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This research first verified that prokaryotic algae are more sensitive to toxicity of CuO nanoparticles (CuO NPs) than eukaryotic algae and that CuO NPs damaged photosynthesis of prokaryotic algae (Arthrospira sp.) but had no effect on respiration. The Cu2+ released by CuO NPs caused a bending deformation of the thylakoid, which was an important cause of the decline in photosynthetic capacity. In addition, the D1 protein was the most susceptible site to CuO NPs. The degradation of D1 protein reduced photosynthetic electron transport, which enhanced the excess excitation energy to cause the accumulation of reactive oxygen species (ROS) to further result in oxidative stress on algae. Dissolved organic matter (DOM) increased the toxicity of CuO NPs to photosynthesis of Arthrospira sp. The damage of photosynthesis caused by CuO NPs is an important reason why CuO NPs have a serious toxicity to algae.

Whether photosynthesis has improved with increasing yield in major crops remains controversial. Research in this area has often neglected to account for differences in light intensity experienced by cultivars released in different years. Light intensity is expected to be positively associated with photosynthetic capacity and the resistance of the photosynthetic apparatus to high light but negatively associated with light-utilization efficiency under low light. Here, we analyzed the light environment, photosynthetic activity, and protein components of leaves of 26 winter wheat cultivars released during the past 60 years in China. Over time, light levels on flag leaves significantly decreased due to architectural changes, but photosynthetic rates under high or low light and the resistance of the photosynthetic apparatus to high light remained steady, contrary to expectations. We propose that the difference between the actual and expected trends is due to breeding. Specifically, breeding has optimized photosynthetic performance under high light rather than low light. Moreover, breeding selectivity altered the stoichiometry of several proteins related to dynamic photosynthesis, canopy light distribution, and photoprotection. These results indicate that breeding has significantly altered the photosynthetic mechanism in wheat and its response to the light environment. These changes likely have helped increase wheat yields.

Reactivation of photosynthetic carbon assimilation during high light after a low light interval is slower in C4 than in C3 leaves.

This study primarily explored the limiting factor for photosynthesis during the development of Euonymus japonicus leaves. The analysis of the chlorophyll fluorescence transient, pulse-modulated fluorescence, 820-nm reflection, and expression of core proteins for photosystems demonstrated that photosystem II (PSII) activity developed more rapidly than did photosystem I (PSI) activity. The slower development of the PSI activity restricted linear and cyclic electron transport and thus inhibited the production of ATP and NADPH, which inhibits the activation of Rubisco, resulting in low activity of carboxylation efficiency. The application of exogenous NADPH (50 μM) and ATP (100 μM) to leaves remarkably increased the Pn and CE in the youngest leaf but not in the fully expanded leaf, which indicated that an inadequate supply of the assimilatory power significantly inhibited CE and Pn. We concluded that the slower development of the PSI activity was one of the most important limiting factors for photosynthesis during the development of E. japonicus leaves.

The mitochondrial alternative pathway (AP) represents an important photoprotective mechanism for the chloroplast, but the temperature sensitivity of its photoprotective role is unknown. In this study, using the aox1a Arabidopsis mutant, the photoprotective role of the AP was verified under various temperatures, and the mechanism underlying the temperature sensitivity of the AP's photoprotective role was clarified. It was observed that the photoprotective role of the AP increased with rising temperature but was absent at low temperature. The photoprotective role of the AP was severely reduced under non-photorespiratory conditions. Disturbance of the AP inhibited the conversion of glycine to serine in mitochondria, which may restrain upstream photorespiratory metabolism and aggravate photoinhibition. With rising temperatures, photorespiration accelerated and the restraint of photorespiration caused by disturbance of the AP also increased, determining the temperature sensitivity of the AP's photoprotective role. We also verified that not only the AP but also the cytochrome pathway in mitochondria contributes to photoprotection by maintaining photorespiration.

Background Glycine soja is a halophytic soybean native to saline soil in Yellow River Delta, China. Photosystem I (PSI) performance and the interaction between photosystem II (PSII) and PSI remain unclear in Glycine soja under salt stress. This study aimed to explore salt adaptability in Glycine soja in terms of photosystems coordination. Results Potted Glycine soja was exposed to 300 mM NaCl for 9 days with a cultivated soybean, Glycine max, as control. Under salt stress, the maximal photochemical efficiency of PSII (Fv/Fm) and PSI (△MR/MR0) were significantly decreased with the loss of PSI and PSII reaction center proteins in Glycine max, and greater PSI vulnerability was suggested by earlier decrease in △MR/MR0 than Fv/Fm and depressed PSI oxidation in modulated 820 nm reflection transients. Inversely, PSI stability was defined in Glycine soja, as △MR/MR0 and PSI reaction center protein abundance were not affected by salt stress. Consistently, chloroplast ultrastructure and leaf lipid peroxidation were not affected in Glycine soja under salt stress. Inhibition on electron flow at PSII acceptor side helped protect PSI by restricting electron flow to PSI and seemed as a positive response in Glycine soja due to its rapid recovery after salt stress. Reciprocally, PSI stability aided in preventing PSII photoinhibition, as the simulated feedback inhibition by PSI inactivation induced great decrease in Fv/Fm under salt stress. In contrast, PSI inactivation elevated PSII excitation pressure through inhibition on PSII acceptor side and accelerated PSII photoinhibition in Glycine max, according to the positive and negative correlation of △MR/MR0 with efficiency that an electron moves beyond primary quinone and PSII excitation pressure respectively. Conclusion Therefore, photosystems coordination depending on PSI stability and rapid response of PSII acceptor side contributed to defending salt-induced oxidative stress on photosynthetic apparatus in Glycine soja. Photosystems interaction should be considered as one of the salt adaptable mechanisms in this halophytic soybean.

Background Plants are always exposed to dynamic light. The photosynthetic light use efficiency of leaves is lower in dynamic light than in uniform irradiance. Research on the influence of environmental factors on dynamic photosynthesis is very limited. Nitrogen is critical for plants, especially for photosynthesis. Low nitrogen (LN) decreases ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and thus limits photosynthesis. The decrease in Rubisco also delays photosynthetic induction in LN leaves; therefore, we hypothesized that the difference of photosynthetic CO2 fixation between uniform and dynamic light will be greater in LN leaves compared to leaves with sufficient nitrogen supply. Results To test this hypothesis, soybean plants were grown under low or high nitrogen (HN), and the photosynthetic gas exchange, enzyme activity and protein amount in leaves were measured under uniform and dynamic light. Unexpectedly, dynamic light caused less photosynthetic suppression, rather than more, in LN leaves than in HN leaves. The underlying mechanism was also clarified. Short low-light (LL) intervals did not affect Rubisco activity but clearly deactivated fructose-1,6-bisphosphatase (FBPase) and sedoheptulose-1,7-bisphosphatase (SBPase), indicating that photosynthetic induction after a LL interval depends on the reactivation of FBPase and SBPase rather than Rubisco. In LN leaves, the amount of Rubisco decreased more than FBPase and SBPase, so FBPase and SBPase were present in relative excess. A lower fraction of FBPase and SBPase needs to be activated in LN leaves for photosynthesis recovery during the high-light phase of dynamic light. Therefore, photosynthetic recovery is faster in LN leaves than in HN leaves, which relieves the photosynthetic suppression caused by dynamic light in LN leaves. Conclusions Contrary to our expectations, dynamic light caused less photosynthetic suppression, rather than more, in LN leaves than in HN leaves of soybean. This is the first report of a stress condition alleviating the photosynthetic suppression caused by dynamic light.

Oxybenzone (OBZ), avobenzone (AVB), octocrylene (OCR) and octinoxate (OMC) are ultraviolet (UV) filters commonly added to chemical sunscreens. These UV filters are known to widely contaminate the environment through a variety of anthropogenic sources, including sewage discharge. However, systematic studies of the damage caused by these four UV filters and their toxicopathological differences in a variety of plant species are lacking. In this study, we demonstrated that irrigation with water containing these four UV filters could significantly inhibit the aboveground growth of cucumber plant. All of the UV filters decreased photosynthesis through nonstomatal factors but via different inhibitory mechanisms. Only OBZ inhibited photosynthesis by directly inhibiting photosynthetic electron transport, while the other three (AVB, OCR, and OMC) inhibited photosynthesis by inhibiting the Calvin-Benson cycle. Additionally, these four UV filters also decreased plant respiration under long-term treatment. Photosynthesis and respiration inhibition led to the over production of reactive oxygen species (ROS) and the formation of lipid peroxidation damage products, which further damaged the structure and function of plant cells, causing secondary pathologies and potentially leading to reduced crop yields. The study also demonstrated that these four UV filters caused different degrees of phototoxic damage to cucumber plants. On the basis of comprehensive evaluation, we speculated that the order of the four UV filters in terms of plant damage was OBZ > AVB > OMC > OCR. Because of the severe damaging effects of these UV filters on plant growth, the application of contaminated biosolids/reclaimed water in agriculture reduces agricultural production and may damage ecosystems. The results of this study can advance recognition of the hazards associated with environmental and agricultural pollution via UV filters and encourage consumers and the industry to limit or reduce the application of cosmetics and over-the-counter drugs containing these substances.

Bisphenol A (BPA), a widely distributed pollutant, suppresses photosynthesis in leaves. In previous studies on higher plants, the plants were treated by BPA through irrigation to root. This method cannot distinguish whether the BPA directly suppresses photosynthesis in leaves, or indirectly influences photosynthesis through affecting the function of root. Here, only the leaves but not the roots of cucumber were infiltrated with BPA solution. The photosystem II and I (PSII, PSI) were insensitive to BPA under darkness. BPA aggravated the PSII but not the PSI photoinhibition under light. BPA also inhibited CO2 assimilation, and the effect of BPA on PSII photoinhibition disappeared when the CO2 assimilation was blocked. The H2O2 accumulated in BPA-treated leaves under light. And the BPA-caused PSII photoinhibition was prevented under low (2%) O2. We also proved that the BPA-caused PSII photoinhibition depend on the turnover of D1 protein. In conclusion, this study proved that BPA could directly suppress photosynthesis in leaves, however, BPA does not damage PSII directly, but inhibits CO2 assimilation and over-reduces the electron transport chain under light, which increases the production of reactive oxygen species (H2O2), the over-accumulated ROS inhibits the turnover of D1 protein and consequently aggravates PSII photoinhibition.

Wheat embryo globulin nutrient (WEGN), with wheat embryo globulin (WEG) as the main functional component, is a nutritional combination that specifically targets memory impairment. In this study, we explored the protective role of WEGN on Alzheimer's disease (AD)-triggered cognitive impairment, neuronal injury, oxidative stress, and acetylcholine system disorder. Specifically, we established an AD model via administration of d -galactose ( d -gal) and Aluminum chloride (AlCl3) for 70 days, then on the 36th day, administered animals in the donepezil and WEGN (300, 600, and 900 mg/kg) groups with drugs by gavage for 35 days. Learning and memory ability of the treated rats was tested using the Morris water maze (MWM) and novel object recognition (NOR) test, while pathological changes and neuronal death in their hippocampus CA1 were detected via HE staining and Nissl staining. Moreover, we determined antioxidant enzymes by measuring levels of superoxide dismutase (SOD), malondialdehyde (MDA), and glutathione peroxidase (GSH-Px) in serum, cortex, and hippocampus, whereas changes in the acetylcholine system were determined by evaluating choline acetyltransferase (ChAT), and acetylcholinesterase (AChE) activities, as well as choline acetylcholine (Ach) content. Results revealed that rats in the WEGN group exhibited significantly lower escape latency, as well as a significantly higher number of targeted crossings and longer residence times in the target quadrant, relative to those in the model group. Notably, rats in the WEGN group spent more time exploring new objects and exhibited lower damage to their hippocampus neuron, had improved learning and memory activity, as well as reversed histological alterations, relative to those in the model group. Meanwhile, biochemical examinations revealed that rats in the WEGN group had significantly lower MDA levels and AChE activities, but significantly higher GSH, SOD, and ChAT activities, as well as Ach content, relative to those in the model group. Overall, these findings indicate that WEGN exerts protective effects on cognitive impairment, neuronal damage, oxidative stress, and choline function in AD rats treated by d -gal/AlCl3.

Phenanthrene is a polycyclic aromatic hydrocarbon (PAH) that is widely distributed in the environment and seriously affects the growth and development of plants. To clarify the mechanisms of the direct effects of phenanthrene on the plant photosynthetic apparatus, we measured short-term phenanthrene-treated cucumber leaves. Phenanthrene inhibited Rubisco carboxylation activity, decreasing photosynthesis rates (Pn). And phenanthrene inhibited photosystem II (PSII) activity, thereby blocking photosynthetic electron transport. The inhibition of the light and dark reactions decreased the photosynthetic electron transport rate (ETR) and increased the excitation pressure (1−qP). Under high light, the maximum photochemical efficiency of photosystem II (F v /F m ) in phenanthrene-treated cucumber leaves decreased significantly, but photosystem I (PSI) activity (Δ I/I o ) did not. Phenanthrene also caused a J-point rise in the OJIP curve under high light, which indicated that the acceptor side of PSII Q A to Q B electron transfer was restricted. This was primarily due to the net degradation of D1 protein, which is caused by the accumulation of reactive oxygen species (ROS) in phenanthrene-treated cucumber leaves under high light. This study demonstrated that phenanthrene could directly inhibit photosynthetic electron transport and Rubisco carboxylation activity to decrease net Pn. Under high light, phenanthrene caused the accumulation of ROS, resulting in net increases in D1 protein degradation and consequently causing PSII photoinhibition.

To improve light environment, photosynthetic capacity, and thus the yield of maize, the effects of directional planting on light distribution in canopy and photosynthetic characteristics of ear leaves, as well as the performance of PSII that closely related with photosynthetic characteristics and reflected by the rapid chlorophyll fluorescence kinetic curves were examined in Zhengdan 958 maize variety. The results showed that the orientation of leaves remarkably affected photosynthetically active radiation (PAR) interception of ear leaves, with PAR interception of ear leaves in southward treatment being 271.8% higher than that under northward treatment. The orientation of leaves affec-ted photosynthetic light use efficiency of ear leaves under high and low light conditions. The southward treatment increased net photosynthetic rate (P为改善玉米群体内光环境,进一步提高玉米单株光合能力以获得高产,本研究以郑单958为试验材料,通过设置种子定向入土方式,研究了定向有序种植条件下群体内光分布特征,以及单株玉米穗位叶花后光合性能,并借助快速叶绿素荧光动力学曲线分析了与叶片光合性能有密切联系的光系统II(PSII)的性能特征.结果表明: 叶片不同朝向显著改变夏玉米群体内穗位叶处光合有效辐射截获量,朝南处理(S)平均比朝北处理(N)高271.8%.不同朝向的叶片对高光与弱光的利用能力差异显著,朝南处理饱和光强下净光合速率(

To date, it remains unclear how saline stress induces leaf phenolic synthesis in honeysuckle (Lonicera japonica Thunb.), a traditional Chinese medicinal crop. This study was to investigate hormone signaling for salt-induced leaf phenolic synthesis in honeysuckle from molecular, physiological and ecological levels by pot experiments and field trials. As a major ingredient of soil salinity in coastal land, NaCl was used in salt treatment for simulating coastal saline environment. NaCl stress significantly increased leaf abscisic acid (ABA) and salicylic acid (SA) concentrations, potentiated leaf phenolic synthesis by elevating phenylalanine ammonia-lyase (PAL) activity and PALs transcription, and enhanced leaf phenolic accumulation. Under NaCl stress, tungstate and paclobutrazol pretreatments inhibited leaf ABA and SA accumulation, respectively, and lowered PAL activity, PALs transcription, and phenolic accumulation. Thus, ABA and SA participated in transducing salt-induced leaf phenolic synthesis. Paclobutrazol pretreatment hardly influenced leaf ABA concentration in salt-treated plants, but tungstate pretreatment abated salt-induced leaf SA accumulation, indicating that ABA was an upstream signal of SA. Under NaCl stress, SA application restored leaf SA level with no effect on ABA level but incompletely restored PAL activity and phenolic accumulation in tungstate-pretreated plants, confirming the downstream role of SA and implying the existence of other downstream signals. Leaf ABA and SA concentrations were higher in plants in coastal saline plots than non-saline plots, and the roles of ABA and SA in mediating salt-induced leaf phenolic synthesis passed ecological test, according to a significant positive correlation of leaf ABA and SA levels with phenolic concentration. In conclusion, ABA acted as an upstream signal of SA to elicit leaf phenolic synthesis in honeysuckle under coastal saline environment.

The purpose of this study was to explore the effect of reducing nitric oxide (NO) in Rumex K-1 leaves on the photoprotection of photosystem II (PSII) under high temperature with strong light. Reducing the content of NO in Rumex K-1 leaves significantly aggravated the PSII photoinhibition and net degradation of D1 protein under high temperature with strong light, but not under high temperature in the darkness. The reduction of NO remarkably inhibited the electron transport of PSII in the leaves under high temperature and strong light, which resulted in an increase in excitation pressure and an over-accumulation of reactive oxygen species (ROS). The over-accumulation of ROS further damaged PSII. However, when the synthesis of D1 protein was inhibited, the D1 protein content and PSII activity were no longer influenced by reducing NO content in the leaves. The reduction of NO in leaves decreased the activities of ROS scavenger enzymes after treatment with high temperature and strong light for 2 h, which enhanced the over-accumulation of ROS to damage photosynthetic apparatus severely. All of these results suggest that NO was involved in the synthesis of D1 protein. Maintaining physiologically appropriate NO content in leaves will alleviate net degradation of D1 protein under high temperature with strong light to keep photosynthetic electrons flowing smoothly, which mitigates the accumulation of ROS in photosystems to avoid damage to the photosynthetic apparatus. Therefore, NO plays an important role in maintaining higher PSII photosynthetic performance under high temperature with strong light.

Oxybenzone (OBZ), an active ingredient in most sunscreens, was recently shown to be toxic to humans, corals and other animals. This study is the first to demonstrate that OBZ can significantly inhibit photosynthesis and respiration in the leaves of a higher plant, cucumber. An OBZ suspension content as low as 0.228 mg/L obviously inhibited the photosynthesis and respiration of cucumber. OBZ instantly inhibits the electron transport of chloroplasts and mitochondria in cucumber leaves. Probit analysis demonstrated that the effective content for 20% inhibition of photosynthetic electron transport was 11.7 mg/L (95% confidence level). The inhibition of photosynthesis and respiration restricts carbohydrate synthesis and ATP regeneration, respectively, limiting the energy available for metabolic processes including the synthesis of vital organic macromolecules such as proteins and nucleic acids in plant cells. The inhibition of photosynthesis also enhanced the excess excitation energy in chloroplasts, resulting in overproduction of reactive oxygen species (ROS), and the inhibition of respiration aggravated this process. ROS accumulation adversely affects the structure and function of proteins, DNA and membrane lipids in plant cells, interfering with normal metabolism and even leading to plant death. Therefore, reducing the use of OBZ is important for protecting global ecological security.

Oxybenzone (OBZ; benzophenone-3, CAS# 131-57-7) is a known pollutant of aquatic and marine ecosystems, and is an ingredient in over 3000 personal care products, as well as many types of plastics. The aim of this study is to explore the different toxicities of OBZ on an eukaryotic (Chlorella sp.) and a prokaryotic algae (Arthrospira sp.). OBZ is a photo-toxicant, with all observed toxicities more sever in the light than in the dark. Cell growth and chlorophyll inhibition were positively correlated with increasing OBZ concentrations over time. Twenty days treatment with OBZ, as low as 22.8 ng L−1, significantly inhibited the growth and chlorophyll synthesis of both algae. Both algae were noticeably photo-bleached after 7 days of exposure to OBZ concentrations higher than 2.28 mg L−1. Relatively low OBZ concentrations (0.228 mg L−1) statistically constrained photosynthetic and respiratory rates via directly inhibiting photosynthetic electron transport (PET) and respiration electron transport (RET) mechanisms, resulting in over production of reactive oxygen species (ROS). Transmission and scanning electron microscopy showed that the photosynthetic and respiratory membrane structures were damaged by OBZ exposure in both algae. Additionally, PET inhibition suppressed ATP production for CO2 assimilation via the Calvin-Benson cycle, further limiting synthesis of other biomacromolecules. RET restriction limited ATP generation, restricting the energy supply used for various life activities in the cell. These processes further impacted on photosynthesis, respiration and algal growth, representing secondary OBZ-induced algal damages. The data contained herein, as well as other studies, supports the argument that global pelagic and aquatic phytoplankton could be negatively influenced by OBZ pollution.

Little is known regarding the detailed mechanism of CuO NPs’ toxicity to microalgal primary metabolism pathway. Photosynthesis and respiration are the most important primary metabolism and the main sources of production of reactive oxygen species (ROS), but the effect of CuO NPs on both of them has not been systematically studied to date. Our research demonstrated that long-term treatment with CuO NPs significantly inhibited activities of photosynthesis and respiration in microalgae, and the photosynthesis was more sensitive to the toxicity of CuO NPs than respiration. CuO NPs could be absorbed by microalgae and be converted into Cu2O NPs concentrated in chloroplast. The internalized Cu, regardless of whether the exposure was Cu2+ or CuO NPs had the same capacity to damage chloroplast structure. The result also shows that the oxygen-evolving complex (OEC) in the photosynthetic electron transport chain was the most sensitive site to CuO NPs and Cu2+-treated microalgae had the same damage site as that of CuO NPs, which may be related to the Mn cluster that is dissociated by Cu ions released from CuO NPs. The damage of OEC inhibited photosynthetic electron transport to increase excess excited energy, which caused the accumulation of ROS in chloroplast. The accumulation of ROS damaged the structure of cell membrane and aggravated the PSII photoinhibition, further decreasing the efficiency of light energy utilization. In conclusion, the Cu ionic toxicity of photosynthetic apparatus by CuO NPs resulted in the carbon starvation and the accumulation of ROS to inhibit the growth of microalgae.

The microstructures and dielectric properties of Y-Mg-Ga-Si co-doped barium titanate ceramics were investigated. Y3+ dissolved in the lattice of BaTiO3 replacing both Ba2+ site and Ti4+ site, and Mg2+ replaced Ti4+ site. The replacements of Y3+ and Mg2+ inhibit the grain growth, cause tetragonal-to-pseudocubic phase transition, reduce the dielectric loss, and flatten the temperature dependence of capacitance curve. The incorporation of Ga3+ can improve sintering and increase permittivity. Y3+ and Ga3+ tended to remain close to the grain boundaries, and play an important role as a shell maker in the formation of the core–shell structure in the co-doped BaTiO3 ceramics. Excellent dielectric properties: ϵr = ∼2487, tanδ = ∼0.7% (at 1 kHz), ΔC/C25

The effects of LiF addition on phase evolution, microstructure and dielectric properties of BaTiO3 based ceramics were investigated. The LiF addition lowered the sintering temperature of BaTiO3 based ceramics effectively from 1,380 to 1,100 °C. Orthorhombic-to-pseudocubic phase transition appeared as increasing sintering temperature or LiF content. Li+ dissolved in the lattice of BaTiO3, replaced Ti4+ site, produced oxygen vacancies and further inhibited the grain growth. F− entered into O2− site, might cause weakening of oxygen bond strength, together with the low-melting-point liquid phase sintering of LiF, facilitated the formation of uniform dense and fine-grained structure and maintained good dielectric properties. Samples with 2 wt% LiF sintered at 1,100 °C showed excellent dielectric properties: er = 1,956, tanδ = 0.8 % (at 1 kHz), ΔC/C25 4.50 kV/mm.

The rapid induction of photosynthesis is critical for plants under light-fleck environment. Most previous studies about photosynthetic induction focused upon single leaf, but they did not consider the systemic integrity of plant. Here, we verified whether systemic signalling is involved in photosynthetic induction. Rumex K-1 (Rumex patientia × Rumex tianschaious) plants were grown under light-fleck condition. After whole night dark adaptation, different numbers of leaves (system leaf or SL) were pre-illuminated with light, and then the photosynthetic induction of other leaves (target leaf or TL) was investigated. This study showed that the pre-illumination of SL promoted photosynthetic induction in TL. This promotion was independent of the number of SL, the light intensity on SL and the distance between SL and TL, indicating that this systemic signalling is non-dose-dependent. More interestingly, the photosynthetic induction was promoted by only the pre-illumination of morphological upper leaf rather than the pre-illumination of morphological lower leaf, indicating that the transfer of this signal is directional. The results showed that the transfer of this systemic signalling depends upon the phloem. This systemic signalling helps plants to use light energy more efficiently under light flecks.

Although the effect of dehydration on photosynthetic apparatus has been widely studied, the respective effect of dehydration duration and dehydration degree was neglected. This study showed that, when leaves dehydrated in air, the PSII activities of leaves decreased with the decline of leaf relative water content (RWC). Unexpectedly, when leaves dehydrated to same RWC, the decreases in Fv/Fm, Ψo and RC/CSm were lower in leaves dehydrating at 43 °C than those at 25 °C. However, to reach the same RWC, leaves dehydrating at 43 °C experienced 1/6 of the dehydration duration for leaves dehydrating at 25 °C. To distinguish the respective effect of dehydration degree and dehydration duration on photosynthetic apparatus, we studied the PSII activities of leaves treated with different concentration of PEG solutions. Increasing dehydration degree aggravated the decline of Fv/Fm, Ψo and RC/CSm in leaves with the same dehydration duration, while prolonging the dehydration duration also exacerbated the decline of Fv/Fm, Ψo and RC/CSm in leaves with identical dehydration degree. With the same dehydration degree and duration, high temperature enhanced the decrease of Fv/Fm, Ψo and RC/CSm in the leaves. When leaves dehydrated in air, the effect of high temperature was underestimated due to reduction of dehydration duration. The results demonstrated that, dehydration degree and duration both play important roles in damage to photosynthetic apparatus. We suggest that, under combined stresses, the effects of dehydration degree and duration on plants should be considered comprehensively, otherwise, partial or incorrect results may be obtained.

The doping behaviors of yttrium, zinc and gallium and their effects on the dielectric properties and microstructures of BaTiO3 were investigated. Y3+ dissolved in the lattice of BaTiO3, replacing both Ba2+ site and Ti4+ site; while Zn2+ and Ga3+ tended to occupy Ti4+ site. Compared with Y2O3 and Ga2O3, ZnO suppressed grain growth of BaTiO3 more effectively and promoted greater uniformity of grains, thus reducing the dielectric loss. The addition of Ga2O3 inhibited the appearance of second phase Y2Ti2O7 and enhanced the sinterability, which was ascribed to the compensation mechanism and synergistic effect. Proper amount of Y2O3, Ga2O3 and ZnO significantly improved the dielectric temperature characteristics due to the formation of the core–shell structure in the codoped BaTiO3 ceramics. High performance dielectrics with er of 2,690, tanδ of 1.0 % (at 1 kHz), and alternating current breakdown voltage E > 3.73 kV/mm, were achieved.

The mechanism by which the mitochondrial alternative oxidase (AOX) pathway contributes to photosystem II (PSII) photoprotection is in dispute. It was generally thought that the AOX pathway protects photosystems by dissipating excess reducing equivalents exported from chloroplasts through the malate/oxaloacetate (Mal/OAA) shuttle and thus preventing the over-reduction of chloroplasts. In this study, using the aox1a Arabidopsis mutant and nine other C3 and C4 plant species, we revealed an additional action model of the AOX pathway in PSII photoprotection. Although the AOX pathway contributes to PSII photoprotection in C3 leaves treated with high light, this contribution was observed to disappear when photorespiration was suppressed. Disruption or inhibition of the AOX pathway significantly decreased the photorespiration in C3 leaves. Moreover, the AOX pathway did not respond to high light and contributed little to PSII photoprotection in C4 leaves possessing a highly active Mal/OAA shuttle but with little photorespiration. These results demonstrate that the AOX pathway contributes to PSII photoprotection in C3 plants by maintaining photorespiration to detoxify glycolate and via the indirect export of excess reducing equivalents from chloroplasts by the Mal/OAA shuttle. This new action model explains why the AOX pathway does not contribute to PSII photoprotection in C4 plants.

Pseudomonas syringae pv. tabaci (Pst) is a hemibiotrophic bacterial pathogen responsible for tobacco wildfire disease. Although considerable research has been conducted on the tobacco plant’s tolerance to Pst, the role of light in the responses of the photosystems to Pst infection is poorly understood. This study aimed to elucidate the underlying mechanisms of the reduced photosystem damage in tobacco leaves due to Pst infection under light conditions. Compared to dark conditions, Pst infection under light conditions resulted in less chlorophyll degradation and a smaller decline in photosynthetic function. Although the maximal quantum yield of photosystem II (PSII) and the activity of the photosystem I (PSI) complex decreased as Pst infection progressed, damage to PSI and PSII after infection was reduced under light conditions compared to dark conditions. Pst was 17-fold more abundant in tobacco leaves under dark compared to light conditions at 3 days post-inoculation (dpi). Additionally, hydrogen peroxide (H2O2) accumulated to a high level in tobacco leaves after Pst infection under light conditions; although to a lesser extent, H2O2 accumulation was also significant under dark conditions. Pretreatment with H2O2 alleviated chlorotic lesions and decreased Pst abundance in tobacco leaves at 3 dpi under dark conditions. Methyl viologen (MV) pretreatment had the same effects under light conditions, whereas 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU) pretreatment aggravated chlorotic lesions and increased the Pst population. These results indicate that chlorotic symptoms and the size of the bacterial population are each negatively correlated with H2O2 accumulation. In other words, light appears to suppress the Pst population in tobacco leaves through the accumulation of H2O2 during infection.

Background Pseudomonas syringae pv. tabaci (Pst), which is the pathogen responsible for tobacco wildfire disease, has received considerable attention in recent years. The objective of this study was to clarify the responses of photosystem I (PSI) and photosystem II (PSII) to Pst infection in tobacco leaves. Results The net photosynthetic rate (Pn) and carboxylation efficiency (CE) were inhibited by Pst infection. The normalized relative variable fluorescence at the K step (Wk) and the relative variable fluorescence at the J step (VJ) increased while the maximal quantum yield of PSII (Fv/Fm) and the density of QA-reducing PSII reaction centers per cross section (RC/CSm) decreased, indicating that the reaction centers, and the donor and acceptor sides of PSII were all severely damaged after Pst infection. The PSI activity decreased as the infection progressed. Furthermore, we observed a considerable overall degradation of PsbO, D1, PsaA proteins and an over-accumulation of reactive oxygen species (ROS). Conclusions Photoinhibition and photoinhibition-like damage were observed under light and dark conditions, respectively, after Pst infection of tobacco leaves. The damage was greater in the dark. ROS over-accumulation was not the primary cause of the photoinhibition and photoinhibition-like damage. The PsbO, D1 and PsaA proteins appear to be the targets during Pst infection under light and dark conditions. Electronic supplementary material The online version of this article (doi:10.1186/s12870-016-0723-6) contains supplementary material, which is available to authorized users.

Ultraviolet-B radiation (UV-B) is generally considered to negatively impact the photosynthetic apparatus and plant growth. UV-B damages PSII but does not directly influence PSI. However, PSI and PSII successively drive photosynthetic electron transfer, therefore, the interaction between these systems is unavoidable. So we speculated that UV-B could indirectly affect PSI under chilling-light conditions. To test this hypothesis, the cucumber leaves were illuminated by UV-B prior or during the chilling-light treatment and the leaves were then transferred to 25 °C and low-light conditions for recovery. The results showed that UV-B decreased the electron transfer to PSI by inactivating the oxygen-evolving complex (OEC), thereby protecting PSI from chilling-light-induced photoinhibition. This effect advantages the recoveries of PSI and CO2 assimilation after chilling-light stress, therefore should minimize the yield loss caused by chilling-light stress. Because sunlight consists of both UV-B and visible light, we suggest that UV-B-induced OEC inactivation is critical for chilling-light-induced PSI photoinhibition in field. Moreover, additional UV-B irradiation is an effective strategy to relieve PSI photoinhibition and yield loss in protected cultivation during winter. This study also demonstrates that minimizing the photoinhibition of PSI rather than that of PSII is essential for the chilling-light tolerance of the plant photosynthetic apparatus.

The effects of inhibition of mitochondrial alternative oxidase (AOX) respiratory pathway on photosynthetic apparatus in Rumex K-1 leaves were studied. Under high irradiance, the inhibition of AOX pathway caused over-reduction of photosystem (PS) 2 acceptor side, a decrease in the energy transfer in the PS 2 units, damage of donor side of PS 2 and decrease in pool size of electron acceptors. The inhibition of AOX pathway also decreased photosynthetic performance index (PIABS), actual photochemical efficiency (ΦPS2), photochemical quenching (qP) and photosynthetic O2 evolution rate. The results demonstrate that mitochondrial AOX pathway plays a vital role in photoprotection of photosynthetic apparatus.

The purpose of this study was to explore how the mitochondrial AOX (alternative oxidase) pathway alleviates photoinhibition in Rumex K-1 leaves. Inhibition of the AOX pathway decreased the initial activity of NADP-malate dehydrogenase (EC 1.1.1.82, NADP-MDH) and the pool size of photosynthetic end electron acceptors, resulting in an over-reduction of the photosystem I (PSI) acceptor side. The over-reduction of the PSI acceptor side further inhibited electron transport from the photosystem II (PSII) reaction centers to the PSII acceptor side as indicated by an increase in V(J) (the relative variable fluorescence at J-step), causing an imbalance between photosynthetic light absorption and energy utilization per active reaction center (RC) under high light, which led to the over-excitation of the PSII reaction centers. The over-reduction of the PSI acceptor side and the over-excitation of the PSII reaction centers enhanced the accumulation of reactive oxygen species (ROS), which inhibited the repair of the photodamaged PSII. However, the inhibition of the AOX pathway did not change the level of photoinhibition under high light in the presence of the chloroplast D1 protein synthesis inhibitor chloramphenicol, indicating that the inhibition of the AOX pathway did not accelerate the photodamage to PSII directly. All these results suggest that the AOX pathway plays an important role in the protection of plants against photoinhibition by minimizing the inhibition of the repair of the photodamaged PSII through preventing the over-production of ROS.

By simultaneously analyzing the chlorophyll a fluorescence transient and light absorbance at 820 nm as well as chlorophyll fluorescence quenching, we investigated the effects of different photon flux densities (0, 15, 200 μmol m(-2) s(-1)) with or without 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU) on the repair process of cucumber (Cucumis sativus L.) leaves after treatment with low temperature (6°C) combined with moderate photon flux density (200 μmol m(-2 )s(-1)) for 6 h. Both the maximal photochemical efficiency of Photosystem II (PSII) (F (v)/F (m)) and the content of active P700 (ΔI/I (o)) significantly decreased after chilling treatment under 200 μmol m(-2 )s(-1) light. After the leaves were transferred to 25°C, F (v)/F (m) recovered quickly under both 200 and 15 μmol m(-2 )s(-1) light. ΔI/I (o) recovered quickly under 15 μmol m(-2) s(-1) light, but the recovery rate of ΔI/I (o) was slower than that of F (v)/F (m). The cyclic electron transport was inhibited by chilling-light treatment obviously. The recovery of ΔI/I (o) was severely suppressed by 200 μmol m(-2) s(-1) light, whereas a pretreatment with DCMU effectively relieved this suppression. The cyclic electron transport around PSI recovered in a similar way as the active P700 content did, and the recovery of them was both accelerated by pretreatment with DCMU. The results indicate that limiting electron transport from PSII to PSI protected PSI from further photoinhibition, accelerating the recovery of PSI. Under a given photon flux density, faster recovery of PSII compared to PSI was detrimental to the recovery of PSI or even to the whole photosystem.

Although root-to-shoot communication has been intensively investigated in plants under drought, few studies have examined root-to-shoot communication under chilling. Here we explored whether root-to-shoot communication contributes to the chilling-light tolerance of cucumber shoots and clarified the key signal involves in this communication. After leaf discs chilling-light treatment, the photoinhibitions of Photosystem I (PSI) and Photosystem II (PSII) were similar in leaf discs of two cucumber varieties (JY-3 and JC-4). When the whole plants, including roots, were chilled under light, the photosynthetic performances in JC-4 leaves decreased more seriously than that in JY-3 leaves. However, when the water status of leaves was maintained by warming roots or floating the attached leaves on water, the PSII activity and amount of PSI in the leaves of the two varieties were similar after chilling-light treatment. In addition, the differences of PSII activities and amount of PSI between the two varieties under whole plant chilling-light treatment were independent of ABA pretreatment. Above results indicate that (1) the better water status in leaves under chilling contributes to the higher chilling tolerance of JY-3; (2) the water status, rather than an ABA signal, dominates root-to-shoot communication under chilling and the chilling tolerance of cucumber shoot.

The rapid induction of photosynthesis is critical for plants under light-fleck environment. Most previous studies about photosynthetic induction focused upon single leaf, but they did not consider the systemic integrity of plant. Here, we verified whether systemic signalling is involved in photosynthetic induction. Rumex K-1 (Rumex patientia × Rumex tianschaious) plants were grown under light-fleck condition. After whole night dark adaptation, different numbers of leaves (system leaf or SL) were pre-illuminated with light, and then the photosynthetic induction of other leaves (target leaf or TL) was investigated. This study showed that the pre-illumination of SL promoted photosynthetic induction in TL. This promotion was independent of the number of SL, the light intensity on SL and the distance between SL and TL, indicating that this systemic signalling is non-dose-dependent. More interestingly, the photosynthetic induction was promoted by only the pre-illumination of morphological upper leaf rather than the pre-illumination of morphological lower leaf, indicating that the transfer of this signal is directional. The results showed that the transfer of this systemic signalling depends upon the phloem. This systemic signalling helps plants to use light energy more efficiently under light flecks.

The effects of chilling-light stress combined with additional stress on PSI and PSII photoinhibition and their interrelationship have not been known. To explore whether NaCl affects the PSI and PSII photoinhibition and their interrelationship under chilling-light treatment, the PSI and PSII activities were studied under chilling-light with or without NaCl treatment. The results showed that the extent of PSI and PSII photoinhibition both increased under chilling-light, while NaCl aggravated PSII photoinhibition and severely damaged cytochrome b6/f complex but alleviated PSI photoinhibition. Moreover, DCMU had a similar effect as NaCl in this study, which indicates that NaCl alleviated PSI photoinhibition through reducing electrons transported to PSI. It was also showed that the increased damage to PSII by NaCl did not depend on the inhibition of PSII repair and PSI electron transportation. In conclusion, NaCl alleviated PSI photoinhibition by inhibiting electron transport from PSII under chilling-light conditions. In addition, PSII photoinhibition was not affected by PSI photoinhibition because of a full inhibition of PSII repair by chilling-light treatment. We also speculate that NaCl aggravates PSII photoinhibition by enhancing the damage instead of inhibiting the repair of it under chilling-light conditions.

Background Sclerotinia sclerotiorum (Lib.) de Bary is a necrotrophic fungal pathogen which causes disease in a wide range of plants. An observed decrease in photosynthetic performance is the primary reason for the reduction of crop yield induced by S. sclerotiorum. The H2C2O4 is the main pathogenic material secreted by S. sclerotiorum, but the effects of H2C2O4 acidity and the C2O42− ion on photosynthetic performance remain unknown. Results S. sclerotiorum infection significantly decreased photosynthetic O2 evolution and the maximum quantum yield of photosystem II (Fv/Fm) in tobacco leaves under high-light. H2C2O4 (the main pathogenic material secreted by S. sclerotiorum) with pH 4.0 also significantly decreased photosynthetic performance. However, treatment with H3PO4 and HCl at the same pH as H2C2O4 caused much less decrease in photosynthetic performance than H2C2O4 did. These results verify that the acidity of the H2C2O4 secreted by S. sclerotiorum was only partially responsible for the observed decreases in photosynthesis. Treatment with 40 mM K2C2O4 decreased Fv/Fm by about 70% of the levels observed under 40 mM H2C2O4, which further demonstrates that C2O42− was the primary factor that impaired photosynthetic performance during S. sclerotiorum infection. K2C2O4 treatment did not further decrease photosynthetic performance when D1 protein synthesis was fully inhibited, indicating that C2O42− inhibited PSII by repressing D1 protein synthesis. It was observed that K2C2O4 treatment inhibited the rate of RuBP regeneration and carboxylation efficiency. In the presence of a carbon assimilation inhibitor, K2C2O42 treatment did not further decrease photosynthetic performance, which infers that C2O42− inhibited PSII activity partly by repressing the carbon assimilation. In addition, it was showed that C2O42− treatment inhibited the PSII activity but not the PSI activity. Conclusions This study demonstrated that the damage to the photosynthetic apparatus induced by S. sclerotiorum is not only caused by the acidity of H2C2O4, but also by C2O42− which plays a much more important role in damaging the photosynthetic apparatus. C2O42− inhibits PSII activity, as well as the rate of RuBP regeneration and carboxylation efficiency, leading to the over production of reactive oxygen species (ROS). By inhibiting the synthesis of D1, ROS may further accelerate PSII photoinhibition. Electronic supplementary material The online version of this article (doi:10.1186/s12870-014-0240-4) contains supplementary material, which is available to authorized users.

Taking Rumex K-1 leaves as test materials, this paper studied the role of mitochondrial alternative oxidase (AOX) pathway in photoprotection under different light intensities. Under low light intensity (200 micromol x m(-2) x s(-1)), and after treated with salicylhydroxamic acid to inhibit the AOX pathway, the leaf actual photochemical efficiency of PS II, linear electron transport rate of photosynthesis, and photosynthetic O2 evolution rate all decreased significantly while the non-Q(B) reducing reaction center had a significant increase, indicating that under low light, the photoinhibition was aggravated while the scavenging enzymes of reactive oxygen species (ROS) increased, which avoided the over-accumulation of ROS and partially alleviated the photoinhibition of Rumex K-1 leaves. Under high light intensity (800 micromol x m(-2) x s(-1)), the inhibition of AOX pathway caused more severe photoinhibition, and the increased activities of ROS scavenging enzymes were insufficient to prevent the over-accumulation of ROS. This study demonstrated that AOX pathway played an important role in the photoprotection in Rumex K-1 leaves under both high and low light intensities, and the role of AOX pathway in photoprotection under high light could be irreplaceable by the other photoprotection pathways in chloroplast.

The net photosynthetic rate, chlorophyll content, chlorophyll fluorescence and 820 nm transmission were investigated to explore the behavior of the photosynthetic apparatus, including light absorption, energy transformation and the photoactivities of photosystem II (PSII) and photosystem I (PSI) during senescence in the stay-green inbred line of maize (Zea mays) Q319 and the quick-leaf-senescence inbred line of maize HZ4. The relationship between the photosynthetic performance and the decrease in chlorophyll content in the two inbred lines was also studied. Both the field and laboratory data indicated that the chlorophyll content, net photosynthetic rate, and the photoactivities of PSII and PSI decreased later and slower in Q319 than in HZ4, indicating that Q319 is a functional stay-green inbred line. In order to avoid the influence of different development stages and environmental factors on senescence, age-matched detached leaf segments from the two inbred lines were treated with ethephon under controlled conditions to induce senescence. The net photosynthetic rate, light absorption, energy transformation, the activities of PSII acceptor side and donor side and the PSI activities decreased much slower in Q319 than in HZ4 during the ethephon-induced senescence. These results suggest that the retention of light absorption, energy transformation and activity of electron transfer contribute to the extended duration of active photosynthesis in Q319. Although the chlorophyll content decreased faster in HZ4, with decrease of chlorophyll content induced by ethephon, photosynthetic performance of Q319 deteriorated much more severely than that of HZ4, indicating that, compared with Q319, HZ4 has an advantage at maintaining higher photosynthetic activity with decrease of chlorophyll although HZ4 is a quick-leaf-senescence inbred line. We conclude that attention should be paid to two favorable characteristics in breeding long duration of active photosynthesis hybrids: 1) maintaining more chlorophyll content during senescence and 2) maintaining higher photosynthetic activity during the loss of chlorophyll.

Taking Cucumis sativus L. (Jinchun No. 4) as test material, through the determination of chlorophyll-a fluorescence transient and light absorbance at 820 nm, and in combining with chlorophyll quenching, this paper studied the recovery of cucumber leaf' s PS I and PS I activities and the interactions between PS I and PS II in the recovery process at room temperature (25 degrees C) and under different light intensities (0, 15, and 200 micromol x m(-2) x s(-1)) after six hours of low temperature (4 degrees C) and strong light (200 micromol x m(-2) x s(-1)) stress. Different extent of photoinhibition of the PS II and PS I occurred after the stress. During the recovery process at room temperature, the PS II activity recovered quickly and was insensitive to light intensity, while the PS I activity recovered quickly under weak light intensity (15 micromol x m(-2) x s(-1)) but slowly under strong light intensity (200 micromol x m(-2) x s(-1)), suggesting that after the chilling-induced photoinhibition, the reduced electron transfer from PS II to PS I protected the PS I from further inhibition, accelerating the recovery of PS I activity. In the breeding of chilling-resistant species of cucumber, it should not only pursue the higher chilling-resistance of PS II and faster recovery of PS II after chilling-induced photoinhibition, but also pay more attention to the coordinating of PS I and PS II during and after the chilling-induced photoinhibition. In the culture of cucumber, after chilling happened, a practical method to reduce light intensity would help the recovery of cucumber leaf PS I activity to protect the photosynthetic apparatus against photoinhibition.

Alternaria alternata has received considerable attention in current literature and most of the studies are focused on its pathogenic effects on plant chloroplasts, but little is known about the characteristics of programmed cell death (PCD) induced by metabolic products (MP) of A. alternata, the effects of the MP on mitochondrial respiration and its relation to PCD. The purpose of this study was to explore the mechanism of MP-induced PCD in non-green tobacco BY-2 cells and to explore the role of mitochondrial inhibitory processes in the PCD of tobacco BY-2 cells. MP treatment led to significant cell death that was proven to be PCD by the concurrent cytoplasm shrinkage, chromatin condensation and DNA laddering observed in the cells. Moreover, MP treatment resulted in the overproduction of reactive oxygen species (ROS), rapid ATP depletion and a respiratory decline in the tobacco BY-2 cells. It was concluded that the direct inhibition of the mitochondrial electron transport chain (ETC), alternative pathway (AOX) capacity and catalase (CAT) activity by the MP might be the main contributors to the MP-induced ROS burst observed in tobacco BY-2 cells. The addition of adenosine together with the MP significantly inhibited ATP depletion without preventing PCD; however, when the cells were treated with the MP plus CAT, ROS overproduction was blocked and PCD did not occur. The data presented here demonstrate that the ROS burst played an important role in MP-induced PCD in the tobacco BY-2 cells.

Background It is known that excess reducing equivalents in the form of NADPH in chloroplasts can be transported via shuttle machineries, such as the malate-oxaloacetate (OAA) shuttle, into the mitochondria, where they are efficiently oxidised by the mitochondrial alternative oxidase (AOX) respiratory pathway. Therefore, it has been speculated that the AOX pathway may protect plants from photoinhibition, but the mechanism by which this protection occurs remains to be elucidated. Results The observation that the malate-OAA shuttle activity and the AOX pathway capacity increased markedly after intense light treatment in Rumex K-1 leaves indicates that excess NADPH was transported from the chloroplasts and oxidised by the AOX pathway. The inhibition of the AOX pathway by salicylhydroxamic acid (SHAM) caused the over-reduction of the photosystem I (PSI) acceptor side, as indicated by the increases in the extent of reduction of P700+. Furthermore, the photosynthetic linear electron flow was restricted, which was indicated by the decreases in the PSII electron transport rate (ETR) and the photosynthetic O2 evolution rate. The restriction of the photosynthetic linear electron flow, which generates the thylakoid ΔpH, inevitably decreased the de-epoxidation of the xanthophyll cycle (ΔPRI). Therefore, the induction of non-photochemical quenching (NPQ) was suppressed when the AOX pathway was inhibited. The effect of the inhibition of the AOX pathway on NPQ induction was less at 20 mM NaHCO3 than at 1 mM NaHCO3. The suppression of NPQ induction by the inhibition of the AOX pathway was also observed during the induction phase of photosynthesis. In addition, the inhibition of the AOX pathway increased the accumulation of hydrogen peroxide (H2O2), suggesting that the AOX pathway functions as an antioxidant mechanism. Conclusions The inhibition of the AOX pathway resulted in the rapid accumulation of NADPH in the chloroplasts, which caused the over-reduction of the PSI acceptor side. Furthermore, the restriction of the photosynthetic linear electron flow due to the inhibition of the AOX pathway limited the generation of the thylakoid ΔpH and suppressed the induction of NPQ. Therefore, the mitochondrial AOX pathway protected the photosynthetic apparatus against photodamage by alleviating the over-reduction of the PSI acceptor side and accelerating the induction of NPQ in Rumex K-1 leaves.