutate to hydrogen peroxide which reacts with a platinum working electrode. In the presence of the electron donor durohydroquinone, the photosystem generated relatively strong photocurrents under various conditions. The results demonstrate the suitability of the photoelectrochemical cell to study pseudocyclic electron transport with isolated photosystem I submembrane fraction instead of whole thylakoid membranes.

OXYGEN MEDIATED PHOTOSYSTEM I ACTIVITY IN THYLAKOID MEMBRANES MONITORED BY A PHOTOELECTROCHEMICAL CELL

Abstract— A photoelectrochemical cell has been used to monitor the effects of three enzymes on the photocurrent produced by isolated spinach thylakoids. The enzymes were glucose oxidase, superoxide dismutase and catalase. It is shown that all three inhibit the photocurrent to varying degrees. The results demonstrate that electron transport to the working electrode is mediated by oxygen. Further, the activity monitored originated from photosystem I with oxygen as the acceptor and photosystem II/plastoquinone as the donor. Thus, the photoelectrochemical cell constitutes a potential new approach for the monitoring of pseudocyclic electron transport.

C4 Photosynthesis: Quantum Requirement, C4 and Overcycling and Q-Cycle Involvement

The relationship between overcycling of the C4 acid cycle in C4 photosynthesis (due to CO2 leakage) and the quantum yield of photosynthesis is considered. From a comparison of theoretical and measured quantum yields we suggest that the high efficiency of light utilisation by most C4 plants can only be explained by the mandatory involvement of both the Q-cycle and cyclic or pseudocyclic electron transport in the proton partitioning process. The existence of the Q-cycle mechanism may have been a prerequisite for the evolution of the C4 pathway.