Adjustment of photosynthetic activity to drought and fluctuating light in wheat

Drought is a major cause of losses in crop yield. Under field conditions, plants exposed to drought are usually also experiencing rapid changes in light intensity. Accordingly, plants need to acclimate to both, drought and light stress. Two crucial mechanisms in plant acclimation to changes in light conditions comprise thylakoid protein phosphorylation and dissipation of light energy as heat by non-photochemical quenching (NPQ). Here, we analyzed the acclimation efficacy of two different wheat varieties, by applying fluctuating light for analysis of plants, which had been subjected to a slowly developing drought stress as it usually occurs in the field. This novel approach allowed us to distinguish four drought phases, which are critical for grain yield, and to discover acclimatory responses which are independent of photodamage. In short-term, under fluctuating light, the slowdown of NPQ relaxation adjusts the photosynthetic activity to the reduced metabolic capacity. In long-term, the photosynthetic machinery acquires a drought-specific configuration by changing the PSII-LHCII phosphorylation pattern together with protein stoichiometry. Therefore, the fine-tuning of NPQ relaxation and PSII-LHCII phosphorylation pattern represent promising traits for future crop breeding strategies.     

PGR5 ensures photosynthetic control to safeguard photosystem I under fluctuating light conditions

In a plant’s natural environment, the intensity of light can change rapidly due to sunflecks, cloudiness and intermittent shading. Fluctuations between high and low illumination phases expose the photosynthetic machinery to rapidly changing signals that can be overlapping or contradictory, and accordingly plants have developed astute acclimation strategies to maintain optimal photosynthetic performance in these conditions. Continuous exposure to high light induces an array of protective mechanisms at anatomical, chemical and molecular levels, but when high light phases are short, such as under fluctuating light conditions, the protective strategies that afford protection to constant high light are not employed by plants. One mechanism that is engaged under both constant and fluctuating high light is the photosynthetic control of the Cyt b₆f complex, which prevents hyper-reduction of the electron transfer chain in order to protect PSI from photodamage. The PGR5 protein was recently shown to play an indispensable role in this protective mechanism. This review revisits the findings of earlier studies into photosynthetic control and places PGR5 within the broader context of photoprotection and light acclimation strategies.     

Photosynthetic response to fluctuating environments and photoprotective strategies under abiotic stress

Plants in natural environments must cope with diverse, highly dynamic, and unpredictable conditions. They have mechanisms to enhance the capture of light energy when light intensity is low, but they can also slow down photosynthetic electron transport to prevent the production of reactive oxygen species and consequent damage to the photosynthetic machinery under excess light. Plants need a highly responsive regulatory system to balance the photosynthetic light reactions with downstream metabolism. Various mechanisms of regulation of photosynthetic electron transport under stress have been proposed, however the data have been obtained mainly under environmentally stable and controlled conditions. Thus, our understanding of dynamic modulation of photosynthesis under dramatically fluctuating natural environments remains limited. In this review, first I describe the magnitude of environmental fluctuations under natural conditions. Next, I examine the effects of fluctuations in light intensity, CO₂ concentration, leaf temperature, and relative humidity on dynamic photosynthesis. Finally, I summarize photoprotective strategies that allow plants to maintain the photosynthesis under stressful fluctuating environments. The present work clearly showed that fluctuation in various environmental factors resulted in reductions in photosynthetic rate in a stepwise manner at every environmental fluctuation, leading to the conclusion that fluctuating environments would have a large impact on photosynthesis.     

Photoinactivation of catalase at low temperature and its relevance to photosynthetic and peroxide metabolism in leaves

Abstract In green as well as in etiolated leaves of rye (Secale cereale L. ev. ‘Halo’), exposed to strong light at low temperature (0.4°C) catalase was inactivated. Other heme-containing enzymes (peroxidases) and various enzymes of photosynthetic, photorespiratory or peroxide metabolism were not photoinactivated. After returning plants from a low to a physiological temperature (22°C), catalase activity recovered within 12 h through new synthesis. The leaf contents of H₂O₂ and organic peroxides were not affected by the photoinactivation of catalse. The content of malondialdehyde generally increased after exposure to a higher light intensity. High-light-induced increases of ascorbate, and particularly of glutathione, were more marked in catalase-deficient than in normal leaves. Photoinactivation of catalase was accompanied by severe inhibition of photosynthesis. Photoinhibition of photosynthesis was not related to the lack of catalase because photosynthesis was not impaired when catalase activity was kept low by growing the plants under non-photorespiratory conditions. Photoinhibition appeared to result from photodamage in primary photochemistry of photosystem II, as indicated by a decrease of the maximal variable fluorescence. Photoinhibition of photosynthesis and of catalase have in common that in both instances proteins are involved that are continuously inactivated in light and, therefore, particularly sensitive to stress conditions that prevent their replacement by repair synthesis.     

植物低温光抑制及可能的光保护机制研究进展

综述了近年来有关植物低温光抑制和光保护机制的研究进展。与以往对光抑制的定义不同,现在认为光抑制既包括光对光合作用反应中心的损伤,也包括植物为避免光破坏而形成的生理生化保护机制。该文主要从三个方面展开论述:低温下光抑制发生的原因及光抑制的位点;低温光抑制时可能的光保护机制;低温光抑制下过剩光能的耗散机制。     

喜光榕树和耐荫榕树光适应机制的差异

100%和36%光强下生长的喜光的斜叶榕的光合能力高于耐荫的假斜叶榕,而热耗散能力与之相似,说明强光下斜叶榕主要通过光合作用利用光能和热耗散、假斜叶榕主要通过热耗散防御光破坏.100%光强下生长的两种榕树的日间光抑制程度相似,但叶表光强相同情况下各光强下生长的假斜叶榕的光抑制均比斜叶榕严重.100%光强下假斜叶榕叶片悬挂角大于斜叶榕,导致日间叶表光强低于斜叶榕,这可能是两种榕树日间光抑制程度相似的原因,表明叶片悬挂角的适应变化对假斜叶榕有重要的意义.     

Light and oxygenic photosynthesis: energy dissipation as a protection mechanism against photo-oxidation

Efficient photosynthesis is of fundamental importance for plant survival and fitness. However, in oxygenic photosynthesis, the complex apparatus responsible for the conversion of light into chemical energy is susceptible to photodamage. Oxygenic photosynthetic organisms have therefore evolved several protective mechanisms to deal with light energy. Rapidly inducible non-photochemical quenching (NPQ) is a short-term response by which plants and eukaryotic algae dissipate excitation energy as heat. This review focuses on recent advances in the elucidation of the molecular mechanisms underlying this protective quenching pathway in higher plants.     

快速叶绿素荧光动力学及其在植物抗逆生理研究中的应用

快速叶绿素荧光动力学是研究光合原初反应的无损探针,可监测光合反应的多个事件,并能反应植物的生理状况,可以应用于环境物理及化学条件胁迫对植物的影响(如化肥及除草剂的使用;CO2、O2、O3浓度的变化及温度、光强度胁迫等);也应用于农业生产(如确定耕作方式及日常管理;除草剂、杀虫剂及激素的使用;品种选优等);更是光合生理研究的便捷工具。介绍了快速叶绿素荧光动力学中的JIP-test分析方法,并从光、水分、温度、盐等多个方面介绍了JIP-test方法在植物抗逆生理研究中的应用。     

温州蜜柑叶片光系统反应中心光能分配的变化

为深入了解果树光化学反应中心光能分配的状况,以柑橘为试材,采用调制荧光法对叶片光系统在高光强和低光强下的状态转换进行了研究．结果表明,光系统在100μmol·m^-2·s^-1的低光强下,由于QA的还原使PQ库处于还原状态,导致光能由PSⅡ转向PSⅠ分配,光系统处于状态2;在1000μmol·m^-2·s^-1的高光强下,PQ库无法得到电子而处于氧化状态,导致光能分配由PSⅠ转向PSⅡ,光系统处于状态1,叶片经磷酸酯酶抑制剂NaF处理后,光系统从高光强下状态2到状态1的转换受到抑制,高光强下过多的光能由PSⅠ向PSⅡ分配是导致PSⅡ光破坏的重要原因．     

The phenomenon of photoinhibition of photosynthesis and its importance in reforestation

Photoinhibition, defined as the inhibition of photosynthesis caused by excessive radiance, affects field production to a great extent. This phenomenon is particularly relevant in reforestation practices, when one deals with forests of rapid growth such asEucalyptus. The imposition of additional stress factors during exposure to high radiance increases the potential for photoinhibitory effects, so the inhibition of photosynthesis indicates that the plant is submitted to stressful conditions. Photoinhibition can be reversible, playing a protective role for the photosynthetic systems, but it can also reflect damage that has already occurred in the photosynthetic apparatus, being irreversible in this case. In this review, we present the physiological and molecular mechanisms of photoinhibition and discuss the interaction between light and other stress factors and its effects on plants destined for reforestation. In addition, the present work analyzes some of the features and strategies that help plants avoid or restrict the occurrence of photoinhibition. For instance, pigments and enzymes which naturally occur in plants can prevent photoinhibition, while preadaptation to nonideal conditions can enhance tolerance to a certain stress factor. Most of these morphological, metabolic, and biochemical mechanisms of defense are related to the dissipation of excessive energy such as heat. Understanding these mechanisms can help improve cultivation procedures, avoid the plants’ death, and increase productivity in the field.     

Physiological Mechanisms Underlying Fruit Sunburn

Sunburn is a physiological disorder that can be observed in fruits of several crops growing in areas with warm climates, as a result of photodamage due to an excess of heat and/or light irradiance (visible and ultraviolet light). The main cause is thought to be an increase in reactive oxygen species production which causes oxidative damage due to the incapacity of the fruit to recover from stress. This can result in a characteristic morphological and structural phenotype unacceptable to consumers, leading to severe losses in productivity for farmers. Fruits have a great array of mechanisms to mitigate or reduce reactive oxygen species production and the inactivation of photosynthetic apparatus, such as enhanced xanthophyll cycle-dependent energy dissipation, accumulation of photoprotective pigments and heat-shock proteins, and the biosynthesis of antioxidants, among others. Nevertheless, these mechanisms become inefficient when the stress factors altering the fruit surface exceed a certain threshold (of both duration and intensity). Although this disorder has been studied in detail and previous efforts have provided significant advances in understanding the underlying mechanisms causing sunburn in a number of fruits, further research is still needed. This will undoubtedly provide new approaches and tools for improving current mitigation strategies.     

Adaptation of Grapevine Flowers to Cold Involves Different Mechanisms Depending on Stress Intensity

Grapevine flower development and fruit set are influenced by cold nights in the vineyard. To investigate the impact of cold stress on carbon metabolism in the inflorescence, we exposed the inflorescences of fruiting cuttings to chilling and freezing temperatures overnight and measured fluctuations in photosynthesis and sugar content. Whatever the temperature, after the stress treatment photosynthesis was modified in the inflorescence, but the nature of the alteration depended on the intensity of the cold stress. At 4°C, photosynthesis in the inflorescence was impaired through non-stomatal limitations, whereas at 0°C it was affected through stomatal limitations. A freezing night (−3°C) severely deregulated photosynthesis in the inflorescence, acting primarily on photosystem II. Cold nights also induced accumulation of sugars. Soluble carbohydrates increased in inflorescences exposed to −3°C, 0°C and 4°C, but starch accumulated only in inflorescences of plants treated at 0 and −3°C. These results suggest that inflorescences are able to cope with cold temperatures by adapting their carbohydrate metabolism using mechanisms that are differentially induced according to stress intensity.     

Differences in the physiological state between triticale and maize plants during drought stress and followed rehydration expressed by the leaf gas exchange and spectrofluorimetric methods

The studies were carried out in order to estimate differences in the physiological state between triticale and maize plants subjected to drought stress followed by rehydration. The physiological state of the plants was evaluated by measurements of leaf water potential, net photosynthesis, transpiration and stomatal conductance. Spectrofluorimetric methods for the study of blue, green and red fluorescence were applied. We observed that the soil drought induced a greater water loss in triticale leaves than in maize and consequently caused greater injuries to the photosynthetic apparatus. Moreover, triticale plant recovery was slower than in maize plants during the rehydration phase. The effect was probably connected with the higher functional and structural disorganisation of the photosynthetic apparatus observed during drought stress in triticale. Water stress is responsible for damages to photosystem PS II. The worst light utilisation in photosynthetic light conversion was recorded as an increase in the intensity of red fluorescence. Drought stress induced a strong increase in the intensity of blue and green fluorescence in the studied species and it was still high in maize plants during the first day of rehydration. Increase in the intensity of blue and green fluorescence in maize seems to be the effect of the photoprotection mechanism which prevents damage to PS II through utilisation of excess energy.     

Salt stress and fluctuating light have separate effects on photosynthetic acclimation,but interactively affect biomass

In nature, soil salinity and fluctuating light (FL) often occur concomitantly. However, it is unknown whether salt stress interacts with FL on leaf photosynthesis, architecture, biochemistry, pigmentation, mineral concentrations, as well as whole-plant biomass. To elucidate this, tomato (Solanum lycopersicum) seedlings were grown under constant light (C, 200 μmol m⁻² s⁻¹) or FL (5–650 μmol m⁻² s⁻¹), in combination with no (0 mM NaCl) or moderate (80 mM NaCl) salinity, for 14 days, at identical photoperiods and daily light integrals. FL and salt stress had separate effects on leaf anatomy, biochemistry and photosynthetic capacity: FL reduced leaf thickness as well as nitrogen, chlorophyll and carotenoid contents per unit leaf area, but rarely affected steady-state and dynamic photosynthetic properties along with abundance of key proteins in the electron transport chain. Salt stress, meanwhile, mainly disorganized chloroplast grana stacking, reduced stomatal density, size and aperture as well as photosynthetic capacity. Plant biomass was affected interactively by light regime and salt stress: FL reduced biomass in salt stressed plants by 17%, but it did not affect biomass of non-stressed plants. Our results stress the importance of considering FL when inferring effects of salt-stress on photosynthesis and productivity under fluctuating light intensities.     

Relationship between xanthophyll cycle and non-photochemical quenching in rice (Oryza sativa L.) plants in response to light stress

Thirty days old rice plants grown under low and moderate light conditions were transferred to full sunlight to observe the extent of photoinhibitory damage and protective mechanism, and the relationship between xanthophyll cycle and nonphotochemical quenching (qN) under changing light environment. Control plants (low, moderate and sun grown) exhibited similar Fv/Fm ratio, indicating similar photosynthetic efficiency prior to light stress. On exposure to the high light treatment, low light grown plants exhibited faster and higher degree of photoinhibition compared to moderate and high light grown plants. Moderate and high light grown plants showed relatively less photoinhibition and also showed higher qN, indicating better capacity of energy dissipation. Increase in qN in moderate light and sun grown plants was accompanied by conversion of violaxanthin (V) to antheraxanthin (A) and zeaxanthin (Z) indicating operation of Z-dependent thermal dissipation. Rice plants fed with ascorbate (AsA), a stimulator of the de-epoxidation state of V to Z, showed higher Fv/Fm ratio and qN than the plants fed with dithiothreitol (DTT) an inhibitor of xanthophyll cycle. This indicated that an increased amount of energy reached PS II reaction centre, due to absence of A and Z formation, thereby causing greater damage to photosynthesis in DTT fed rice plants. The present data confirmed the relationship between qN and Z in dissipating the excess light energy, thereby protecting plants against photodamage.     

Behavioural versus physiological photoprotection in epipelic and epipsammic benthic diatoms

Benthic diatoms are dominant primary producers in intertidal marine sediments, which are characterized by widely fluctuating and often extreme light conditions. To cope with sudden increases in light intensity, benthic diatoms display both behavioural and physiological photoprotection mechanisms. Behavioural photoprotection is restricted to raphid pennate diatoms, which possess a raphe system that enables motility and hence positioning in sediment light gradients (e.g. via vertical migration into the sediment). The main physiological photoprotection mechanism is to dissipate excess light energy as heat, measured as Non-Photochemical Quenching (NPQ) of chlorophyll fluorescence. A trade-off between vertical migration and physiological photoprotection (NPQ) in benthic diatoms has been hypothesized before, but this has never been formally tested. We exposed five epipelic diatom species (which move in between sediment particles) and four epipsammic diatom species (which live in close association with individual sand grains) to high light conditions, and characterized both NPQ and the relative magnitude of the migratory response to high light. Our results reveal the absence of a significant downward migratory response in an araphid diatom, but also in several raphid epipsammic diatoms, while all epipelic species showed a significant migratory response upon high light exposure. In all epipsammic species the upregulation of NPQ was rapid and pronounced; NPQ relaxation in low light conditions, however, occurred faster in the araphid diatom, compared with the raphid epipsammic species. In contrast, all epipelic species lacked a strong and flexible NPQ response and showed higher susceptibility to photodamage when not able to migrate. While overall our results support the vertical migration-NPQ trade-off, the lack of strong relationships between the capacity for vertical migration and NPQ within the epipsammic and epipelic groups suggests that other factors as well, such as cell size, substrate type and photoacclimation, may influence photoprotective strategies.     

MPH1 is a thylakoid membrane protein involved in protecting photosystem II from photodamage in land plants

Photosystem II (PSII) is highly susceptible to photoinhibition caused by environmental stimuli such as high light; therefore plants have evolved multifaceted mechanisms to efficiently protect PSII from photodamage. We previously published data suggesting that Maintenance of PSII under High light 1 (MPH1, encoded by AT5G07020), a PSII-associated proline-rich protein found in land plants, participates in the maintenance of normal PSII activity under photoinhibitory stress. Here we provide additional evidence for the role of MPH1 in protecting PSII against photooxidative damage. Two Arabidopsis thaliana mutants lacking a functional MPH1 gene suffer from severe photoinhibition relative to the wild-type plants under high irradiance light. The mph1 mutants exhibit significantly decreased PSII quantum yield and electron transport rate after exposure to photoinhibitory light. The mutants also display drastically elevated photodamage to PSII reaction center proteins after high-light treatment. These data add further evidence that MPH1 is involved in PSII photoprotection in Arabidopsis. MPH1 homologs are found across phylogenetically diverse land plants but are not detected in algae or prokaryotes. Taken together, these results suggest that MPH1 protein began to play a role in protecting PSII against excess light following the transition from aquatic to terrestrial conditions.     

Energy balance of a plant under normal and unfavourable conditions

Two main components of plant energy balance are analyzed--photosynthesis and dark respiration. Different plant species, growing in optimal conditions and differing in photosynthetic metabolism, productivity and potential resistance to stress, have constant ratio between total dark respiration and grossphotosynthesis. The ratio about 38-40% at the phase of active growth. This value reflect plant state, when income (assimilation) is maximized, and expense (total oxidation) is minimized. Intensities of dark respiration of plants in darkness and light are similar despite the fact that the plants have considerable differences: 1) different carbon sources--young assimilates in light and reserve ones in the dark; 2) biochemical modifications (or inhibition) of some stages of dark respiration in light; 3) intensification of alternative respiration in unfavourable conditions differing in dark and light variants. Ratio between intensity of dark respiration and photosynthesis increases in plants growing in unfavourable conditions. This increase is more considerable in plants that are less resistant to the given stress. Characters of energy balance can be used for estimation of physiological status of plants, prediction of resistance and potential productivity at seedling stage.     

The cost of photoinhibition

Photoinhibition is an inevitable consequence of oxygenic photosynthesis. However, the concept of a 'photoinhibition-proof' plant in which photosystem II (PSII) is immune to photodamage is useful as a benchmark for considering the performances of plants with varying mixes of mechanisms which limit the extent of photodamage and which repair photodamage. Some photodamage is bound to occur, and the energy costs of repair are the direct costs of repair plus the photosynthesis foregone during repair. One mechanism permitting partial avoidance of photodamage is restriction of the number of photons incident on the photosynthetic apparatus per unit time, achieved by phototactic movement of motile algae to places with lower incident photosynthetically active radiation (PAR), by phototactic movement of plastids within cells to positions that minimize the incident PAR and by photonastic relative movements of parts of photolithotrophs attached to a substrate. The other means of avoiding photodamage is dissipating excitation of photosynthetic pigments including state transitions, non-photochemical quenching by one of the xanthophyll cycles or some other process and photochemical quenching by increased electron flow through PSII involving CO? and other acceptors, including the engagement of additional electron transport pathways. These mechanisms inevitably have the potential to decrease the rate of growth. As well as the decreased photosynthetic rates as a result of photodamage and the restrictions on photosynthesis imposed by the repair, avoidance, quenching and scavenging mechanisms, there are also additional energy, nitrogen and phosphorus costs of producing and operating repair, avoidance, quenching and scavenging mechanisms. A comparison is also made between the costs of photoinhibition and those of other plant functions impeded by the occurrence of oxygenic photosynthesis, i.e. the competitive inhibition of the carboxylase activity of ribulose bisphosphate carboxylase-oxygenase by oxygen via the oxygenase activity, and oxygen damage to nitrogenase in diazotrophic organisms.     

PROTON GRADIENT REGULATION5 Is Essential for Proper Acclimation of Arabidopsis Photosystem I to Naturally and Artificially Fluctuating Light Conditions

In nature, plants are challenged by constantly changing light conditions. To reveal the molecular mechanisms behind acclimation to sometimes drastic and frequent changes in light intensity, we grew Arabidopsis thaliana under fluctuating light conditions, in which the low light periods were repeatedly interrupted with high light peaks. Such conditions had only marginal effect on photosystem II but induced damage to photosystem I (PSI), the damage being most severe during the early developmental stages. We showed that PROTON GRADIENT REGULATION5 (PGR5)-dependent regulation of electron transfer and proton motive force is crucial for protection of PSI against photodamage, which occurred particularly during the high light phases of fluctuating light cycles. Contrary to PGR5, the NAD(P)H dehydrogenase complex, which mediates cyclic electron flow around PSI, did not contribute to acclimation of the photosynthetic apparatus, particularly PSI, to rapidly changing light intensities. Likewise, the Arabidopsis pgr5 mutant exhibited a significantly higher mortality rate compared with the wild type under outdoor field conditions. This shows not only that regulation of PSI under natural growth conditions is crucial but also the importance of PGR5 in PSI protection.