Light acclimation maintains the redox state of the PS II electron acceptor Q2 within a narrow range over a broad range of light intensities

Chrysanthemum inducum-hybrid `Coral Charm', Hibiscus rosa-sinensis L. `Cairo Red' and Spathiphyllum wallisii Regel `Petit' were grown in natural light in a greenhouse at three levels of irradiance using permanent shade screens. Light acclimation of photosynthesis was characterized using modulated chlorophyll a fluorescence of intact leaves. A close correlation was found between the degree of reduction of the primary electron acceptor Q_A of Photosystem II (PS II) approximated as the fluorescence parameter 1−q_P, and light acclimation. The action range of 1−q_P was 0–0.4 from darkness to full irradiance around noon, within the respective light treatments in the greenhouse, indicating that most PS II reaction centres were kept open. In general, the index for electron transport (ETR) measured by chlorophyll fluorescence was higher for high-light (HL) than intermediate-(IL) and low-light (LL) grown plants. However, HL Chrysanthemum showed 40% higher ETR than HL Hibiscus at light saturation, despite identical redox states of Q_A. The light acclimation of the non-radiative dissipation of excess energy in the antenna, NPQ, varied considerably between the species. However, when normalized against q_P, a strong negative correlation was found between thermal dissipation and ETR measured by chlorophyll fluorescence. To be able to accommodate a high flux of electrons through PS II, the plants with the highest light-saturated ETR had the lowest NPQ/q_P. The possibility of using chlorophyll fluorescence for quantification of the energy balance between energy input and utilization in PS II in intact leaves is discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

光强转换对不同生长环境下桑树叶片光化学效率的影响

以桑树品种‘蒙古桑’为试验材料,利用叶绿素荧光技术研究了光强转换对生长在不同光强下的桑树叶片实际光化学效率(ΦPSⅡ)、电子传递速率(ETR)和非光化学淬灭(NPQ)的影响,分析了非光化学淬灭(NPQ)3个组分的变化.结果表明:当光强从黑暗或弱光转换到自然光条件下,自然光桑树叶片的光量子转化效率高于弱光叶片,ΦPSⅡ、ETR诱导平衡较快,NPQ诱导呈先升后降趋势.自然光叶片在强光下状态转换淬灭组分(qT)占NPQ的18%,而弱光叶片qT仅占NPQ的7%.与弱光桑树叶片相比,自然光桑树叶片可以通过较高的光量子转化效率和较强的调节激发能在PSⅠ和PSⅡ之间的分配能力来适应光强的变化.     

田间条件下棉花幼叶光合特性及光保护机制

通过比较棉花(Gossypium hirsutum)幼叶和完全展开叶气体交换参数及叶绿素荧光特性的差异, 探讨高光强下幼叶的光抑制程度及明确光保护机制间的协调机理。在田间自然条件下, 以棉花刚展平的幼嫩叶片(幼叶)和面积已达到最大的完全展开叶片为研究对象, 通过测定不同发育阶段叶片气体交换参数及叶绿素a荧光参数的变化, 并运用Dual-PAM100对不同发育阶段的叶片进行快速光响应曲线的拟合。结果表明: 幼叶和完全展开叶片在光合、荧光特性方面表现出明显的差异。与完全展开叶相比, 较低的叶绿素(Chl)含量和气孔导度(G_s)是幼叶较低净光合速率(P_n)的限制因素, 从而直接导致其光系统II (PSII)实际光化学效率(Φ_PSII)和光化学猝灭系数(q_P)的降低。在1800 μmol·m^-2·s^-1光强以下, 完全展开叶具有较强的围绕PSI循环的电子流(CEF), 有利于合成ATP, 是其具有较高光合能力的原因之一。相同光强下, 幼叶较低的光饱和点(LSP)更易受光抑制, 但其PSII原初光化学效率(F_v/F_m)的日变化幅度显著小于完全展开叶, 说明强光下幼叶通过类胡萝卜素(Car)猝灭单线态氧、光呼吸(P_r)、热耗散(NPQ)以及PSI-CEF等光保护机制能有效地耗散过剩的光能, 从而避免其光合机构发生光抑制。     

On the relationship between the quantum yield of Photosystem II electron transport,as determined by chlorophyll fluorescence and the quantum yield of CO2-dependent O2 evolution

We tested the two empirical models of the relationship between chlorophyll fluorescence and photosynthesis, previously published by Weis E and Berry JA 1987 (Biochim Biophys Acta 894: 198–208) and Genty B et al. 1989 (Biochim Biophys Acta 990: 87–92). These were applied to data from different species representing different states of light acclimation, to species with C₃ or C₄ photosynthesis, and to wild-type and a chlorophyll b-less chlorina mutant of barley. Photosynthesis measured as CO₂-saturated O₂ evolution and modulated fluorescence were simultaneously monitored over a range of photon flux densities. The quantum yields of O₂ evolution (ØO₂) were based on absorbed photons, and the fluorescence parameters for photochemical (q_p) and non-photochemical (q_N) quenching, as well as the ratio of variable fluorescence to maximum fluorescence during steady-state illumination (F'_v/F'_m), were determined. In accordance with the Weis and Berry model, most plants studied exhibited an approximately linear relationship between ØO₂/q_p (i.e., the yield of O₂ evolution by open Photosystem II reaction centres) and q_N, except for wild-type barley that showed a non-linear relationship. In contrast to the linear relationship reported by Genty et al. for q_p×F'_v/F'_m (i.e., the quantum yield of Photosystem II electron transport) and ØCO₂, we found a non-linear relationship between q_p×F'_v/F'_m and ØO₂ for all plants, except for the chlorina mutant of barley, which showed a largely linear relationship. The curvilinearity of wild-type barley deviated somewhat from that of other species tested. The non-linear part of the relationship was confined to low, limiting photon flux densities, whereas at higher light levels the relationship was linear. Photoinhibition did not change the overall shape of the relationship between q_p×F'_v/F'_m and ØO₂ except that the maximum values of the quantum yields of Photosystem II electron transport and photosynthetic O₂ evolution decreased in proportion to the degree of photoinhibition. This implies that the quantum yield of Photosystem II electron transport under high light conditions may be similar for photoinhibited and non-inhibited plants. Based on our experimental results and theoretical analyses of photochemical and non-photochemical fluoresce quenching processes, we conclude that both models, although not universal for all plants, provide useful means for the prediction of photosynthesis from fluorescence parameters. However, we also discuss that conditions which alter one or more of the rate constants that determine the various fluorescence parameters, as well as differential light penetration in assays for oxygen evolution and fluorescence emission, may have direct effect on the relationships of the two models.Abbreviations F₀ and F'₀ fluorescence when all Photosystem II reaction centres are open in dark- and light-acclimated leaves, respectively - F_m and F'_m fluorescence when all Photosystem II reaction centres are closed in dark and light, respectively - F_v variable fluorescence equal to F_m-F₀ - F_s steady state level of fluorescence in light - F'_v and F'_m variable (F'_m-F'₀) and maximum fluorescence under steady state light conditions - HEPES N-2-hydroxyethylpiperazine-N-2-ethane-sulphonic acid - Q_A the primary, stabile quinone acceptor of Photosystem II - q_N non-photochemical quenching of fluorescence - q_p photochemical quenching of fluorescence - ØO₂ quantum yield of CO₂-saturated O₂ evolution based on absorbed photons     

SINGLE‐CELL PULSE AMPLITUDE MODULATION FLUORESCENCE MEASUREMENTS OF THE GIANT DIATOM ETHMODISCUS (BACILLARIOPHYCEAE)1

Diurnal changes in effective yield (ΔF:F_m′), rapid light curves (RLCs), and induction/dark recovery time series were measured on individual cells of the giant diatom Ethmodiscus Castracane using active fluorescence (pulse amplitude modulation fluorometry). Unlike the co‐occurring diatom Hemiaulus and bulk phytoplankton, there was no observable diurnal down‐regulation of yield or relative electron transport rates in Ethmodiscus. Yields were constant at or near maximum values (0.7–0.8). Increases in ΔF:F_m′ during the initial actinic levels are consistent with dark nonphotochemical quenching mechanisms. Sustained actinic illumination (660 μmol photon·m^?2·s^?1) resulted in a ΔF:F_m′ of 0.2–0.3, but rapid recovery to near‐maximum values occurred in subsequent dark periods. Such recovery occurred even after exposure to full sunlight for 28 min, but not at 60 min. Thus, the lack of diurnal down‐regulation in Ethmodiscus is apparent, not real, and is an artifact of the time scale of sample extraction from net tows. These positively buoyant cells showed no evidence of routine photodamage, probably due to mixing and reduction in the average light exposure. The general patterns seen in RLCs from light‐and dark‐adapted higher plants are significantly different from those observed in Ethmodiscus. These results suggest that active fluorescence characteristics require careful examination to differentiate habitat‐ and taxon‐specific characteristics from light‐history effects. It is unclear whether the rapid recovery seen in Ethmodiscus is unique. The differences seen between Hemiaulus and Ethmodiscus from the same samples suggest that changes in community yield values measured in countertop systems could be the result of species replacement in addition to experimental or environmental perturbations.     

Reversible photoinhibition of unhardened and cold-acclimated spinach leaves at chilling temperatures

The photoinhibition of photosynthesis at chilling temperatures was investigated in cold-acclimated and unhardened (acclimated to +18° C) spinach (Spinacia oleracea L.) leaves. In unhardened leaves, reversible photoinhibition caused by exposure to moderate light at +4° C was based on reduced activity of photosystem (PS) II. This is shown by determination of quantum yield and capacity of electron transport in thylakoids isolated subsequent to photoinhibition and recovery treatments. The activity of PSII declined to approximately the same extent as the quantum yield of photosynthesis of photoinhibited leaves whereas PSI activity was only marginally affected. Leaves from plants acclimated to cold either in the field or in a growth chamber (+1° C), were considerably less susceptible to the light treatment. Only relatively high light levels led to photoinhibition, characterized by quenching of variable chlorophyll a fluorescence (F_V) and slight inhibition of PSII-driven electron transport. Fluorescence data obtained at 77 K indicated that the photoinhibition of cold-acclimated leaves (like that of the unhardened ones) was related to increased thermal energy dissipation. But in contrast to the unhardened leaves, 77 K fluorescence of cold-acclimated leaves did not reveal a relative increase of PSI excitation. High-light-treated, cold-acclimated leaves showed increased rates of dark respiration and a higher light compensation point. The photoinhibitory fluorescence quenching was fully reversible in low light levels both at +18° C and +4° C; the recovery was much faster than in unhardened leaves. Reversible photoinhibition is discussed as a protective mechanism against excess light based on transformation of PSII reaction centers to fluorescence quenchers.Abbreviations F_O initial fluorescence - F_M maximal fluorescence - F_V devariable fluorescence (f_m-f_o) - PFD photon flux density - PS photosystem - SD standard deviation The authors thank the Deutsche Forschungsgemeinschaft and the Academy of Finland for financial support.     

Light Scattering as an Indicator of the Energy State in Leaves of the Crassulacean Acid Metabolism Plant Kalanchoë pinnata

Both transmittance changes in a weak beam of green light (light scattering) and the slow decay of chlorophyll a fluorescence were used as indicators of the energy state of leaves of a Crassulacean acid metabolism plant, Kalanchoë pinnata, at frequent intervals during 12-hour light/12-hour dark cycles. To induce light scattering and fluorescence changes, leaves were exposed to red light for 6 minutes. When measurements were made during the light period, the leaves were kept in darkness for 6 minutes before illumination. In the middle of the light period, when malic acid decarboxylation was very active and stomatal conductance was low, light scattering changes were small and indicated that the energy state of leaves was low. This result was supported by determination of adenylate levels. Light scattering and ATP/ADP ratios increased during the late light period when the tissue was deacidified. Illumination produced maximum light scattering changes between the 2nd and 5th hour of the dark period, when rates of dark CO₂ fixation were highest. Light scattering and fluorescence measurements taken from leaves, which were illuminated with red or far-red light in the presence or absence of O₂ showed that, in addition to linear electron transport, K. pinnata has the potential for both cyclic and pseudocyclic electron transport. The results are relevant with regard to the high ATP demand during Crassulacean acid metabolism.     

Chilling-Induced Photoinhibition in Two Oak Species: Are Evergreen Leaves Inherently Better Protected Than Deciduous Leaves?

We compared the sensitivity to cold stress, in terms of photosynthetic capacity and changes in chlorophyll fluorescence of photosystem 2 (PS2), of an evergreen and a deciduous oak species, which co-occur in the southeastern United States. We predicted that the evergreen species, Quercus virginiana, which must endure winter, is likely to have an inherently greater capacity for energy dissipation and to be less susceptible to chilling stress than the deciduous species, Quercus michauxii. Short-term cold stress in both species lead to greater than 50 % reduction in maximum photosynthetic rates, 60-70 % reduction in electron transport, and irreversible quenching of PS2 fluorescence. The kinetics of recovery in the dark after exposure to 1 h high irradiance (1000 μmol m^-2 s^-1) and chilling (5 °C) showed that the evergreen Q. virginiana exhibited more protective q_E and less irreversible quenching (q_I) than the deciduous Q. michauxii. The large q_E which we observed in Q. virginiana suggests that the capacity for photoprotection at low temperatures is not induced by a long-term acclimation to cold but preexists in evergreen leaves. This capacity may contribute to the ability of this species to maintain leaves during the winter. This revised version was published online in June 2006 with corrections to the Cover Date.     

Chilling-Induced Photoinhibition in Two Oak Species: Are Evergreen Leaves Inherently Better Protected Than Deciduous Leaves?

We compared the sensitivity to cold stress, in terms of photosynthetic capacity and changes in chlorophyll fluorescence of photosystem 2 (PS2), of an evergreen and a deciduous oak species, which co-occur in the southeastern United States. We predicted that the evergreen species, Quercus virginiana, which must endure winter, is likely to have an inherently greater capacity for energy dissipation and to be less susceptible to chilling stress than the deciduous species, Quercus michauxii. Short-term cold stress in both species lead to greater than 50 % reduction in maximum photosynthetic rates, 60-70 % reduction in electron transport, and irreversible quenching of PS2 fluorescence. The kinetics of recovery in the dark after exposure to 1 h high irradiance (1000 µmol m^-2 s^-1) and chilling (5 °C) showed that the evergreen Q. virginiana exhibited more protective q_E and less irreversible quenching (q_I) than the deciduous Q. michauxii. The large q_E which we observed in Q. virginiana suggests that the capacity for photoprotection at low temperatures is not induced by a long-term acclimation to cold but preexists in evergreen leaves. This capacity may contribute to the ability of this species to maintain leaves during the winter.     

NPQ(T): a chlorophyll fluorescence parameter for rapid estimation and imaging of non‐photochemical quenching of excitons in photosystem‐II‐associated antenna complexes

In photosynthesis, light energy is absorbed by light‐harvesting complexes and used to drive photochemistry. However, a fraction of absorbed light is lost to non‐photochemical quenching (NPQ) that reflects several important photosynthetic processes to dissipate excess energy. Currently, estimates of NPQ and its individual components (q_E, q_I, q_Z and q_T) are measured from pulse‐amplitude‐modulation (PAM) measurements of chlorophyll fluorescence yield and require measurements of the maximal yield of fluorescence in fully dark‐adapted material (F_m), when NPQ is assumed to be negligible. Unfortunately, this approach requires extensive dark acclimation, often precluding widespread or high‐throughput use, particularly under field conditions or in imaging applications, while introducing artefacts when F_m is measured in the presence of residual photodamaged centres. To address these limitations, we derived and characterized a new set of parameters, NPQ_(T), and its components that can be (1) measured in a few seconds, allowing for high‐throughput and field applications; (2) does not require full relaxation of quenching processes and thus can be applied to photoinhibited materials; (3) can distinguish between NPQ and chloroplast movements; and (4) can be used to image NPQ in plants with large leaf movements. We discuss the applications benefits and caveats of both approaches.     

Rapid light-response curves of chlorophyll fluorescence in microalgae: relationship to steady-state light curves and non-photochemical quenching in benthic diatom-dominated assemblages

Rapid light-response curves (RLC) of variable chlorophyll fluorescence were measured on estuarine benthic microalgae with the purpose of characterising its response to changes in ambient light, and of investigating the relationship to steady-state light-response curves (LC). The response of RLCs to changes in ambient light (E, defined as the irradiance level to which a sample is acclimated to prior to the start of the RLC) was characterised by constructing light-response curves for the RLC parameters α _RLC, the initial slope, ETR_m,RLC, the maximum relative electron transport rate, and E _k,RLC, the light-saturation parameter. Measurements were carried out on diatom-dominated suspensions of benthic microalgae and RLC and LC parameters were compared for a wide range of ambient light conditions, time of day, season and sample taxonomic composition. The photoresponse of RLC parameters was typically bi-phasic, consisting of an initial increase of all parameters under low ambient light (E < 21–181 μmol m⁻² s⁻¹), and of a phase during which α _RLC decreased significantly with E, and the increase of ETR_m,RLC and E _k,RLC was attenuated. The relationship between RLC and LC parameters was dependent on ambient irradiance, with significant correlations being found between α _RLC and α, and between ETR_m,RLC and ETR_m, for samples acclimated to low and to high ambient irradiances, respectively. The decline of α _RLC under high light (Δα _RLC) was strongly correlated (P < 0.001 in all cases) with the level of non-photochemical quenching (NPQ) measured before each RLC. These results indicate the possibility of using RLCs to characterise the steady-state photoacclimation status of a sample, by estimating the LC parameter E _k, and to trace short-term changes in NPQ levels without dark incubation.     

Changes in the distribution of light energy between the two photosystems in spinach leaves

Time courses of chlorophyll fluorescence at room temperature and fluorescence spectra at 77 K were measured to investigate the light-induced changes in the distribution of light energy between the two photosy stems in young spinach leaves. Illumination of the dark adapted leaves with primarily system II light induced typical fluorescence transients at room temperature. Fluorescence spectra at 77 K showed that the intensity of system II fluorescence at 77 K changed nearly in parallel with the fluorescence transients at room temperature within the range from M₁ to T during illumination of the leaf. Illumination of the dark adapted leaves with light I produced an increase of system II fluorescence measured at 77 K. The characteristics of the changes induced by light I or II were different, showing that these two effects are related to different mechanisms. These results suggest that the dark state in spinach leaves is state II, that light I induces a state II to I transition, while light II induces fluorescence changes that are produced by mechanisms other than state I-state II transitions.     

The carotenoid zeaxanthin and ‘high-energy-state quenching’ of chlorophyll fluorescence

The possibility that zeaxanthin mediates the dissipation of an excess of excitation energy in the antenna chlorophyll of the photochemical apparatus has been tested through the use of an inhibitor of violaxanthin de-epoxidation, dithiothreitol (DTT), as well as through the comparison of two closely related organisms (green and blue-green algal lichens), one of which (blue-green algal lichen) naturally lacks the xanthophyll cycle. In spinach leaves, DTT inhibited a major component of the rapidly relaxing high-energy-state quenching' of chlorophyll fluorescence, which was associated with a quenching of the level of initial fluorescence (F₀) and exhibited a close correlation with the zeaxanthin content of leaves when fluorescence quenching was expressed as the rate constant for radiationless energy dissipation in the antenna chlorophyll. Green algal lichens, which possess the xanthophyll cycle, exhibited the same type of fluorescence quenching as that observed in leaves. Two groups of blue-green algal lichens were used for a comparison with these green algal lichens. A group of zeaxanthin-free blue-green algal lichens did not exhibit the type of chlorophyll fluorescence quenching indicative of energy dissipation in the pigment bed. In contrast, a group of blue-green algal lichens which had formed zeaxanthin slowly through reactions other than the xanthophyll cycle, did show a very similar response to that of leaves and green algal lichens. Fluorescence quenching indicative of radiationless energy dissipation in the antenna chlorophyll was the predominant component of high-energy-state quenching in spinach leaves under conditions allowing for high rates of steady-state photosynthesis. A second, but distinctly different type of high-energy-state quenching of chlorophyll fluorescence, which was not inhibited by DTT (i.e., it was zeaxanthin independent) and which is possibly associated with the photosystem II reaction center, occurred in addition to that associated with zeaxanthin in leaves under a range of conditions which were less favorable for linear photosynthetic electron flow. In intact chloroplasts isolated from (zeaxanthin-free) spinach leaves a combination of these two types of rapidly reversible fluorescence quenching occurred under all conditions examined.Abbreviations DTT dithiothreitol - F₀ (or F₀) yield of instantaneous fluorescence at open PS II reaction centers in the dark (or during actinic illumination) - F_M (or F_M) yield of maximum fluorescence induced by a saturation pulse of light in the dark (or during actinic illumination) - F_V (or F_V) yield of variable fluorescence induced by a saturating pulse of light in the dark (or during actinic illumination) - k _D rate constant for radiationless energy dissipation in the antenna chlorophyll - SV Stern-Volmer equation - PFD photon flux density - PS I photosystem I - PS II photosystem II - Q_A acceptor of photosystem II - q_N coefficient of nonphotochemical chlorophyll fluorescence quenching - q_P coefficient of photochemical chlorophyll fluorescence quenching     

Strategies providing success in a variable habitat: II. Ecophysiology of photosynthesis of Cladophora glomerata

Cladophora glomerata (L.) Kütz. is the dominant filamentous algae of the river Ilm, Thuringia, Germany. For most of the year it can be found at open as well as at shaded sites. Photosynthetic acclimation of C. glomerata to different light intensities was detected by chlorophyll fluorescence measurements and pigment analysis. Cladophora glomerata from highlight sites showed decreased values of efficiency of open photosystem II (F_v/F_m) as compared with C. glomerata from low‐light sites. Winter populations revealed higher F_v/F_m values than summer populations. A light‐induced decrease in efficiency of the closed photosystem II was observed at increasing irradiance intensities. The decrease was higher in C. glomerata from shaded sites compared with plants from open sites. Differences in the photosynthetic electron transport rate of different populations of C. glomerata were shown by photosynthesis–irradiance curves. Summer populations from high‐light sites yielded higher maximum electron transport rates than plants from low‐light sites, whereas winter populations exhibited significantly decreased values compared with the summer populations. Results of the analysis of photosynthetic pigments corresponded with data from chlorophyll fluorescence measurements. In addition to these long‐term acclimation effects, C. glomerata expressed its ability to cope with rapid changes in the light environment by the de‐epoxidation of violaxanthin during exposure to high light intensities.     

Diurnal changes in chlorophyll fluorescence and light utilization in Colocasia esculenta leaves grown in marshy waterlogged area

Diurnal cycle of chlorophyll fluorescence parameters was done in Colocasia esculenta L. (swamp taro) grown in marshy land under sun or under shade. The sun leaves maintained higher electron transport rate (ETR) and steady state to initial fluorescence ratio (F_s/F₀) than shade leaves. In spite of lower ETR, higher photochemical quenching (PQ), and effective quantum yield of photosystem 2 (Φ_PS2) was evident in shade plants compared to plants exposed to higher irradiance. ETR increased linearly with increase in irradiance more under low irradiance (r ² = 0.84) compared to higher irradiance (r ² = 0.62). The maximum quantum yield of PS 2 (F_v/F_m) did not differ much in sun and shade leaves with the exception of midday when excess of light energy absorbed by plants under sun was thermally dissipated. Hence swamp taro plants adopted different strategies to utilize radiation under different irradiances. At higher irradiance, there was faster decline in proportion of open PS 2 centers (PQ) and excess light energy was dissipated through non-photochemical quenching (NPQ). Under shade, absorbed energy was effectively utilized resulting in higher Φ_PS2.     

Photorespiration is Essential for the Protection of the Photosynthetic Apparatus of C3 Plants Against Photoinactivation Under Sunlight

CO₂ assimilation, transpiration and modulated chlorophyll fluorescence of leaves of Chenopodium bonus-henricus (L.) were measured in the laboratory and, at a high altitude location, in the field. Direct calibration of chlorophyll fluorescence parameters against carbon assimilation in the presence of 1 or 0.5% oxygen (plus CO₂) proved necessary to calculate electron transport under photorespiratory conditions in individual experiments. Even when stomata were open in the field, total electron transport was two to three times higher in sunlight than indicated by net carbon gain. It decreased when stomata were blocked by submerging leaves under water or by forcing them to close in air by cutting the petiole. Even under these conditions, electron transport behind closed stomata approached 10 nmol electrons m^?2 leaf area s^?1 at temperatures between 25 and 30 °C. No photoinactivation of photosystem II was indicated by fluorescence analysis after a day's exposure to full sunlight. Only when leaves were submerged in ice was appreciable photoinactivation noticeable after 4 h exposure to sunlight. Even then almost full recovery occurred overnight. Electron transport behind blocked stomata was much decreased when leaves were darkened for 70 min (in order to deactivate light-regulated enzymes of the Calvin cycle) before exposure to full sunlight. Brief exposure of leaves to HCN (to inhibit photoassimilation and photorespiration) also decreased electron transport drastically compared to electron transport in unpoisoned leaves with blocked stomata. Non-photochemical fluorescence quenching and reduction of Q_A, the primary electron acceptor of photosystem II was increased by HCN-poisoning. Very similar observations were made when glyceraldehyde was used instead of HCN to inhibit photosynthesis and photorespiration. In HCN-poisoned leaves, residual electron transport increased linearly with temperature and showed early light saturation revealing characteristics of the Mehler reaction. During short exposure of these leaves to photon flux densities equivalent to 25% of sunlight, no or only little photoinactivation of photosystem II was observed. However, prolonged exposure to sunlight caused inactivation even though non-photochemical quenching of chlorophyll fluorescence was extensive. Simultaneously, oxidation of cellular ascorbate and glutathione increased. Inactivation of photosystem II was reversible in dim light and in the dark only after short times of exposure to sunlight. Glyceraldehyde was very similar to HCN in increasing the sensitivity of photosystem II in leaves to sunlight. We conclude from the observations that the electron transport permitted by the interplay of photoassimilatory and photorespiratory electron transport is essential to prevent the photoinactivation of photosynthetic electron transport. The Mehler and Asada reactions, which give rise to strong nonphotochemical fluorescence quenching, are insufficient to protect the chloroplast electron transport chain against photoinactivation.     

Photo-inhibition and seasonal photosynthetic performance of the seaweed Laminaria saccharina during a simulated tidal cycle: chlorophyll fluorescence measurements and pigment analysis

Laminaria saccharina (Lamouroux) form the largest, most abundant and conspicuous seaweed populations along the French coast of the eastern English Channel. As they are located in the intertidal zone, they are exposed to considerable irradiance variations, mainly related to tidal cycles. The response of these macro‐algae to light variations over a simulated daily tidal cycle was investigated in the laboratory during spring, autumn and winter using chlorophyll fluorescence and pigment analysis. The maximum quantum yield of photosystem II (PSII) photochemistry (F_v/F_m) and the operating PSII efficiency (Φ_PSII) showed clear daily cycles according to the irradiance variation throughout the 12 h simulated tidal cycle, whereas the pattern of the relative photosynthetic electron transport rate (rETR) was not so obvious. The algae reacted to the light increase by developing photoprotective mechanisms able to dissipate the excess energy reaching PSII by the de‐epoxidation of violaxanthin into zeaxanthin. Because of their better acclimation to strong irradiance, spring populations were less affected by this light treatment than were winter populations. In particular, L. saccharina showed more pigments of the xanthophyll cycle in spring to cope with strong irradiance exposure. Alternatively, they developed their antenna complexes in winter to harvest a maximum of light.     

Use of simultaneous analysis of gas-exchange and chlorophyll fluorescence quenching for analysing the effects of water stress on photosynthesis in apple leaves

Summary A convenient system for the rapid simultaneous measurement of both chlorophyll fluorescence quenching using a modulated light system, and of CO₂, and water vapour exchange by leaves is described. The system was used in a study of the effects of water deficits on the photosynthesis by apple leaves (Malus x domestica Borkh.). Apple leaves were found to have low values of steady-state variable fluorescence, and the existence of significant fluorescence with open traps (F_o) quenching necessitated the measurement and use of a corrected F_o in the calculation of quenching components. Long-term water stress had a marked effect on both gas-exchange and chlorophyll fluorescence quenching. Non-photochemical quenching (qn) in particular was increased in water-stressed leaves, and it was particularly sensitive to incident radiation in such leaves. In contrast, rapid dehydration only affected gas exchange. Relaxation of qn quenching in the dark was slow, taking approximately 10 min for a 50% recovery, in well-watered and in draughted plants, and whether or not the plants had been exposed to high light.     

THE SHORT‐TERM EFFECT OF IRRADIANCE ON THE PHOTOSYNTHETIC PROPERTIES OF ANTARCTIC FAST‐ICE MICROALGAL COMMUNITIES1

Although sea‐ice represents a harsh physicochemical environment with steep gradients in temperature, light, and salinity, diverse microbial communities are present within the ice matrix. We describe here the photosynthetic responses of sea‐ice microalgae to varying irradiances. Rapid light curves (RLCs) were generated using pulse amplitude fluorometry and used to derive photosynthetic yield (Φ_PSII), photosynthetic efficiency (α), and the irradiance (E_k) at which relative electron transport rate (rETR) saturates. Surface brine algae from near the surface and bottom‐ice algae were exposed to a range of irradiances from 7 to 262 μmol photons · m^?2 · s^?1. In surface brine algae, Φ_PSII and α remained constant at all irradiances, and rETR_max peaked at 151 μmol photons · m^?2 · s^?1, indicating these algae are well acclimated to the irradiances to which they are normally exposed. In contrast, Φ_PSII, α, and rETR_max in bottom‐ice algae reduced when exposed to irradiances >26 μmol photons · m^?2 · s^?1, indicating a high degree of shade acclimation. In addition, the previous light history had no significant effect on the photosynthetic capacity of bottom‐ice algae whether cells were gradually exposed to target irradiances over a 12 h period or were exposed immediately (light shocked). These findings indicate that bottom‐ice algae are photoinhibited in a dose‐dependent manner, while surface brine algae tolerate higher irradiances. Our study shows that sea‐ice algae are able to adjust to changes in irradiance rapidly, and this ability to acclimate may facilitate survival and subsequent long‐term acclimation to the postmelt light regime of the Southern Ocean.     

Studies on the Mechanism of Photoinhibition in Higher Plants: I. EFFECTS OF HIGH LIGHT INTENSITY ON CHLOROPLAST ACTIVITIES IN CUCUMBER ADAPTED TO LOW LIGHT

Cucumber plants (Cucumis sativus L.), grown at low quantum flux density (120-150 microeinsteins per square meter per second) were photoinhibited by a three-hour exposure in air to ten times the light intensity experienced during growth. Chloroplasts were isolated from photoinhibited and control leaves and the following activities determined: O₂ evolution in the presence of ferricyanide, photosystem I activity, noncyclic and cyclic photophosphorylation, and light-induced proton uptake. Chlorophyll and chloroplast absorbance spectra, and chloroplast fluorescence were also measured. It was found that photosystem II electron transport and non-cyclic photophosphorylation were inhibited by about 50%, while cyclic photophosphorylation was less inhibited and photosystem I electron transport and light-induced proton uptake were unaffected. Electron transport to methylviologen could not be fully restored by electron donation to photosystem II. Chloroplast fluorescence induction at room temperature was strongly reduced following photoinhibition. There was no difference in the absorption spectra of the extracted chlorophylls from control and photoinhibited chloroplasts, but an increase of the absorption in the blue wavelength region was observed in the photoinhibited chloroplasts. It is suggested that high light stress does not result in alteration of the membrane properties, as is the case in low-temperature stress for example, but affects directly the photosynthetic reaction centers, primarily of photosystem II.